SMITHSONIAN

NATURAL
THE ULTIMATE VISUAL GUIDE TO EVERYTHING ON EARTH

history
FOREWORD 6
ABOUT THIS BOOK 8

LIVING EARTH MINERALS, MICROSCOPIC PLANTS

A living planet 12
ROCKS, AND LIFE MOSSES 108
Active Earth 14 FOSSILS ARCHAEA AND BACTERIA 90 LIVERWORTS 110
Changing climates 16
Habitats for life 18 MINERALS 38 PROTISTS 94 HORNWORTS 111
Human impact 20 ROCKS 62 Amoebas and relatives 96
LYCOPHYTES 111
Origins of life 22 Flagellates 97
FOSSILS 74 Rhizarians 98 FERNS AND RELATIVES 112
Evolution and diversity 24
Evolution in progress 26 Alveolates 100
CYCADS, GINKGOS,
Classification 28 Heterokonts 101
AND GNETOPHYTES 116
Animal genealogy 30 Red algae 103
Tree of life 32 Green algae 105 CONIFERS 118
Stoneworts and relatives 105 FLOWERING PLANTS 122
Basal angiosperms 124
Magnoliids 128
Monocots 130
Eudicots 150

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 FISHES 324 Turacos 464
Jawless fishes 326 Hoatzin 465
Cartilaginous fishes 327 Cuckoos 465
Ray-finned fishes 334 Owls 467
Lobe-finned fishes 353 Mesites 470
Nightjars 471
AMPHIBIANS 354 Hummingbirds and swifts 473
Frogs and toads 356 Trogons 476
Caecilians 369 Mousebirds 476
Salamanders and newts 370 Kingfishers and relatives 477
REPTILES 374
Cuckoo-roller 479
Turtles and tortoises 376
Hornbills, hoopoes, and
Tuatara 383
wood hoopoes 482
Lizards 384
Woodpeckers and toucans 483
Amphisbaenians 393
Seriemas 487
Snakes 394
Passerines 488

FUNGI ANIMALS Crocodiles and alligators 404 MAMMALS 506

BIRDS 408
Egg-laying mammals 508
MUSHROOMS 212 INVERTEBRATES 250
Tinamous 410
Pouched mammals 509
Sponges 252 Sengis 518
SAC FUNGI 238 Cnidarians 254 Ratites 410
Tenrecs and golden moles 519
Flatworms 258 Fowl, game birds, and relatives 412
Aardvark 520
LICHENS 244 Waterfowl 416
Roundworms 259 Dugong and manatees 521
Segmented worms 260 Penguins 420
Hyraxes 521
Velvet worms 260 Loons 424
Elephants 522
Water bears 261 Albatrosses, petrels, Armadillos 525
Arthropods 262 and shearwaters 425
Sloths and anteaters 528
Arachnids 264 Grebes 427
Rabbits, hares, and pikas 529
Sea spiders 270 Flamingos 428
Rodents 533
Horseshoe crabs 270 Storks 429
Tree shrews 543
Crustaceans 271 Ibises, bitterns, herons, Colugos 543
Insects 278 and pelicans 430
Primates 544
Ribbon worms 304 Cormorants, gannets, Bats 562
Bryozoans 304 and relatives 433
Hedgehogs, moles, and relatives 570
Lampshells 305 Tropicbirds 433
Pangolins 573
Mollusks 305 Birds of prey 434
Carnivores 574
Bivalves 306 Falcons and caracaras 441
Odd-toed ungulates 602
Gastropods 308 Bustards 442
Even-toed ungulates 608
Cephalopods 313 Cranes, rails, and relatives 443
Whales, porpoises, and dolphins 628
Chitons 317 Kagu and sunbittern 445
Tusk shells 317 Waders, gulls, and auks 448
Echinoderms 318 Sandgrouse 456 GLOSSARY 634
Pigeons and doves 457 INDEX 638
CHORDATES 322 Parrots and cockatoos 460 ACKNOWLEDGMENTS 663

Established in 1846, the Smithsonian Institution—the ADDITIONAL CONSULTANTS CONSULTANT EDITOR

world’s largest museum and research complex—includes Dr. Matthew D. Kane, Dr. James D. Lawrey,
19 museums and galleries and the National Zoological David Burnie is a former winner of the Aventis Prize for
Dr. Diana Lipscomb, Dr. Robert Lücking, Dr. Thorsten Science Books, and the editor of DK’s highly successful
Park. The total number of objects, works of art, and
specimens in the Smithsonian’s collections is estimated at Lumbsch, Andrew M. Minnis, Dr. Ashleigh Smythe, Animal. He has written or contributed to more than 100
155.5 million, the bulk of which is contained in the Dr. William B. Whitman. books and is a fellow of the Zoological Society of London.
National Museum of Natural History, which holds more
than 126 million specimens and objects. The Smithsonian
is a renowned research center, dedicated to public
CONTRIBUTORS
education, national service, and scholarship in the Richard Beatty, Dr. Amy-Jane Beer, Dr. Charles Deeming,
arts, sciences, and history. SMITHSONIAN ENTERPRISES Dr. Kim Dennis-Bryan, Dr. Frances Dipper, Dr. Chris
Product develoPment m anager Gibson, Derek Harvey, Professor Tim Halliday, Rob Hume,
SMITHSONIAN CONSULTANTS Kealy Gordon Geoffrey Kibby, Dr. Richard Kirby, Joel Levy, Chris
Mattison, Felicity Maxwell, Dr. George C. McGavin,
Dr. Stephen Cairns, Dr. Allen Collins,
director, licensed Publishing Dr. Pat Morris, Dr. Douglas Palmer, Dr. Katie Parsons,
Dana M. De Roche, Dr. Carla Dove, Leslie Hale,
Jill Corcoran Chris Pellant, Helen Pellant, Michael Scott, Carol Usher,
Dr. M. G. (Jerry) Harasewych, Gary Hevel, Dr. Rafael Professor Mark Viney, Dr. David J. Ward,
Lemaitre, Dr. Chris Meyer, Dr. Jon Norenburg, Dr. Elizabeth Wood.
DMM, ECom and D-to-C
Dr. David L. Pawson, Paul Pohwat, Dr. Jeffrey E. Post, Janet Archer
Dr. Klaus Rutzler, Dr. Hans-Dieter Sues, Dr. Michael
Vecchione, Dr. Warren Wagner, Dr. Jeffrey T. Williams, President, smithsonian enterPrises
Dr. Don E. Wilson, Dr. George Zug. Carol LeBlanc
F O R E WO R D
When I was a boy, I would visit the local public library and spend
as long as I was allowed looking at science and reference books.
The books that especially enthralled me were the forerunners of
this book. Packed with color diagrams, images of exotic species,
and faraway places, coupled with informative text, they were the
siren call for a lifetime of studying and teaching. The natural world
around me was then an unknown entity and I wanted to learn
about all of it. It is a very human desire to learn, but nobody
knows which bit of biology they will be most attracted to and I
went on to pick the largest and most diverse animal group of all.
If I were to live a lifetime of lifetimes, I would never know
everything there is to know about insects.
No book could ever be a comprehensive study of Earth’s extant
life forms, which incidentally comprise only one percent of those
that have ever lived. For that you would need a library of hefty
tomes. What you have in your hands is much more serviceable. It
is a clear roadmap to guide you on a journey that will take you
from the steaming forests of the tropics to the freezing cold of
the polar regions, and from mountain peaks to the depths of the
ocean. It is an abridged exploration of the products of a couple of
billion years of evolution that will show you some of the several
million ways of living on Earth.
It is relatively easy to write screeds of detailed information on
any topic you care to name, but this approach is not going to be
of use to anyone other than experts and it is unlikely to encourage
anyone else to study the living world. No, the real skill and the
very purpose of this book is to distil a vast body of natural history
knowledge amassed by countless researchers to provide an
authoritative, yet accessible, account.
There may come a time in the distant future when we are
able to venture elsewhere in our galaxy and beyond. Will we find
extraterrestrial life on a far distant rocky planet and if so, will it
be simple, single-celled organisms or complex creatures to rival
humans or humpbacked whales? To imagine that in the vastness
of the Cosmos, our inconsequential planet is the single, solitary
place where life has evolved—an unbelievable fluke of stellar
serendipity—seems vanishingly improbable. What is certain is
that we inhabit a planet of staggering complexity and breathtaking
beauty, and one that is constantly changing. What we must realize,
and quickly, is that we have no dominion over it but are simply
part of an immense community of interdependent species.
Newcomers to natural history will find themselves engrossed
and even specialists, steeped in the minutiae of one tiny part of
the tree of life, will value the majestically broad sweep of this
meticulously researched volume. If I could have my life all over
again, I’d start by reading this.

DR. GEORGE McGAVIN

Honorary Research Associate, 
Oxford University Museum of Natural History
Senior Principal Research Fellow, Imperial College London
ABOUT THIS BOOK
The Natural History Book begins with a general introduction to life on
Earth: the geological foundations of life, the evolution of life forms, and how for easy reference, visual
organisms are classified. The next five chapters form an extensive, accessible contents panels list the
subgroups within each section,
catalog of specimens—from minerals to mammals—interspersed with and the page number where
each subgroup can be found
fact-filled introductions to each group and in-depth feature profiles.

SECTION INTRODUCTION
Each chapter is divided into sections 374 REPTILES
r e p t i l e g ro u p s
By the Mesozoic Era, reptiles had

representing major taxonomic groupings.

diversified. Turtles and crocodilians
had diverged early on, and later the
Reptiles are a sophisticated, diverse, and successful group of squamates—lizards, snakes, and

ANIMALS • REPTILES
amphisbaenians—appeared.
ectothermic (cold-blooded) vertebrates. Although commonly
The section introduction highlights associated with hot, dry environments, they are found in a
wide range of habitats and climates around the world.
t u rt l e s a n d
t o rt o i s e s

the characteristics and behaviors that PHYLUM

CLASS
CHORDATA
REPTILIA
The first reptiles evolved from amphibians more
than 295 million years ago. They are the ancestors
not only of modern-day reptiles but also of mammals
376
t u ata r a

define the group, and discusses their

ORDERS 4
FAMILIES 92
and birds. During the Mesozoic Era, reptiles such
SPECIES 11,050
as dinosaurs, aquatic ichthyosaurs and plesiosaurs, 383

evolution over time.

and aerial pterosaurs dominated Earth. Reptile lizards
groups that still exist today evolved in this period
and survived the mass extinction event that killed
off the dinosaurs 66 million years ago. 384
All reptiles share certain characteristics, such as scaly a m ph i s ba e n i a n s
skin and behavioral thermoregulation—using outside
sources to maintain a constant body temperature, by
basking in the sun, for example. But the various groups 393
exhibit obvious differences. Turtles and tortoises are s na k e s
Reptile scales are a protective layer of protected by a unique, heavily armored shell. Lizards,
PHYLUM CHORDATA skin with overlapping patches thickened
with keratin and sometimes bone. crocodilians, and tuataras have four limbs and a long tail.
394

on each introduction, CLASS REPTILIA

Many squamates, or scaled reptiles—lizards, snakes,
and amphisbaenians—have evolved limbless forms.
Hot desert ecosystems are typically dominated by
c ro c o d i l e s a n d
a l l i g at o r s

classification boxes display lizards and snakes but reptiles of all types are found in
all habitats in the tropics and subtropics. Fewer species
404

the current taxonomic ORDERS 4 are found in cooler temperate climates. Within each
ecosystem reptiles are important both as predators
and as prey. Most are carnivores, feeding on a wide

hierarchy—the level of the FAMILIES 92

The shelled egg allows reptiles to
reproduce out of water. The outer layer
protects the embryo from dehydration.
range of other animals. A few large lizards and
tortoises are exclusively herbivores, but many
species are opportunistic omnivores.

group under discussion SPECIES 11,050 b e h av i o r a n d s u rv i va l

is highlighted
Some species are solitary whereas others are highly
social. Although sluggish and inactive when cool, once
their preferred body temperature is reached by basking
or pressing against a warm rock, they can be very active.
Reproductive behavior can be complex, with males
Crocodiles can walk with their body actively defending territories and courting females.
raised well off the ground, whereas Although some squamates give birth to live young, in
turtles manage only a sprawling gait. most cases the male reptile fertilizes the female, who
then lays eggs in an underground nest. All reptiles are
independent and self-feeding from birth, although some
crocodilians give parental care for two or more years.
Reptiles are exploited by humans for their skin or
as food. This, along with habitat loss, pollution, and
climate change, threatens the survival of many species.

Reptiles modify their behavior to SEA TURTLE IN ITS ELEMENT

regulate body temperature. They bask in The green turtle may appear primitive but, with efficient

debate boxes tackle scientific the morning sun to absorb heat energy. flippers and a flattened shell, is well adapted to its aquatic life.

controversies and taxonomic discussions

arising from new discoveries
a male of a female of
the species the species
384 LIZARDS C H I S E L-T E E T H L I Z A R D S
Common in Africa, southern Asia, and Australasia, the small lizards
385
of the family Agamidae—which also includes flying lizards and water
Typically, lizards have four legs and a mechanisms depend on agility, camouflage, PHYLUM CHORDATA dragons—are the Old World equivalents of the iguanas. Spines, flaps, AUSTRALIAN WATER DRAGON
Intellagama lesueurii
3¼ ft
1m
long, thin tail, but there are also many and bluff. Many species are able to shed their
REPTILES • LIZARDS

CLASS REPTILIA and crests often adorn the head and back of these lizards, which lay The largest water dragon in Australia, this
legless species. They have scaly skin tail to distract the attention of predators as ORDER SQUAMATA soft-shelled eggs. Males are brightly colored while the females are duller. species lives beside water into which it
they escape. The tail grows back. In some FAMILIES (LIZARDS) 38 dives to escape predators. Adults eat
and firm jaw articulation. invertebrates and small vertebrates.
families of lizards, the skin can change color. SPECIES 6,687
16 in
All lizards are ectothermic, obtaining heat This ability can be used for camouflage or for 40 cm
energy from the environment. Although seen sexual or social signaling. Unusually, in many D E B AT E
as predominately tropical or desert animals, they lizards the pineal gland on the top of the head S Q U A M ATA : A N
have a global distribution; lizards are found from acts as a light-sensitive “third eye.” ALL-INCLUSIVE ORDER ORIENTAL 3¼ ft long limbs
beyond the Arctic Circle in Europe to the tip of GARDEN LIZARD 1m
South America. Highly adaptable, they occupy va r i e d l i f e s t y l e s Traditionally, lizards and snakes were
considered two distinct groups, but
Calotes versicolor
This agile, diurnal lizard is often
ASIAN WATER
DRAGON PA R ROT S A N D C O C K ATO O S black mask
a wide range of terrestrial habitats with many Although some species are solitary, many lizards
being arboreal or rock-dwelling. Many legless have complex social structures, with the males
modern genetic research makes it clear
that they are not. Primitive squamates
found hunting insects in trees
around human habitation. It
is common in S. Asia.
Physignathus cocincinus
A powerful swimmer, this lizard seeks
refuge in water when threatened. It is
462
species are adapted for burrowing, a few lizards maintaining their territories through visual (scaled reptiles) were lizardlike animals
long, laterally
flattened tail used found in riverside trees in S. Asia.
are effective gliders from trees, and others are signals. Many species lay eggs in underground that arose in the mid-Jurassic. Available in swimming
semiaquatic, including a marine species on nests, while others retain the eggs in the oviduct evidence suggests that snakes evolved long tail helps
with balance
16 in NORTH AFRICAN MASTIGURA
Uromastyx acanthinura
40 cm
the Galapagos Islands. until hatching. There are also truly viviparous from lizards during the mid-Cretaceous. when climbing This herbivorous lizard inhabits 14 in 8 in 18½ in 12 in
BIR DS • PAR ROTS A ND COCK ATOOS

species, in which the mother provides nutrition By this time various families of lizards the harsh desert environments of 36 cm 20 cm 47 cm 30 cm
had branched off on their own BOULENGER’S N. Africa. It uses its clublike,
s t r at e g i e s f o r s u rv i va l via a placenta. Males have paired hemipenes— PRICKLENAPE spiny tail in defense.
Lizards range in length from about 1/2 in (1.5 cm) to sexual organs used for internal fertilization— evolutionary course. Some lizard 10 in Acanthosaura crucigera
families are thus closer to snakes 26 cm This slow-moving lizard from Asia
10 ft (3 m) in the case of the Komodo dragon, but although some species are parthenogenetic, with than they are to other lizards. Of living preys on insects while perched on a
the majority are between 4 and 12 in (10–30 cm). females reproducing without the participation lizards, the monitors are thought to be
branch. Males use the long spines on
Although most are carnivores, about two percent of a male. Some lizards tend their eggs during those most closely related to snakes.
their neck when fighting each other.
yellow belly
of species are primarily herbivorous. Many lizards incubation but very few exhibit any maternal chisel-shaped teeth on
outer rim of mouth
are prey for other carnivores. Their defense care of the hatchlings.
CENTRAL BEARDED DRAGON 20 in
Pogona vitticeps 50 cm
Found in arid woodland in Australia, this
CHAMELEONS lizard has a beard of spines. It lays eggs
in an underground nest.
3¼ in
Restricted to the Old World, the members of the 8 cm
TURQUOISE PARROT EASTERN ROSELLA
family Chamaeleonidae have long limbs with grasping
feet and a prehensile tail for gripping branches—an Neophema pulchella Platycercus eximius
adaptation for life in trees. Their eyes are capable of
8 in
20 cm f: Psittacidae f: Psittacidae
moving independently in any direction to locate GREEN-STRIPED TREE DRAGON
insects or small vertebrates. They catch their prey BEARDED PYGMY-CHAMELEON PARSON’S CHAMELEON
Diploderma splendidum A member of a group of small, Rosellas are broad-tailed parrots
28 in
with a long, sticky tongue that shoots out from the Rieppeleon brevicaudatus Calumma parsonii 70 cm This colorful lizard is from humid
montane forests of China, where it
green grass parrots from with a range from Australia to
mouth. Their ability to change color is for display This unusual, small chameleon is from
E. Africa. Drab coloration and patterning
The largest chameleon in the world, this
species is restricted to Madagascar. It hunts hunts insects. It lays small clutches Australia, the turquoise parrot nearby Pacific islands. A typical
and camouflage. Many species are threatened of five to seven eggs.
with extinction.
along its body allow it to resemble a dead leaf. invertebrates in the canopy of montane forests.
12 in is an open woodland bird from rosella, this variable E.
prehensile tail
30 cm
the southeast of the continent. Australian bird has white cheeks.
green skin coloration 5–7 in
sometimes changes to brown 15–18 cm
RED-FAN PARROT ROSE-RINGED MASKED SHINING PARROT
12 in
30 cm Deroptyus accipitrinus PARAKEET Prosopeia personata blue plumage
MEDITERRANEAN
CHAMELEON THORNY DEVIL f: Psittacidae Psittacula krameri f: Psittacidae and upperparts
Chamaeleo chamaeleon
This chameleon spends its
JACKSON’S
CHAMELEON
Moloch horridus
This Australian lizard feeds exclusively on ants. The Possibly related more closely f: Psittacidae One of the three species of
time in bushes searching for Triocerus jacksonii large skin frill body is covered with large, thorny spines, which make to macaws than to other This is the most widespread Asian shining parrots restricted to Fiji,
insects. It is found in N. This diurnal, arboreal species is found in it difficult for a predator to swallow. The spines also
dorsal
spines
Africa and around the E. Africa. The male is identifiable by the three
makes head
look larger channel rainwater or morning dew into its mouth. short-tailed parrots, this parakeet that reaches westward this parrot is declining in numbers
Mediterranean. horns on its snout, which are for display.
S. American parrot raises a into N. Africa. This species has also due to loss of its native forest.
grasping, red neck ruff when excited. been introduced in Europe.
231/2 in
four-toed feet 60 cm 35 in
90 cm
20 in 22 in 16 in
51 cm 56 cm PANTHER FRILLED LIZARD 40 cm 15–16½ in 16–18½ in
CHAMELEON Chlamydosaurus kingii
Furcifer pardalis Common in subtropical woodland 38–42 cm 40–47 cm
GIANT SPINY CHAMELEON
Furcifer verrucosus
Found only in dry forests VEILED CHAMELEON
Chamaeleo calyptratus
strong hind legs
allow this lizard to
in Australia, this species feeds on RAINBOW LIZARD
Agama agama
gray-black
in Madagascar, this lizard insects and other lizards in trees and
This large chameleon is a native of the humid, hunts insects in trees by This species is from the southern coast of the run away bipedally on the ground. When threatened, it This common, insectivorous lizard from Africa is gray at undertail
coastal region of Madagascar. A shy species, it stealth. Males are Arabian Peninsula. Males have a large casque unfurls its frill and opens night, but becomes brightly colored in sunlight. Males
relies on camouflage to ambush its insect prey. highly territorial. on the head, while females have a smaller one. its mouth wide. use their bright skin colors during territorial disputes.

16 in
white face 40 cm

GROUP INTRODUCTION 13 in
33 cm
SUPERB PARROT
Within each section—for example, species-specific
Polytelis swainsonii
f: Psittacidae
reptiles—lower-ranking taxonomic groups, information
PRINCESS PARROT
Polytelis alexandrae
f: Psittacidae
This S.E. Australian bird
belongs to a group of long-tailed
Australian parrots and breeds in

such as lizards, are explored. Key features accompanies A nomad from central
Australia, this bird follows
watercourses and gathers
red gum forests. It is rapidly
declining in number.

are described, including their distribution, each image near spinifex grass. It often
breeds in small colonies yellow
undertail
habitat, physical characteristics, life cycle, in eucalyptus trees.

behavior, and reproductive habits. pale gray

underparts

6 in 6½–7 in
15 cm 17–18 cm

SPECIES CATALOG
Pictorial galleries profile some 5,000
species, showing the distinctive visual GRAY PARROT
YELLOW-COLLARED
LOVEBIRD
Agapornis personatus
ROSY-FACED
LOVEBIRD
Agapornis roseicollis
14–14½ in
35–37 cm

features of each one. Closely related Psittacus erithacus

f: Psittacidae
A native of African rainforests,
f: Psittacidae
Lovebirds are small African parrots
that live in small flocks. Unlike
f: Psittacidae
This social lovebird from dry
woodland and semidesert
BROWN-NECKED PARROT
Poicephalus robustus
f: Psittacidae
species are placed together for useful red tail
this highly intelligent species is an
accomplished mimic. This has led to
most parrots, this Tanzanian
species builds a nest—a domed
habitats of S.W. Africa
habitually gathers near
The largest of a group of mainly green-gray
parrots from Africa, this bird occurs in forests

comparison, while essential data

its exploitation in the bird trade. structure in a tree cavity. watering holes. from Gambia to the Cape of Good Hope.

highlights unique and interesting

aspects of each organism.
 data sets display key size 4½–9½ ft (1.4–2.9 m)
D ATA
details at a glance, habitat Forests, swamps, scrub thickets, MEASUREMENTS
such as size, habitat, savanna, and rocky landscapes
distribution, and diet distribution India to China, Siberia, the Malay The approximate sizes of the organisms in this book are given in
peninsula, and Sumatra data sets and size boxes. Below is a list of the dimensions used:
diet Mainly hoofed animals, such as deer and
pigs; may also catch smaller mammals and birds MICROSCOPIC LIFE
FEATURE PROFILE Length
Zooming in on single specimens,
feature profiles use close-up PLANTS
photographs to provide in-depth each feature includes a side Maximum height above ground, except for:
portraits of some of the world’s profile of the animal, plant, or fungus Height above water rushes
most spectacular species. Spread aquatic plants
FUNGI
Width (of widest part), except for:
TIGER
furry ears can swivel

Height stinkhorn, dog stinkhorn

independently to scan
surroundings for sounds 593
of prey or danger
Panthera tigr is
The largest and most striking of the big

MAMMALS • CARNIVORES
cats, the tiger is a powerful predator with
almost supernatural grace and agility. Its
natural range extends from the tropical
jungles of Indonesia to the snowy expanses
INVERTEBRATES
of Siberia, where the largest individuals
are found. A full-grown male may weigh
anything up to 660 lb (300 kg), but despite
Adult body length, except for:
Height sponges, feather stars, common hydra, feather
this bulk it can leap up to 33 ft (10 m) in a
single bound. Adult tigers live alone, except size 4½–9½ ft (1.4–2.9 m)
for females with cubs—mothers watch over habitat Forests, swamps, scrub
their young for two years or more, while
hydroid, pink-hearted hydroid, fire “coral” hydroid, tall
thickets, savanna, and rocky landscapes
teaching them vital survival skills. distribution India to China, Siberia,
the Malay peninsula, and Sumatra
diet Mainly hoofed animals, such as
deer and pigs; may also catch smaller

anemones, tall corals

ROUND PUPIL mammals and birds
Unlike small cats, whose pupils
contract to vertical slits, tiger
pupils are always round. They

Diameter snailfur, blue button, eight-ribbed hydromedusa,

expand to provide excellent
sense of smell is surprisingly night vision and contract to
poor, although tigers do pinpricks in bright light.
use scent marks to define
their territory

cup hydromedusa, freshwater jellyfish

WHITE EAR FLASH
The prominent white spot
on the back of each ear is
thought to aid communication.

Diameter, excluding spines echinoderms

Cubs following their mother
will notice ear movements
that may signal danger.

Diameter of medusa jellyfish

STAB AND SLICE
Four long canine teeth deliver the killing bite
Wingspan butterflies and moths
Length of colony bryozoans
to a tiger’s prey. Blade-edged cheek teeth called
carnassials slice through the meat with ease.
dew claw
retracted
claw

Length of shell mollusks, shelled gastropods

Spread of tentacles octopuses
FORELIMBS
The tiger has long legs and
long whiskers allow large feet, which allow it to
the tiger to feel its way run fast, leap large distances,
through dense undergrowth
in almost total darkness and knock prey as large as
oxen to the ground with a
single, deadly swipe.

nonslip pad
FISHES, AMPHIBIANS, AND REPTILES
KILLER IN STRIPES
A fiery orange coat marked
with bold black stripes
provides superb camouflage
PADDED PAW
Tigers have five toes on their front
paws—four of which are
Adult body length from head to tail
as the tiger moves through weight-bearing and the fifth is the
sun-dappled vegetation. The dew claw. The claws are withdrawn
white tigers seen in zoos are fully into the paw when not in use.
usually bred in captivity and
are extremely rare in the
wild. Indeed, tigers have been
TAIL END
The long tail is typically held
BIRDS
hunted almost to extinction; curved just above the ground.
globally, there are fewer than
3,900 left in the wild.
The tiger uses it to aid balance
when chasing prey or climbing. Adult body length from bill to tail
MAMMALS
Adult body length, excluding tail, except for:
Height to shoulder elephants, apes, even-toed ungulates,
odd-toed ungulates

PLANT ICONS
white facial 463
patch with lines
of black feathers
The basic shape of all trees, shrubs, and woody plants is
BIR DS • PAR ROTS A ND COCK ATOOS

15 in 16 in
38 cm 40 cm

described using one of the following symbols. Herbaceous

perennials which die back each winter are not given symbols.
BLUE-FRONTED PARROT ST. VINCENT PARROT ST VINCENT PARROT TREES SHRUBS
Amazona aestiva
f: Psittacidae
Amazona guildingii
f: Psittacidae Amazona guildingii
f: Psittacidae Broadly columnar Bushy, mound-forming
34 in
85 cm A member of a large group Several Caribbean Amazona parrots
of mostly green parrots, this are faced with extinction. This
BLUE-AND-YELLOW powerful bill species occurs in open forest large species is being conserved by
MACAW habitats of central-eastern an ongoing breeding program
Broadly conical Bushy, suckering
Ara ararauna
f: Psittacidae
S. America. in its native island of St. Vincent.
species’ common names are
Macaws are large, long-tailed
parrots with sparsely feathered 43/4–5½ in
12–14 cm highlighted in bold, scientific Large weeping Compact, bushy
facial patches. One of two blue and
yellow species, this bird comes
from northern S. America. names are in italics; in some Small weeping Erect, treelike
31–35 in
79–89 cm
9½–11 in
24–28 cm
cases the family (F) name Multi-stemmed tree Loose, open
is given below Narrowly columnar, Open, spreading
22–23½ in
55–60 cm flame-shaped Rounded, bushy
bright yellow
BLUE-HEADED PARROT
Pionus menstruus
PACIFIC PARROTLET
Forpus coelestis Narrowly columnar Spreading, prostrate
feathers on f: Psittacidae f: Psittacidae
blue wings A relative of the Amazona
parrots, this small parrot is
Parrotlets are tiny green American
parrots; only New Guinean pygmy
12 in Narrowly conical Upright
RED-FRONTED SCARLET MACAW common in lowland forests parrots are smaller. This species
MACAW
Ara rubrogenys
f: Psittacidae
Ara macao
f: Psittacidae
Like other macaws, this
from Costa Rica to Bolivia. occurs in western Ecuador and Peru.
30 cm Rounded, broadly Upright, arching
columnar
This small macaw is restricted noisy, flocking bird has a
to the arid scrub of central
Bolivia. Its small population
heavy bill for breaking nuts
and palm fruit. Its range
12 in
30 cm
white eye-ring
size boxes give the most Upright, vigorous,
is threatened by habitat loss
and wildlife trafficking.
extends from S. Mexico
to C. Brazil. appropriate measurement Rounded, broadly bushy
31/4 ft
1m
long, red tail
JANDAYA PARAKEET
Aratinga jandaya
f: Psittacidae
for the organism they spreading Sprawling, climbing
HYACINTH MACAW
Anodorhynchus hyacinthinus 8–10 in
Aratinga is a genus
of predominantly green accompany (see panel, right) Single-stemmed palm
20–25 cm “mini-macaws” found in
f: Psittacidae northeastern Brazil. Some
Unlike other macaws, this giant Brazilian
parrot has its facial patch reduced to an
have extensive splashes
of golden coloration. Multi-stemmed palm,
eye-ring—similar to that of the related
Aratinga parakeets. cycad or similar
10 in
25 cm

17½–18 in
44–46 cm
11½ in
29 cm
red belly A B B R E V I AT I O N S

YELLOW-CHEVRONED
MAROON-BELLIED
PARAKEET sp.: species (used where species name is unknown)
PARAKEET MONK PARAKEET Pyrrhura frontalis
BURROWING PARAKEET
Cyanoliseus patagonus
Brotogeris chiriri
f: Psittacidae
Myiopsitta monachus
f: Psittacidae
f: Psittacidae
Relatives of Aratinga, most mya: million years ago
f: Psittacidae Native to central S. America, This colonial breeder from Pyrrhura parakeets have
A Patagonian relative of macaws,
this colonial breeder nests in
escaped cagebirds of this
species have established
temperate S. America is
the only parrot to build a
prominent splashes of
maroon or red, such
h: hardness of a mineral, measured on the Moh’s scale
tunnels in earth banks. It forms feral populations in stick nest, often a large as this species from
faithful pair-bonds, unlike most birds. warmer parts of the US. communal structure. eastern S. America. sg: specific gravity—a mineral’s density is measured by
comparing its weight to that of an equal volume of water
Living
Earth
Our blue planet, spinning in the vastness of space, is the only
proven home of living things. Over nearly 4 billion years, life has
evolved from the simplest of beginnings. Most species have become
extinct, but life itself has flourished, endlessly diversified, and
recovered from repeated extinctions. The result is an extraordinary
variety of living things, which scientists continue to study as they
piece together the story of life on Earth.
12 A LIVING PLANET WATER AND LIFE
Life depends upon water, which forms
more than 50 percent of all living tissues.
Earth is uniquely equipped to support a wide diversity of life, Most rainfall is driven by evaporation
both on land and in the seas. Without heat and light from the from the oceans, which hold 97 percent
LIVING EARTH • A LIVING PLANET

of Earth’s surface water, and a vital

sun, plentiful supplies of water, the protection provided by network of rivers flow in all but the
the atmosphere, and chemicals from the rocks that form the world’s hottest, coldest, and driest places.
basis of Earth’s ecosystems, life would perish.

DYNAMIC EARTH
Within our solar system, Earth seems to be uniquely placed to support abundant
life. The third planet from the sun, Earth is neither too close to nor too far from
the sun’s heat. It therefore retains an outer atmosphere of oxygen and other gases,
and a hydrosphere of plentiful surface water. Together, these form a protective,
insulating layer that enables life to flourish. In contrast, the other planets in the
solar system are either too hot or too cold, and devoid of the levels of water and
oxygen required to support detectable life.
Earth has a layered structure, with an extremely hot, solid metallic core at its
center, surrounded by an outer molten layer. This, in turn, is surrounded by a thick
and hot silicate mantle, which rises to a thin, cool, and brittle outer crust. The
mantle is constantly churned by heat rising from the core, which generates new
ocean floor rocks and breaks the crust
outer
into large “plates.” Over geological core
time, some oceans expand, carrying
continents with them. Other inner
core
oceans become smaller as old, cool
crust descends into the mantle.
Continents collide to form
mountain ranges. Life has had
to adapt to the constant change
in the Earth’s environment.
upper
mantle
EARTH’S STRUCTURE
The liquid mantle is constantly
stirred by heat rising from the core. lower
This moves the plates of the outer mantle
crust, causing earthquakes and
volcanic eruptions on the surface. crust

SUN AND MOON

The sun and the moon both have a direct
impact on life on Earth. Without the sun’s
energy, in the form of heat and light, there
would be no life. Solar energy heats Earth’s
atmosphere, the oceans, and the land,
producing our varied climate. Because Earth
rotates at an angle while it orbits the sun, the
sun’s radiant energy is unevenly distributed
over its surface. This results in daily, seasonal,
and annual variations in light, heat, and living
conditions for plants and animals. Even at
the equator, there are marked temperature
changes between day and night. The orbit of
SOLAR FLARES Earth’s satellite moon and its gravitational pull
The sun’s energy is dramatically raise tides in Earth’s oceans and seas. Tidal
released from the surface in periodic
explosions, which heat its atmosphere cycles are especially influential on coastal life,
to form solar flares of hot ionized gas. which has to adapt to changing conditions.
FRAGILE ATMOSPHERE
Earth’s atmosphere is 75 miles (120 km) thick. It is made 13
up of several layers, each with its own temperature and gas
composition. Its density decreases with height, until it

LIVING EARTH • A LIVING PLANET

becomes the sparse, outermost layer, called the ionosphere.
The ozone layer, in the lower atmosphere, plays a vital role in
protecting life, because it absorbs harmful radiation such
as ultraviolet light, which damages living cells. Before the
ozone layer formed, life was confined to the seas, whose
waters offered some protection against ultraviolet light.
BLUE PLANET
The majority of water vapor and weather activity is About two-thirds of Earth’s surface is
restricted to the lowest 10 miles (16 km) of atmosphere, covered with water, which supports
known as the troposphere. Earth’s surface water and gaseous the abundance and diversity of life.
atmosphere interact to recycle water from the surface into
the atmosphere and—through clouds, rain, and snow—
redistribute it over the land
and sea. From the land, water
flows back into the sea, other gases, argon
including carbon 0.9%
although large quantities are dioxide, methane,
held back in lakes, ice, and and ozone
oxygen 21%
under the ground. nitrogen 78%

ATMOSPHERIC GASES
Nitrogen and oxygen make up over LAYERS OF ATMOSPHERE
99 percent of Earth’s atmosphere, Earth is surrounded by a thin, layered
along with small but important atmosphere, composed of water vapor
volumes of water vapor, carbon and various gases, which trap solar
dioxide, and several other gases. energy and heat the surface.

VARIED ROCKS
There are about 500 different kinds result from the alteration of existing rocks
of rocks on Earth, made up of varying within Earth’s crust. These different types of
combinations of thousands of naturally rocks are exposed on the surface by a
occurring minerals. All rocks have a mixture of uplift, driven by Earth’s moving
specific composition and properties, and crust, and surface processes such as
can be divided into three main categories: weathering and erosion. Erosion also
igneous rocks were originally molten; modifies the rocks to produce multiple kinds
sedimentary rocks are deposited on the of landforms, soils, and sediments. These are
Earth’s surface; and metamorphic rocks the inorganic elements on which life depends.

IGNEOUS ROCKS METAMORPHIC ROCKS SEDIMENTARY ROCKS

The cooling and solidification of The application of heat and Layers of sediment and the remains
molten rock produces crystalline pressure to existing rocks deep of plants and animals are deposited
igneous rocks. Their composition within Earth’s crust can change by wind and water. Over time,
and texture vary—rapid cooling their form and mineral composition, older and deeper buried sediments
produces fine resulting in undergo compression
grained rocks, metamorphic and chemical
and slow cooling rocks such as change which
produces coarse slate, schist, transforms
grained rocks. and marble. them into rock.
biotite
basalt schist sandstone
14 AC T I V E E A RT H
Earth’s surface is constantly changing, thanks to dynamic geological processes
driven by its internal heat energy. The plates of Earth’s brittle outer crust are
LIVING EARTH • ACTIVE EARTH

always in motion, altering the shape of oceans and continents as they do so.

PLATE TECTONICS
Over geological time, Earth’s surface—and the to form new ocean floor crust. This spreads away
distribution and size of the continents and on either side of the rift to form divergent plate
oceans—has constantly changed, driven by the boundaries, mainly beneath the oceans. And, since
process of plate tectonics. The cool and brittle Earth itself cannot expand, the creation of new
rocks of Earth’s outermost crust are broken into a oceanic crust requires that the crust is shortened
number of semirigid slabs known as tectonic plates. elsewhere by the same amount. This reduction SAN ANDREAS FAULT
There are seven major continent-sized plates and occurs at convergent boundaries where either one Stretching some 810 miles (1,300 km)
about a dozen smaller ones. Over time, crustal plate overrides the other—a process known as through California, this dramatic fault
is the product of a transform boundary
plates have jostled against one another. Hot magma subduction—or the plate margins are compressed between the Pacific and North American
rising from the mantle erupts from rifts in the crust and buckled to form mountains. plates, which slide against one another.

PLATE BOUNDARIES MAP KEY plates moving

ridge appears at site of This map shows the main plates that
new plate formation Convergent boundary together
form the jigsaw surface of Earth’s Divergent boundary
crust. A study of earthquake Deep-sea trench
plates moving locations worldwide established Transform boundary the thinner,
apart where the plate boundaries lie. denser plate is
forced down into
Earth’s crust

eurasian plate
north american
plate
DIVERGENT BOUNDARY SUBDUCTION ZONE
As plates are pulled apart they arabian Where two plates converge and
stretch and break, forming faulted plate indian philippine
pacific
one plate is thicker, it pushes
plate plate
rifts and volcanically active ridges. african plate
plate the thinner plate beneath it in
pacific plate a process called subduction.
south american
nazca plate
australian
plate
plate
plates
moving elevated
plates moving together mountain
past each other ridges form
antarctic plate

TRANSFORM BOUNDARY CONVERGENT BOUNDARY

Also known as conservative plate Where two converging plates are of similar
boundaries, these occur when two plates density and thickness, their leading edges are
slide horizontally past one another, neither crumpled, faulted, and thickened to form
destroying or creating new crust. mountain ranges such as the Himalayas.

FOLD MOUNTAINS
The intense pressure caused by MOUNTAINS AND VOLCANOES
converging plate margins folds, One of the major factors controlling the movement and distribution
faults, and thickens the rocks of of life on Earth is its varied topography—its surface features—
the crust. Crustal thickening
elevates mountain ranges. including the towering mountains and volcanoes on land and under
the sea. On land, mountains not only hinder the movement of
wildlife but alter weather, climate, and local plant life, which in
turn impacts upon animal life. Active volcanoes also affect their
ACTIVE VOLCANOES surroundings when they erupt—initially by destroying life, but
Most volcanoes form at plate margins. also, in the longer term, when the weathering and erosion of erupted
Deep down, rocks melt to produce hot lava and ash provide new mineral nutrients that fertilize the area.
magma which rises and erupts at the
surface. Even dormant volcanoes may one Submarine mountains and volcanic eruptions affect the movement
day erupt as plates shift beneath them. of water, marine life, and the fertility of ocean waters.
 WORN AWAY
Weathering by wind and water can
result in the significant erosion of rocks
and the sculpting of land surfaces, as
seen here in the dramatic landscape
of Bryce Canyon, Utah.

WEATHERING AND EROSION SCIENCE

S O I L F O R M AT I O N
Many rocks are formed below Earth’s surface, is loosened, dissolved, and transported away is
and when they are exposed by uplift from pressure known as erosion. The combination of weathering The production of a soil requires the
within Earth’s crust or by retreating seas or rivers, and erosion wears down Earth’s rocky surfaces, initial weathering and erosion of
they react in many ways with the atmosphere, water, layer by layer. Exposed rocks on mountain tops and the parent rock, which is broken down
and living organisms. The physical and chemical on the exterior of buildings, for example, are subject into small, mineral-bearing particles
processes resulting from the interaction of rocks to chemical weathering by acid rain and to physical known as regolith. The addition of
humus—organic matter formed from
and minerals with the atmosphere are known as weathering by temperature change and the
the remains of plants and animals—
weathering. The processes by which rock material splintering freeze-thaw effect of ice formation. Bare forms the basis for a soil. The soil
rock surfaces may also be physically eroded by sand in turn becomes the bed in
particles carried by the wind. The combined effect which more life flourishes.
of weathering and erosion dissolves some rocks, and
reduces others to fragments. As rock debris is broken plants flourish
in soil bed
down and transported by wind, water, and ice,
the sediment created becomes increasingly available
humus-rich
to life forms, providing important mineral nutrients layer
and a new surface for anchorage and growth.
mineral-rich
layer of
LANDSLIDE, RIO DE JANEIRO regolith
Even with a well-developed plant cover, the
impact of high rainfall on an area of steep slopes
has the potential to cause landscape-altering parent rock
and even life-threatening occurrences such as
avalanches and landslides.
16 C H A N G I N G C L I M AT E S
The features of our seasons—for example, hot, dry summers and cold, icy
winters—make up the climate of each region. Earth’s climate has always
LIVING EARTH • CHANGING CLIM ATES

changed at different rates, places, and times, and this variation in conditions
has considerable and continuing impact upon the evolution of life.

WHAT IS CLIMATE?
Climate is a region’s average weather over a long in different regions of the world.
period of time, produced by all the atmospheric For instance, there is a major
conditions such as temperature, rainfall, wind difference between climates arctic fox
in summer
strength, and pressure. The climate of any given of polar regions, which receive
place is also partly controlled by other factors the least light and heat, and
such as human input, height above sea level, local of tropical regions around the
topography, proximity to seas and oceans—with equator, which receive the most.
their prevailing winds and
water currents—and, most c h a ng i ng c on di t ion s
importantly, its latitudinal Global climates are generally classified according
position between the to the average temperature and the amount of
equator and pole. Latitude rainfall each area receives, and their combined arctic fox
in winter
controls the amount of effect on plant growth. For instance, equatorial
solar radiation received regions at present are hot and wet because oceans SEASONAL ADAPTATION
dominate there, whereas deserts are dry and polar Annual climate change can bring acutely
CHANGING TREE LINE regions are cold. However, this has not always been different living conditions from one season
As the air temperature declines the case. Factors controlling weather conditions to the next. Animals and plants have various
with increasing altitude, plant life means of adapting to these changes; for
changes. Broad-leaved trees are over geological time have affected the climate of example, the Arctic fox grows a thick coat
replaced by conifers, then shrubs. the planet, from ice ages through to global warming. in winter, which it sheds in summer.

LIFE IN THE DESERT

Plants such as cacti have adapted to life
in semiarid environments by slowing
growth, reducing the leaves to spines (to
stop water loss from transpiration), and
evolving special tissues that retain water.
 EVIDENCE OF CLIMATE CHANGE
Studies of polar ice core samples have revealed 17
details of past climate change. The chemical analysis
of trapped gas bubbles helps provide an approximate
measure of the temperature of the atmosphere at
the time the ice was formed.

LIVING EARTH • CHANGING CLIM ATES

ICE CORE
SAMPLE
This is a close-up of
an ice core sample
recovered from Lake
Bonney, Antarctica,
which has a permanent
ice cover. It shows
trapped bubbles of air
and sediment particles
from the lake bed.

CLIMATE CYCLES SCIENCE

S T U D Y I N G S T O M ATA
There is clear evidence from rocks and fossils size of these huge landmasses affected regional
that Earth’s climate has changed significantly over climates. In addition, the changing shape of ocean Plant growth depends upon an
time and that this has affected the evolution and basins has altered water circulation, which in exchange of gases between the
distribution of life, and led to the extinction of turn changes the temperature and humidity of atmosphere and the plant tissue,
many species. There are a number of causes of this the atmosphere above, and so affects climate. through special gaps between the
natural climate change, including volcanic activity— leaves’ cells called stomata. The
stomata open and close to take in
which pollutes the atmosphere with ejected gas and g r e e n h o u s e a n d i c e ag e s
carbon dioxide for photosynthesis
dust—and changes in ocean currents, which move Long-term changes in climate are divided into cold and allow waste water and oxygen to
heat around the globe. Change is also driven by the periods—ice ages, when there were long-lasting escape. In general, plants adapt to high
orbital and rotational cycles of the planet, which ice sheets at the poles—and warmer, greenhouse levels of atmospheric carbon dioxide,
affect the amount of solar radiation that reaches periods with largely ice-free poles. Warm phases associated with warm “greenhouse”
Earth’s surface. This, in turn, influences the planet’s are linked to the release of greenhouse gases, such climates, by evolving high densities of
temperature and climate, and triggers Earth’s cycles as carbon dioxide, by plants into the atmosphere. stomata on the leaf surface. Fossil
of ice ages and greenhouse periods. These trapped heat in Earth’s atmosphere and evidence of changes in the stomatal
produced huge, shallow seas, arid zones, and lush density of certain plants tracks changes
in atmospheric carbon dioxide over time.
ch angi ng geogr a ph y forests that, in the time of the dinosaurs, provided
Over time, continents have been displaced as them with plentiful food. Ice ages, lasting millions
oceans have expanded and contracted due to plate of years, can be traced from the impact that
tectonics. As continents moved from one hemisphere glaciation has left on the landscape. Fossils show
to another they passed through different climatic how rapid climate change associated with ice ages
zones and, at times, formed supercontinents. The had a dramatic impact upon life on a global scale.

CARBON DIOXIDE
AND TEMPERATURE
Gas bubbles trapped in polar ice cores indicate
the fluctuating temperature of the planet—the
higher the amount of carbon dioxide detected
in the ice core, the higher the atmospheric eucalyptus stomata
temperature would have been at that time.
CO2 (purple) Temperature
(blue)
PARTS OF CARBON DIOXIDE (CO2) PER
MILLION PARTS OF ATMOSPHERE

TEMPERATURE CHANGE ºC (ºF)

300 7.2 (4)

280 3.6 (2)
260 0 (0)
-3.6 (-2)
240
-7.2 (-4)
220
-10.8 (-6)
200 -14.4 (-8)
180 -18 (-10)
DEVONIAN CORAL REEF
Earth’s changing climate is spectacularly illustrated by this limestone outcrop in
The Kimberleys, W. Australia. In the Devonian Period (400 million years ago), 400,000 350,000 300,000 250,000 200,000 150,000 100,000 50,000 0
the area was under water and the cliff was a barrier reef. YEARS AGO
18 H A B I TAT S F O R L I F E KEY
Earth’s uniquely varied habitats enable it to support an abundantly Polar region Coniferous forest
Desert Mountains
rich diversity of animal and plant life—from the extreme depths of
LIVING EARTH • HABITATS FOR LIFE

Grassland Coral reef

the ocean floors to the highest mountains, and from arid deserts Tropical forest Rivers and wetland
and grasslands to the warm, wet tropics. Temperate forest Oceans

Each life-form has its preferred habitat—the

one to which it has adapted over thousands A R C T I C
or even millions of years. However, Earth’s O C E A N
varied environments allow many different
kinds of animals and plants to live in the same
habitat—a phenomenon known as biodiversity. G R E E N L A N D
Over geological time, life has had to adapt
to environmental change, especially rapid
climate change, with both extinctions and the Arctic Circle
evolution of life-forms to colonize new habitats.
The presence of these pioneering organisms, in
turn, produced changes in the environments N O R T H
they colonized—by forming soils, for A M E R I C A
instance—and these modifications encouraged EUROPE
further colonization by new life forms.
Differences in habitats are produced

A T
by many factors, ranging from an area’s
height above sea level, its distance from

L A
Tropic of
the equator, and its topography (or physical Cancer
shape). Some areas of the globe are

N T
A F R I C A
biodiversity “hotspots,” rich in animal and P A C I F I C
plant life—most notably tropical reefs and O C E A N

I C
rainforests—while other areas with more
extreme conditions support only a few, Equator
though often richly populated, species.

O
S O U T H
S O U T H E R N A M E R I C A

C
SCIENCE Tropic of O C E A N

E
LEVELS OF LIFE Capricorn

A
Organisms rarely exist entirely on their own, even

N
in the remotest places on Earth. The interactions
between wildlife create different levels—from the MAP OF BIOMES
individual organism to the overall ecosystem which A biome unites ecosystems that have
it inhabits and shares with others. developed under similar climatic and
soil conditions in different parts of the
individual world. Biomes are defined by a variety
A single, usually independent of factors such as plant types, climate,
geology, and topography.
and habitat-restricted member
of a population.

population
Antarctic Circle
A group of individuals of the same
species that occupy the same area
and interbreed.

community
A naturally occurring collection of GRASSLAND
plant and animal populations living The evolution of grass plants some 20
million years ago, and the colonization by
within the same area. grazing mammals, transformed Earth’s
landscapes. Temperate grasslands are
ecosystem
generally treeless and have extremely
A biological community and its fertile soils. Savanna grasslands, as shown
physical surroundings, which here, are more like open woodland,
support one another. featuring scatterings of trees and scrubs.
bison
 DESERT TROPICAL FOREST
Extreme lack of rain and soils for sustained The richest wildlife habitats on
plant growth creates deserts, which at present land are found in the forests of 19
account for about a third of Earth’s landscapes, the tropics —Earth’s
although this proportion is increasing. The hottest areas, which lie by
largest desert is the the equator. Their numerous

LIVING EARTH • HABITATS FOR LIFE

African Sahara. ecosystems are important
strawberry poison-
yet increasingly vulnerable dart frog
biodiversity hotspots.
rattlesnake

TEMPERATE FOREST
Temperate environments lie between
the tropics and the polar regions.
A R C T I C O C E A N The influence of both tropical
and polar air masses promotes
vast forests with considerable
biodiversity. However, clearing
by humans has greatly reduced
their extent.
red deer

CONIFEROUS FOREST
Coniferous forest trees, such as
redwoods, spruce, and fir, belong
A S I A to an ancient plant group and are the
world’s toughest trees. Evergreen
with small leaves, they thrive where
few other trees can, in cold areas
and in mountain ranges.
brown bear

MOUNTAINS
Reaching as high as 5½ miles (9 km)
above sea level, Earth’s mountains
are home to many different
environments. A single mountain
P A C I F I C can rise from temperate woodland
O C E A N up to arctic conditions as the
climate changes with altitude.
peregrine falcon
I N D I A N
RIVERS AND WETLAND
O C E A N A wide range of animal and plant
life thrives in Earth’s rivers and lakes.
Landscapes saturated with water, either
A U S T R A L I A permanently or seasonally, form
distinctive wetlands, where
areas of open water mix with
dense vegetation.
dragonfly

S O U T H E R N O C E A N
CORAL REEF
Coral reefs develop from the
skeletons of marine organisms
in shallow, sunlit tropical
waters. Supporting an
immense variety of
life, they are the
rainforests of the
A N TA R C T I C A aquatic world.
yellow tang fish

POLAR REGIONS OCEANS

The Arctic and Antarctic experience Life in oceans is found at all levels, from the sunlit
extreme seasons, with 24-hour daylight surface to the deepest depths. Covering two-thirds
in summer and perpetual darkness in of the planet, oceans form the largest continuous
winter. Dominated by snow and ice, habitat on Earth and support lobster
and with vast, dry areas of polar very varied life forms—
desert, both are highly vulnerable from microscopic plankton
to climate change. to the largest living mammal,
the blue whale.
rockhopper penguin
20 H U M A N I M PAC T
The rapid growth of the human population has had an
enormous impact upon Earth’s natural environments,
LIVING EARTH • HUM A N IMPACT

affecting the climate and countless species of plant

and animal life. Some of these changes are irreversible.

ENVIRONMENTAL CHANGE
Earth has a long history of climate change with dominant warm
“greenhouse” climates, widespread forestation, and no polar ice
punctuated by glacial “ice ages.” Global warming is known to be
SCARRED LANDSCAPE
linked to high levels of greenhouse gases in the atmosphere, such as
The growth of industry has required carbon dioxide and methane, which trap incoming solar energy and
the exploitation of raw materials. Their raise the temperature of the oceans, land, and air. In the past,
extraction, such as at this copper mine, natural increases in atmospheric carbon dioxide have been balanced
has changed global landscapes forever.
by the development of forests on land and lime-rich sediments in
the seas, which eventually became coal and limestone and stored
excess carbon dioxide effectively. Since the industrial revolution
of the 19th century, human activity has released huge amounts
of carbon dioxide and other greenhouse gases back into the
atmosphere through the mining and burning of fossil fuels,
the clearing of forests, and the raising of cattle.

the oceans
The health of Earth’s oceans is vital to all life. Marine life depends
upon the circulation of oceanic waters that contain enough oxygen
and nutrients to support the food chain from plankton and shellfish
to all the other animals that depend upon them for food. Fossil
records show that when the oceanic environment deteriorated
in the past, it led to extinctions of life. Today, overfishing and
ATMOSPHERIC POLLUTION pollution, especially with plastics, by humans are affecting
Slash-and-burn forest clearing for the marine environment.
agriculture not only releases atmospheric
pollutants but also diminishes the capture
of carbon dioxide by the plant life. t h e at m o s p h e r e
For thousands of years human activity has affected the
GREENHOUSE EFFECT atmosphere. Initially, it was restricted to the release of pollutants
An excess of greenhouse gases in the from domestic fires and forest clearance. In Roman times metal
atmosphere creates a shield which
prevents some of the sun’s energy production released the first industrial pollutants into the
from radiating back into space. atmosphere, leaving traces that can be seen in polar ice cores. POLAR ICE SHELF COLLAPSE
Over the last 200 years, pollution by gases and particulates Rising temperatures are leading to the
some heat is reflected collapse of the polar ice shelves. The release
back into space has risen steeply. This has produced acid rain and smog, and of such large volumes of water is raising sea
infrared radiation
greenhouse gases that are linked to global warming and levels, which in turn threatens coastlines.
transfers heat energy depletion of the ozone layer, which helps screen out
from the sun through some of the infrared harmful ultraviolet radiation.
Earth’s atmosphere radiation is reflected
off greenhouse gases
back to Earth’s surface the land
infrared radiation
is reflected back Since settlements and agriculture
off Earth’s surface
Earth’s became widespread 8,000 years
surface also ago, humans have had a growing
radiates heat impact upon Earth’s landscapes.
some heat is
reflected back Worldwide population growth,
Earth’s into space land settlement, and demand
surface
warms up for food has left few untouched
landscapes. Greater awareness
of the impact of human
heat absorbed by activity on the environment
greenhouse gases radiates
back to Earth’s surface is now leading to efforts to
conserve natural habitats.
 AGRICULTURE
Natural landscapes are radically
altered by intensive farming, such
as the cultivation of rice in irrigated
terraces throughout Asia. Such
methods support big populations,
but require large amounts of water.

EXTINCTION SCIENCE
The inability of many organisms to adapt to environmental change MAMMOTHS NO MORE
has led to a huge turnover of species throughout geological time. Woolly mammoths were elephants that
In fact, the vast majority of nature’s species are now extinct. were adapted to the cold. They migrated
Only the fittest organisms survive, usually through gradual across ice-age Europe and Asia in vast
adaptation, but sometimes by the sudden elimination herds. Archaeological evidence such as
of competitors. For example, when a huge asteroid cave paintings shows that they were
hit Earth 66 million years ago, it set in motion a actively hunted by humans about
chain of events that led to the demise of many 30,000 years ago, which may have
life-forms, including the dinosaurs on land and DECLINE OF THE contributed to the extinction of most
CRESTED IBIS woolly mammoths by 11,000 years ago.
the ammonites in the seas. But it also opened Once widespread in Asia, hunting and
the way for the rapid evolution of mammals, habitat loss has restricted the crested
including humans. More recently, the arrival ibis to a small wild population in
of modern humans in different regions around China. Captive breeding has allowed
it to be reintroduced to Japan.
the world has contributed to the extinction of
particular species, such as the woolly mammoth PÈRE DAVID’S DEER
in Europe and Asia (see far right). Today, as the Extinct in the wild, this east Asian deer
human population expands, our actions are has survived only in captive herds bred
in England since 1900. Père David’s deer
placing a growing number of species, such as reintroduced to China in the 1980s now cave painting, pech merle, france
the tiger, in imminent danger of extinction. form a population some 700 strong.
22 ORIGINS OF LIFE EARLY LIFE
The very earliest forms of life evolved
The fossil record shows that life first appeared on Earth at least in the seas, and evidence of this comes
3.7 billion years ago, and that all complex life has evolved from these from two main sources: living primitive
LIVING EARTH • ORIGINS OF LIFE

first simple forms. Today, diverse life-forms range from single-celled organisms and the fossil record. The
most primitive life-forms today are
organisms to mammals with a complex anatomy. archaea and bacteria, which can survive
even in extreme temperatures and acidic
WHAT IS LIFE? conditions. Such microorganisms may
Several features distinguish living organisms be similar to those that first evolved in
from inanimate, inorganic matter, although the the extreme environments of early Earth.
existence of viruses blurs the boundary between Fossil evidence for early life is
the living and nonliving. These features contentious. Chemical evidence of carbon
include an ability to take in and expend energy, derived from living organisms has been
to grow and change, to reproduce, to adapt to found in rocks from Western Greenland
its environment, and—in more complex living and dates back some 3.7 billion years.
organisms—to communicate. The most convincing of ancient records
The cell is the fundamental unit of life, of life are layered, mound-shaped
capable of replicating itself and carrying out all stromatolites (see right).
living processes. Even the smallest independent PHOTOSYNTHESIS
Plants use the pigment chlorophyll to capture light energy
organisms are made up of at least one cell, and and convert water and carbon dioxide into sugars and
almost every cell of every living organism has oxygen. This benefits other life-forms that feed on the
its own set of molecular instructions. Within plants and breathe in oxygen. FLOURISHING LIFE
which are responsible for the particular Ever since life first appeared on Earth, it
each cell, the threadlike chromosomes carry has thrived in the sea, evolving into the
hereditary information in the form of genes, characteristics of an organism. The set biodiversity hotspots of modern, sunlit
of instructions in a gene are mainly reefs, second only to rainforests for the
recorded in the form of a molecule variety and density of life they sustain.
called chromosomal deoxyribonucleic
acid (DNA). The DNA of an organism
carries information from one generation to
another, allowing certain characteristics
to be passed on from parent to offspring.
VITAL ENERGY di v i s ion s of l i f e
To sustain itself, life must obtain energy from the The huge array of life on Earth is divided into
environment. Energy flows mostly from photosynthesis
in plants—but also occasionally from chemosynthesis in three domains or superkingdoms—Archaea,
bacteria—to the animals that feed upon them and are, Bacteria, and Eukaryota –which encompass
in turn, eaten by other animals. all life-forms from plants and fungi to
animals. Archaea and bacteria are primitive,
SCIENCE single-celled organisms, and were probably
VIRUSES
the earliest form of life on Earth. The more
Viruses are the most abundant biological entity on advanced eukaryotes are distinguished from
Earth, and they lie at the boundary between living and archaea and bacteria by having a cell nucleus,
nonliving. While they have features in common with which contains the cell’s
living organisms—they are made of genetic material genetic material, DNA.
and are protected by a protein coat—viruses are Eukaryotes vary
parasitic and can only reproduce within the living cells
of other organisms. They are tiny packets of chemicals
enormously in shape
that copy themselves, without truly being alive.
and size, ranging
from single-celled
organisms to complex,
multicelled plants
and animals.

GROWTH
A capacity to grow and repair
is one of the key defining
features of life. All organisms,
from simple fungi to mammals,
sars-cov-2 virus, the cause of covid-19 grow mostly by cell division
but also increase in cell size.
 STROMATOLITES
For billions of years these layered structures have built up
in shallow tropical seas by alternating layers of sediment 23
and sheets of microorganisms, including cyanobacteria
(blue-green algae).

About 650 million years ago, fossils of various

frond- and disk-shaped, soft-bodied, and mostly
sessile organisms appeared. These marine
animals were known as the Ediacaran Fauna.
By the beginning of the Cambrian Period,
545 million years ago, numerous multicelled BURGESS SHALE
The fossils of the Burgess Shale
marine organisms had evolved, including in Canada show that marine
From the first appearance of simple life, it burrowing worms and a variety of small, life diversified rapidly in
took at least 2.7 billion years before complex shelled mollusks, whose bodies had muscle Cambrian times, from sponges
and arthropods to vertebrates.
life-forms appeared. The fossil of a microscopic tissues and organs such as gills for respiration.
and multicelled red algae called Bangiomorpha About 510 million years ago, the first WIWAXIA
provides the first evidence of the existence of vertebrates appeared, with an internal skeletal Found at the Burgess Shale,
specialized cells. These cells evolved for sexual support for the body. By late Devonian times, Wiwaxia was a 2 in (5 cm) long,
mollusklike seabed crawler
reproduction, and also for the development of about 380 million years ago, vertebrates had with spines and scales,
a holdfast to attach the algae to the sea floor. begun to emerge from the seas onto land. and a soft underside.
24 E VO LU T I ON A N D D I V E R S I T Y
Until the 19th century, when a number of theories were proposed, it was a matter of
speculation as to how such remarkably diverse life-forms had developed on Earth. Today,
LIVING EARTH • EVOLUTION AND DIVERSITY

the theory of evolution and diversification, alongside geological evidence for changes in the
distribution of continents, give a fascinating insight into the ever-changing life on our planet.

CHANGE OVER TIME EVIDENCE OF EVOLUTION

Comparison of the anatomy of vertebrate limb bones from various
All living things can change and adapt to their surroundings. species show that, despite different appearances and functions, they
Changes that are passed from generation to generation are derive from the same basic developmental plan and the same genes.
hard to see, but over time they can alter the way a species FROG upper arm
looks or behaves. This process, known as evolution, is not The frog’s leg, arm, and bone
gradual but has seen significant extinctions and bursts of finger bones are modified finger
for swimming. Large bone
growth over time. muscles enable it to jump
The study of fossils to unravel the history of life was in powerfully—essential
its early stages in Charles Darwin’s day, and since then a vast for catching prey and
escaping from predators.
amount of information supporting the theory of evolution
has emerged. We now know that life evolved in the oceans
at least 3.7 billion years ago, and that it was from these early OWL upper arm
The wing of a bird bone
simple life-forms that all current life on Earth evolved— is powered by flight
including plants, fungi, and animals. muscles attached to finger
the upper arm and bone
As life-forms became more complex and moved from lower
bones of the wrist,
sea to land, the first forests and land-living animals evolved. with greatly modified arm bone
The Mesozoic Era, which began 252 million years ago, with and extended fingers.
its successions of evolving plants and animals, produced the
dominant dinosaur reptiles and their bird descendants. A CHIMPANZEE upper
mass extinction event 66 million years ago ushered in the finger arm bone
The arm of a chimpanzee bone
Cenozoic Era. The reptiles were largely replaced by mammals is anatomically very
similar to our own, lower
both in the seas and on land, where flowering plants and but has slightly different
arm bone
their pollinating insects also became abundant and diverse. proportions, with
elongated fingers wrist
and a short thumb. bone

shoulder
DOLPHIN finger blade
GIANT SALAMANDER The arm bones of bone
whales and dolphins upper
This extremely rare fossil arm bone
skeleton of an Andrias (giant form a flipper—with
salamander) was mistaken for shortened, flattened, and
a human victim of the biblical strengthened arm bones
Flood until French anatomist and greatly lengthened
Georges Cuvier identified it as second and third fingers. lower arm bone
an amphibian in 1812.

LAMARCK LEADS THE WAY

The 18th-century French biologist Jean-Baptiste Lamarck
developed the first comprehensive evolutionary theory that higher
forms of life had “evolved” from simpler organisms. Based on his
extensive research of invertebrates in particular, Lamarck argued
that necessary characteristics could be acquired during an
organism’s lifetime—through the desire to gain food, shelter, and
mates—and those not needed could be lost, with the resulting
transformation passed on to its offspring. Although modern
genetics disproves this theory of “soft inheritance,” Lamarck’s INNER STRIVING
concepts were an important starting point, and were developed Lamarck believed that evolution
further by the Scottish anatomist Robert Grant, who tutored worked through a process of
Charles Darwin in Edinburgh. Darwin himself did not entirely “inner striving.” Thus the giraffe
developed a long neck to reach
rule out a Lamarckian mechanism, which he thought might be the leaves of trees, and the heron
supplementary to natural selection. grew long legs in order to wade.
 25

LIVING EARTH • EVOLUTION AND DIVERSITY

butterfly
GALAPAGOS FINCHES wing
On his voyages, Darwin collected
many different specimens of Galapagos
finches which he believed may have
descended from a common ancestor.
DARWIN AND WALLACE
In the mid-19th century, British naturalists his explanations for the geographical distribution
Charles Darwin and Alfred Russel Wallace of organisms—biogeography—and realized the
independently came up with the theory of evolution role of natural selection in evolution. Meanwhile,
by natural selection. They both had experience of Darwin’s five-year voyage on HMS Beagle around
fieldwork in the tropics—an environment boasting the southern hemisphere as a gentleman naturalist
high biodiversity, competition for resources, and provided him with much material to formulate his COLLECTION BOX
marked differences in the organisms living in own theory of evolution. In 1858, Wallace and Both Darwin and Wallace were fascinated
separate areas. They both wondered how and why Darwin produced a joint publication on natural by insect diversity, especially as found in
such natural phenomena had arisen. On his travels the tropics, and were avid collectors.
selection, and the following year Darwin expanded
Wallace collected specimens for study and sale, and the theory to produce his famous and influential
it was in the Malay Archipelago that he formulated book, On the Origin of Species.
BIOGEOGRAPHY
The distribution of certain
reptile, therapsid, and plant
fossils across the southern
continents shows that they were
africa
Cynognathus Lystrosaurus once joined as a supercontinent,
therapsid fossils india therapsid fossils called Gondwana.
from the from the Triassic
Triassic Period Mesosaurus
reptile fossils from
the Permian Period
south america

australia
EARLY BIRD
The discovery of the fossil Archaeopteryx
Glossopteris in 1861 revealed characteristics that
plant fossils from antarctica provided an evolutionary link between
the Permian Period two major groups: reptiles and birds.
26 E VO LU T I ON
I N PRO G R E S S
LIVING EARTH • EVOLUTION IN PROGRESS

Darwin and Wallace proposed the theory of natural selection,

but it was the discovery of the gene that gave scientists the
mechanism by which selection takes place. Understanding the
gene has since become the key to understanding evolution.

NATURAL SELECTION
A key evolutionary mechanism, natural
selection favors the survival of the
fittest. In other words, individuals that
possess characteristics best adapted to
their current environment have a better
chance of surviving to reproduce and
pass on those favorable traits to
another generation. Natural genetic INDIVIDUAL VARIATION
variation within populations produces Litters of domestic cats commonly include
differences such as size, shape, and individuals with varied coloration, especially
where the parents’ colors differ.
color, and some of these may help
promote survival. For instance, a changes over time, a different coloration
particular body coloration may provide might be more beneficial, so natural
a camouflage that better hides the selection will ensure another change.
animal from predators than other A split into two populations may be
colors would. If this leads to the imposed by a physical barrier, with each
survival of that animal and to its new population becoming adapted to
reproduction, that same improved slightly different conditions. Eventually,
coloration will be passed on to some this may lead to one species becoming
of its offspring. As an environment two, in a process known as speciation.

SEXUAL DIMORPHISM
There are often marked differences
between males and females of the same SCIENCE
species. Male frigate birds have inflatable C R E AT I O N I S M : B E L I E F V E R S U S S C I E N C E
throat pouches for attracting the females.
Most of the world’s religions provide a theory of creation, which
gives an explanation for the formation of Earth and life. Many
of these creation stories originated long before scientific data
and theories were available to offer alternative explanations and
understanding. Some believers in the Western Judeo-Christian
tradition maintain that the complexity of “design” in many organisms
implies that there must have been a “designer”—God—behind their
creation, and for this reason they dispute the theory of evolution.

GENES AND INHERITANCE a forest fire wipes out large

populations of butterflies
GENES AND CHANCE
Sometimes individuals are eliminated at random,
Particular traits pass from parents to offspring through with the result that their genes are not passed on
the transmission of genetic material. Genes preserve, to the next generation.
encoded in their DNA, all the information necessary
for the replication of a cell’s structure and its
maintenance. Genes are therefore the basic units
of heredity. Individual chromosomes—the
threadlike part of a cell—hold thousands of genes
on long strands of DNA. During sexual reproduction,
the fusion of sperm and egg cells produces two by chance, the
complete sets of gene-bearing chromosomes: survivors are only the genes next generation a chance event
mostly yellow of the survivors has fewer purple leads to total loss of
one copy from the father and one from the mother. butterflies are passed on butterflies purple butterflies
 ADAPTING TO EXTREMES
The lesser flamingo has evolved to
occupy a very specialized niche: to feed
on algae in extremely alkaline African
lakes. In the absence of competitors, it
can thrive there in huge numbers.

ISLAND EVOLUTION ARTIFICIAL SELECTION

Isolated islands provide natural laboratories for unusually fast Over millennia, humans have domesticated many different kinds
evolution, with intense competition for limited resources leading of animals and plants, from dogs and cattle to fruit trees and cereal
to rapid speciation. In 1835, Darwin’s visit to the Galapagos crops. Before the discovery of the gene, this was achieved simply
Islands allowed him to collect many bird specimens, particularly by selectively interbreeding organisms bearing the desired
finches. He noted slight variations between the specimens from characteristics—such as the ability to run fast or produce more
island to island. He also heard about the differences between the succulent fruits—over many generations, until the selected traits
giant tortoises on separate islands, and subsequent visits to other became dominant. Today, biotechnology achieves the same result
Pacific islands made him wonder about the possibility of new much faster by directly manipulating genes, both to enhance
species evolving from a common ancestor. The ornithologist John beneficial traits and to remove problematic ones.
Gould was able to identify Darwin’s finches as a new group of
12 separate species, rather than just varieties of the same species.
This persuaded Darwin
that species could change
under certain conditions
such as island isolation.
Island wildlife continues
to be an important
research area for modern
evolutionary biologists.

FLIGHTLESS BIRDS GENETIC MODIFICATION CLONING

Many species of flightless birds, such Alteration to the genetic makeup Genetically identical individuals can
as kiwis, evolved on the islands of of organisms can remove undesired be created by the transfer of a nucleus—
New Zealand, which lacked serious characteristics and add more useful with its genetic information—from
predators until humans arrived. ones, such as resistance to disease. an adult cell into a host egg cell.
28 C L A S S I F I C AT I ON “DOG ROSE”
Known variously as the dog rose, wild
briar, witches’ briar, dogberry, and the
hip tree, this plant has only one
Global diversity is estimated to range from 10 million species to billions. Latin name, Rosa canina, which
Nearly 2 million have been described, with many more added each year. All identifies it for everyone everywhere.
LIVING EARTH • CL ASSIFICATION

species are named and classified using a system devised over 250 years ago.

For centuries people have studied the The science of morphology, along with
natural world. Initially they were limited to other criteria such as behavior and
what they could find locally and to reports modern genetics, forms the basis
from travelers, as it was impossible to of classification today. In 1758, the “PUMA”
The puma is also known
preserve and send specimens any distance. tenth edition of Systema Naturae was as the cougar or mountain
Later, as travel became easier, explorers published. It was written by the lion. Its Latin name,
were paid to collect plants and animals and Swedish botanist Carolus Linnaeus. Puma concolor, alludes
to its uniform color.
ship’s artists would draw them. By the early He and his friend Peter Artedi had decided
1600s, natural history collections in Europe to divide the natural world between them for identifying different organisms to replace
were substantial, and many specimens had and classify everything in it, fitting the 7,300 the arbitrary descriptions that existed
been described, but there was no formal described species into the same hierarchical previously. The new method put an end to
arrangement that made these specimens or framework. Although Artedi died before his confusion caused by the same name being
accounts of them easily accessible. book was finished, Linnaeus completed the given to several species, or to single species
The aim of the early taxonomists, or work and published it along with his own. being know by several names.
scientists who describe and classify species, Within a species, distinct subspecies
was simple—to organize living things so l at i n n a m e s can sometimes be recognized in different
that they reflected God’s plan of creation. All living things now have a unique Latin locations. In the 1800s, Elliot Coues and
Between 1660 and 1713, John Ray published name—such as Panthera leo for the lion— Walter Rothschild adopted a trinomial
works on plants, insects, birds, fishes, and which is made up of the genus name, starting Latin system to accommodate them.
mammals, forming his groups on the basis with a capital letter, and a descriptive species This convention for naming species and
of morphological (structural) similarities. name. Linnaeus devised this binomial method subspecies is still used today.

TRADITIONAL CLASSIFICATION

DOMAIN KINGDOM PHYLUM CLASS

Eukaryota Animalia Chordata Mammalia
The latest taxonomic unit to In recent years, the traditional A phylum (known as a division in Introduced by Carl Linnaeus,
be created is the domain. It is kingdoms of plants and animals plants) is a major subdivision of a a class contains one or more
based on whether organisms have been further subdivided. kingdom and is made up of one or orders. Class Mammalia includes
possess cells with a nucleus Kingdom Animalia now includes more classes that share certain only animals that are warm-
(Eukaryota: protists, plants, fungi, only multicellular organisms that features. Members of the phylum blooded, have fur, a single jaw
and animals) or not (Archaea and must eat other species in order Chordata have a notochord—the bone, and that suckle their
Bacteria). to survive. precursor of the backbone. young on milk.

CONTRIBUTING TO
CLASSIFICATION
Over the years many scientists have ANIMALS AND PLANTS ORDER FROM CHAOS
attempted to organize the natural
world, combining earlier ideas with Aristotle was the first person to classify John Ray classified organisms based on
new research, culminating in the living things, and he introduced the term their overall morphology rather than just
traditional system of classification genos (meaning race, stock, kin)—genus a part of it. In doing so he could establish
shown above and the use of bi- and in Latin. He separated animals into those relationships between species more easily
trinomial Latin names. Some scientists with blood and those without, not realizing and organize them into groups more
were particularly influential, making blood need not be red. This division is very effectively. He also divided the flowering
significant contributions to taxonomy.
close to the modern classification of plants into two major groups of orders—
vertebrates and invertebrates. monocotyledons and dicotyledons.

aristotle, 384 –322 bce john ray, 1627–1705

 CLASSIFYING INSECTS
About 20,000 species of butterfly
have now been described. In order to
identify them or recognize new species,
museums hold vast reference collections,
like this one, for comparative purposes.

ORDER FAMILY GENUS SPECIES

Carnivora Canidae Vulpes Vulpes vulpes
Orders are the next level down A subdivision of an order, a family A term first used by Aristotle The basic unit of taxonomy,
in Linnaeus’s hierarchy, and is made up of genera and the in Ancient Greece, a genus species are populations of
contain one or more families. species within them. The family identifies subdivisions of a family. similar animals that breed
The Carnivora have modified Canidae has 35 living species, all Vulpes is a genus within the only with one another.
cheek teeth (carnassials) and with nonretractile claws and two family Canidae. All foxes have Vulpes vulpes, known for
large canine teeth specialized fused wrist bones. All but one upright, triangular ears and a its bright red fur, breeds
for biting and shearing. species have long, bushy tails. long, narrow, pointed snout. only with other red foxes.

A N I M A L , V E G E TA B L E , O R M I N E R A L A NEW KINGDOM INTRODUCING THE ARCHAEA

Linnaeus divided the natural world Historically organisms were classed Recognized by Carl Woese and
into three kingdoms—animals, as being either animals or plants, George Fox in 1977, the Archaea
plants, and minerals. He then but in 1866 Ernst Haeckel argued are microscopic organisms that
devised a hierarchical system of that microscopic organisms formed live in very extreme environments.
classification based on class, order, a separate group, which he called Initially grouped with the Bacteria,
family, genus, and species, and Protoctista (now Protista). There their DNA turned out to be so
established the convention of giving were now three kingdoms of life: unique that a new three-domain
species binomial Latin names. Animalia, Plantae, and Protista. taxonomic system was introduced.

carl linnaeus, 1707–1778 ernst haeckel, 1834–1919 carl woese, born 1928
30 ANIMAL GENEALOGY SCIENCE
OUTGROUPS
In the 1950s, a revolutionary new way of classifying organisms was proposed. The first step in doing cladistic analysis
of a group is to choose a closely
Called phylogenetics, it allowed taxonomists to investigate evolutionary
LIVING EARTH • ANIMAL GENEALOGY

related but more primitive species (the

relationships between species by placing them in hierarchical groups called clades. outgroup) for comparison. This allows
derived characters to be distinguished
Phylogenetics, also known as cladistics, is based they are more closely related to each other than to from primitive ones. For example, to
on the work of entomologist Willi Hennig (1913– groups without flippers. Characters that are unique determine a phylogenetic tree of birds,
1976). He assumed that organisms with the same to one group are useful for recognition purposes a crocodile might be chosen as the
outgroup, as both birds and crocodiles
morphological characters must be more closely but say nothing about relationships. So cladistic
belong to the clade Archosauria.
related to each other than to those that lacked analysis is based entirely on the identification of
them. Therefore, these organisms must also share synapomorphic characters.
the same evolutionary history and have a more
recent ancestor in common. Like the traditional understanding ancestry
taxonomy of Linnaeus, this method of classifying The more derived characters organisms have in
organisms is hierarchical, but due to the volume common, the closer their relationship is assumed to
of data involved, computers are used to generate be. For instance, brothers and sisters look more alike
the family trees, known as cladograms. than other children—they may have the same eyes,
For a morphological feature to be useful in the same chin, and so on. This is because in terms
cladistic analysis, it must have altered in some way of common ancestry they have the same parents but
from the so-called “primitive” ancestral condition are only distantly related to other people. a primitive bird relative
to a “derived” one. For example, the legs and paws Cladistics is now most commonly based on
of most carnivores are considered primitive in the genetics—except in the study of fossils—and has CLOSE RELATIONS
cladogram on the opposite page, compared to the exposed some unexpected shared ancestry. For The giraffes and kudus are both even-toed
derived condition of flippers in the seals, fur seals example, a genetic cladogram surprisingly revealed ungulates, so are more closely related to
and sea lions, and walruses. This derived character, that the whale’s closest living relative is the land- each other than to the odd-toed zebras.
As fur-bearing mammals, all three are
called a synapomorphy, is useful because it is shared dwelling hippopotamus—a relationship that more closely related to each other than
between at least two taxonomic groups, suggesting Linnaeus would certainly not have expected. to the feathered birds around them.
LOOKING AT CLADOGRAMS MILKY DIET
All mammals possess
To do a cladistic analysis, the different groups of organisms are scored on mammary glands. Unique
31
a set of characters that are either primitive or derived. Their distribution to the class Mammalia, this
feature is synapomorphic
is not always as straightforward as is shown in the diagrams below. Often at this taxonomic level.

LIVING EARTH • ANIMAL GENEALOGY

the resulting cladogram can be constructed in a number of different ways To find additional familial
and taxonomists have to choose between them. To do this they adopt the relationships within the
class, characters that are
principle of parsimony—they choose the cladogram involving the least synapomorphic at familial
number of steps or character transformations to explain the observed level are used.
relationships between the groups.

CHARACTER CANID BEAR SEAL FUR SEAL WALRUS CHARACTER SET

& SEA LION Most modern cladograms are based on
genetics using DNA codes. The codes that
Feeding young on milk 1 1 1 1 1 were used to generate the cladogram below
Short tail 0 1 1 1 1 have been replaced with more familiar
morphological descriptions in this character
Forelimbs modified into flippers 0 0 1 1 1 set. One character, feeding young on milk,
Very flexible spine 0 0 1 1 1 is shared by all the groups shown; some
characters are found in only some of the
Hind limbs turn forward under body 0 0 0 1 1 groups; and one character, the presence
Presence of tusks 0 0 0 0 1 of tusks, is unique to the walrus.

CLADOGRAM KEY
In this cladogram the canids are considered the most primitive group, known 0 primitive character
as the outgroup, and the walrus is the most derived. All characters on the 1 derived character
cladogram are shared by the groups to the right of each number—for example,
a short tail is shared by the bears, seals, fur seals and sea lions, and the walrus.

FUR SEALS
CANIDS BEARS SEALS AND SEA LIONS WALRUS

1 2 3 4 5
two characters differentiate the
the outgroup does not seals, fur seals and sea lions, only the fur seals and sea lions, walrus is the most derived
share character 1 and walruses from the bears and the walruses share character 4 genus in this cladogram

1 2 3 4 5

SHORT TAIL FLIPPERS FLEXIBLE SPINE SUPPORTING LIMBS TUSKS

Bears, seals, fur seals and sea Among carnivores, the Character 3 (flexible spine) In both the fur seals and sea This is a unique character
lions, and the walrus all have modified limbs of seals, fur operates at the same level as lions and the walrus, the (autapomorphy) of the walrus
short tails (character 1). seals and sea lions, and the character 2, further supporting pelvic girdle can be rotated that reveals nothing about its
Canids, however, display the walrus are unique. Character the relationship indicated to allow the hind limbs to relationship to other mammal
primitive condition and have 2 (forelimbs modified into by possession of flippers. aid locomotion on land. This families. The same is true of
long, bushy tails. Character 1 flippers) is therefore The more synapomorphic suggests they share a more the character “feed young
is therefore a synapomorphic synapomorphic at this level, characters that appear at a recent common ancestor with on milk”—present in all the
character shared by all the suggesting these three groups particular level, the more each other than they do with groups, it provides no clues as
carnivore families shown are more closely related to convincing the suggested the seals, which are far less to how they interrelate, so it is
except the canids. one another than to the bears. relationship becomes. mobile when out of water. not plotted on the cladogram.
32 TREE OF LIFE r ea di ng th e tr ee
This diagram shows how life evolved from simple organisms,
such as the Archaea—which appeared about 3.4 billion years
Using a branching tree as a way of showing the diversity ago—to complex life forms, such as animals, which appeared
of life was first suggested by the German naturalist Peter about 650 million years ago. It also shows the diversity of the
LIVING EARTH • TREE OF LIFE

Pallas in 1766. Since then many such trees have been vertebrates (see pp.34–35), which have a disproportionate
constructed. Initially treelike, complete with bark and representation here. The circles indicate points where two
leaves, they later became more diagrammatic, and took or more groups of organisms have branched from a common
account of evolutionary theories. Modern, computer- ancestor at about the same time. Only extant species are shown,
generated trees of life present many different ideas of so extinct groups, such as the reptilian dinosaurs, are missing.
how living organisms are related.

The first person to produce a tree of life that

reflected the concept of evolution was Charles
Darwin. He sketched the first of his ten
evolutionary trees of life in 1837. It was a
simple branching diagram, which he developed LIFE
further before publishing it in the Origin of BEGINS ARCHAEA
Species in 1859. The lettered branches show how
he thought his theory might work—the more
branch points separating an organism from
its ancestor (numbered 1), the more different
the organism will be. In 1879, Ernst Haeckel BACTERIA
took the idea further, with a tree that showed
darwin’s first tree animals evolving from single-celled organisms.
Today, DNA and protein analyses as well as morphology are used to
construct evolutionary trees and establish the genetic relationships between
organisms. Vast data sets require computers to generate the trees, which
are continually refined as new species and information are discovered.
Traditionally, trees of life emphasised vertebrate groups and
relationships as those of the microscopic archaea and bacteria, and
protists (those eukaryotes not classified as plants, animals, or fungi)
were more problematic. However, modern genomic studies have
revealed an extraordinary diversification and complexity of relationships STRUCTURE OF LIFE
within the bacteria and within the early, or basal, eukaryote groups.
All forms of life belong to one of three domains: Archaea, Bacteria, and
Eukaryota. Archaea and bacteria are unicellular and lack a cell nucleus—
m a s s e x t i nc t ion s
they used to be classified together as Prokaryota. Eukaryotic organisms
Mapping all life forms that have ever existed on a tree is difficult as, over tend to be multicellular and each cell contains a nucleus, within which
time, more than 95 percent of all species have become extinct. A mass DNA is stored. This table shows the three domains and the four eukaryote
extinction occurs when a large number of species dies off at the same kingdoms. Despite appearances, the Archaea and Bacteria are the largest
groups—although only about 20,000 species have been described,
time. This has happened five times in the past. The best-known extinction, estimates exceed 4 million species. Among eukaryotes, the groups that
which wiped out the dinosaurs, occurred at the end of the Cretaceous make up the protists and invertebrates are far more numerous in terms
Period; it is thought to have been caused by a meteor impact combined of species than vertebrate groups.
with volcanic activity. As habitats are rapidly destroyed by human activity, ARCHAEA EUKARYOTES
it is likely that there will be another extinction event in the future. BACTERIA PROTISTS
PLANTS
EXTINCTION TIMELINE MOSSES
CARBONIFEROUS

LIVERWORTS
started 2.58 MYA
QUATERNARY
CRETACEOUS

HORNWORTS
ORDOVICIAN

PALEOGENE
CAMBRIAN

DEVONIAN

JURASSIC

LYCOPHYTES
NEOGENE
SILURIAN

PERMIAN

TRIASSIC

FERNS AND RELATIVES

CYCADS, GINKGOS, AND
800 GNETOPHYTES
CONIFERS
FLOWERING PLANTS
NUMBER OF FAMILIES

600
FUNGI
MUSHROOMS
400 mass SAC FUNGI
extinction LICHENS
200 event
100
ANIMALS
541 485 444 419 359 299 252 201 145 66 23 0
INVERTEBRATES
cyanobacteria CHORDATES
MILLIONS OF YEARS AGO (MYA)
 SAC FUNGI, YEASTS GLOMEROMYCETES CHYTRIDIOMYCETES NEOCALLIMASTIGOMYCETES

LICHENS

Lichens are classified

according to the fungal
half of the partnership

MUSHROOMS,
RUSTS, & SMUTS
BLASTOCLADIOMYCETES ZYGOMYCETES

FUNGI CHOANOFLAGELLATES
AMOEBOZOANS
ANIMALS
EUKARYOTES
OPISTHOKONTANS
PROTISTS
EXCAVATES

KINETOPLASTIDS EUDICOTS
HORNWORTS
EUGLENIDS

LYCOPHYTES
LIVERWORTS

MAGNOLIIDS

RHIZARIANS
FERNS &
RELATIVES
BROWN STONEWORTS
RADIOLARIANS & RELATIVES
ALGAE

PLANTS FLOWERING PLANTS

GREEN MOSSES
ALGAE
CYCADS, GINKGOS,
RED GNETOPHYTES
ALGAE
MONOCOTS
FORAMINIFERANS WATER
MOLDS HETEROKONTS

CONIFERS

CERCOZOANS ALVEOLATES
DIATOMS BASAL ANGIOSPERMS

APICOMPLEXANS DINOFLAGELLATES CILIATES

gymnochlora stellata
 HORSESHOE MILLIPEDES
APLACOPHORANS CHITONS GASTROPODS BIVALVES TUSKSHELLS SEA SPIDERS ARACHNIDS CRABS & CENTIPEDES

CEPHALOPODS

RIBBON
WORMS

ARTHROPODS
MOLLUSKS
SEGMENTED FLATWORMS WATER BEARS
WORMS

LOBE-FINNED
LAMPSHELLS FISHES

CARTILAGINOUS
FISHES
BRYOZOANS TUNICATES

ANIMALS CHORDATES VERTEBRATES

AMPHIBIANS
SPONGES ECHINODERMS LANCELETS RAY-FINNED REPTILES
JAWLESS FISHES
FISHES TURTLES &
TORTOISES

CNIDARIANS

SNAKES
TUATARA CROCODILIANS

CAECILIANS
LIZARDS &
AMPHISBAENIANS
BIRDS

TINAMOUS
FROGS
& TOADS

SALAMANDERS
& NEWTS RATITES
GAME BIRDS

WATERFOWL

WADERS,
CRANES & RAILS GULLS, & AUKS

NIGHTJARS HUMMINGBIRDS CUCKOOS TURACOS BUSTARDS MESITES SANDGROUSE PIGEONS GREBES FLAMINGOS KAGU & TROPICBIRDS
& SWIFTS SUNBITTERN
 AARDVARK
PRIMATES
INSECTS CRUSTACEANS
HYRAXES

TREE SHREWS
SENGIS

ELEPHANTS
VELVET
WORMS TENRECS &
ROUNDWORMS GOLDEN MOLES
COLUGOS

POUCHED DUGONG &

EGG-LAYING MAMMALS MANATEES
MAMMALS

MAMMALS PLACENTALS
RABBITS & PIKAS

ANTEATERS
& SLOTHS
RODENTS
IBISES & ARMADILLOS
STORKS RELATIVES HOATZIN

CORMORANTS
& GANNETS
PETRELS LOONS PENGUINS
BATS
HEDGEHOGS
& RELATIVES

KINGFISHERS HORNBILLS
& RELATIVES & HOOPOES

PANGOLINS

BIRDS OWLS MOUSEBIRDS CUCKOO- CARNIVORES

TROGONS WOODPECKERS
OF PREY ROLLER & TOUCANS

ODD-TOED
UNGULATES

EVEN-TOED
UNGULATES
PASSERINES

SERIEMAS FALCONS PARROTS &

COCKATOOS WHALES
Minera
Rocks,
Fossils
als,
Life on Earth is shaped by the rocks
that lie beneath our feet. Made of different
combinations of minerals, they have a
far-reaching influence on the landscape,
on vegetation, and on the soil. Preserved

&
within these rocks, fossils form a highly
detailed record of life in the distant past,
showing the path that evolution has followed
over hundreds of millions of years.

38 62 74
MINERALS ROCKS FOSSILS
The building blocks of Classified according to Most fossils preserve
rocks, minerals typically how they formed, rocks hard remains, such as
have a crystalline are constantly broken teeth and bones. They can
structure. Several down and reformed. also include traces such
thousand exist in Earth’s The oldest rocks date as footprints, and carbon
crust, but fewer than back 3.8 billion years, impressions in rock,
50 minerals are common when Earth’s crust which reveal the past
and widespread. first solidified. existence of living things.
38 MINERALS
Minerals are the materials from which rocks are made. More than 4,000 different
MINERALS, ROCKS, AND FOSSILS • MINERALS

minerals exist, each one with its own chemical composition, found naturally on
Earth. Many minerals are hard and crystalline, and, while some are extremely
abundant, others—including diamond—are very rare, and highly prized.
Minerals are of great economic significance. i den ti fy i ng m i n er a ls
They provide us with countless useful materials, With experience, many minerals can be
from metals to industrial catalysts, and they identified by their appearance alone. Important
also include objects of great intrinsic beauty, clues include their color, their luster—the way
particularly when cut and polished as gems. But light is reflected from their surfaces—and above
on an even broader scale, minerals are essential all, their habit, and/or crystal form. Crystals
for life itself. In soil and in water, soluble are grouped into six systems according to their
Copper often appears in a dendritic minerals release a steady stream of chemical symmetry (see panel, below). Minerals do not
form, branching like a tree. It is a nutrients that plants and other organisms need always occur as crystals but can take different
very important economic mineral.
to grow. Without them, the world’s ecosystems forms: they can be dendritic (branching) or
would no longer operate. botryoidal (like a bunch of grapes), for example.
Minerals are classified according to their Minerals also differ in density, or specific
chemistry. A few minerals, such as gold, silver, gravity (SG)—measured by comparing the
and sulfur, can exist in a native state, meaning weight of a mineral to that of an equal volume
they contain a pure chemical element, and of water—and in their hardness (H). On the
nothing else. All other minerals are chemical 10-point Mohs scale, which measures hardness,
compounds. Quartz, for example, contains talc is rated 1, whereas diamond, the hardest
Malachite can be botryoidal, with two elements—silicon and oxygen—which mineral, is rated 10. A fingernail (hardness 21⁄2),
small, rounded shapes, or massive,
with no definite shape. are bound together very tightly, giving quartz copper coin (31⁄2), and steel knife blade (51⁄2)
its exceptional hardness and strength. Under all make useful benchmarks for testing hardness.
the Strunz classification system, quartz Surprisingly, size is not a useful clue. Gypsum
(silicon dioxide) is classified as an oxide, as crystals, for example, are usually less than
are chalcedony and opal, but under the Dana ⅜ in (1 cm) long, but the largest specimens ever
system, used in this book, they are included discovered are the size of a two-story house.
with the silicates. The largest group of minerals,
the silicates make up about 75 percent of
Earth’s crust. Other common mineral groups
Crocoite, which is lead chromate, VOLCANIC MINERALS
frequently occurs as slender, elongated include sulfides, oxides, carbonates, arsenates, The ground at Dallol in Ethiopia’s Danakil Desert is pockmarked
prismatic crystals. and halides. by volcanic vents and covered in sulfur, a native element.

D E B AT E
MINERALS OR NOT? CRYSTAL SYSTEMS
Minerals are traditionally considered
Crystals in the cubic Hexagonal and Crystals in the
to be inorganic. Although some system are relatively trigonal systems tetragonal system
organic materials are used as gemstones common and easily are very similar to have three axes of
and for other decorative purposes, and recognized. Such each other, with four symmetry, all at
some of these have a similar chemical crystals have three axes of symmetry. right angles. In a
composition to minerals, they are not equal axes at right Their crystals are tall prism (left), the
angles; shapes include often six-sided prisms vertical axis is the
true minerals. Amber is the hardened the cube and the with pyramidal longest. Squat prisms
resin from trees; most amber is of eight-faced octahedron. tops (left). are also common.
Cretaceous, Palaeogene, and Neogene
age and can contain fossilized insects. Crystals in the Crystals in the Crystals in the
Jet, a soft black rock, is a type of monoclinic system orthorhombic triclinic system
impure coal, and became highly have three unequal system are similar have a low degree
fashionable for jewelry in Victorian axes of symmetry, to those in the of symmetry, as the
times. Shell, pearl, and coral (the use only two at right monoclinic system, three axes are unequal
angles. Tabular but all three axes are in length, with no
of which is not sustainable) are rich (flattened, left) and at right angles; the right angles between
in calcite and are used decoratively. prismatic habits usual habits are tabular them. Prismatic
are common. and prismatic (left). habits are common.
 NAT I V E E L E M E N TS
40 Of the many natural elements, only about 20 are found in
the native state; that is, uncombined with other elements.
They are divided into three groups. The metals rarely form
MINER ALS • NATIVE ELEMENTS

as distinct crystals, tend to have a high specific gravity, GRAPHITE

and are relatively soft. Semimetals, such as antimony and Hexagonal • h 1–2 • sg
arsenic, commonly occur as rounded masses. Nonmetals, 2.09–2.23
including sulfur and carbon, often form as crystals. A form of pure carbon
ANTIMONY common in metamorphic
Trigonal rocks, graphite is dark,
native
h 3–3 ½ • sg 6.69 soft, and greasy, and
copper This rare semimetal occurs makes ideal pencil leads.
in hydrothermal veins,
often with arsenic and
silver minerals. The silvery
gray masses are coated
white when oxidized.

COPPER
Cubic • h 2 ½–3 • sg 8.94
Native copper occurs mainly as
irregular masses or branching or
wirelike forms. It is most notably
associated with basaltic lava. A
good conductor, it is widely used
in the electrical industry.

dendritic habit
copper on
goethite
groundmass

distinct, isolated
diamond crystal

DIAMOND
Cubic • h 10 • sg 3.51
The hardest of all minerals,
diamond, a valuable form
of carbon, occurs in igneous
rock rocks called kimberlites, which
groundmass originate in deep volcanic pipes.
resinous luster
uneven surface

ARSENIC
Trigonal • h 3 ½ • sg 5.72–5.73
Highly poisonous arsenic usually forms as
pale gray, rounded masses in hydrothermal platinum nugget
veins. If heated, it smells of garlic.
PLATINUM
Cubic • h 4 – 4 ½ • sg 21.44
SULFUR A rare metal, native platinum forms as
Orthorhombic • h 1 ½–2 ½ • sg 2.07 scales, grains, and nuggets in igneous rocks
Native sulfur forms striking yellow crystals and and is found in alluvial sands. Its high
powdery crusts around volcanic vents. It is mined for melting point means it is useful in industry,
use in sulfuric acid, dyes, insecticides, and fertilizers. for example in aircraft spark plugs. platinum
 gold in
quartz SULFIDES CINNABAR
Trigonal
The sulfides are a large group of minerals, in h 2–2 ½ • sg 8–8.2 41
gold Red-colored mercury
nugget which sulfur is combined with one or more sulfide has been the
metals. Many sulfides have a high specific main source of mercury

MINERALS • SULFIDES
gravity and a metallic luster. They often form through the centuries.
as excellent crystals. Sulfides occur in many It occurs around
hot springs and
geological situations, but frequently in volcanic vents.
hydrothermal veins. The group includes the
majority of the economically important metal
ore minerals.

GOLD long, curved STIBNITE

Cubic • h 2 ½–3 • sg 19.3 crystals resemble Orthorhombic • h 2 • sg 4.63– 4.66
Prized for its color and blades or swords A sulfide of antimony, this dark gray mineral
malleability, gold forms in is the main ore of antimony. Large deposits
hydrothermal veins and often occur in China, Japan, and the W. US.
weathers out to be found
as nuggets in river sands.

bornite
crystals
COBALTITE indistinguishable
Orthorhombic • h 5 ½ • sg 6.33 crystals form
Cobaltite is an uncommon large masses
sulfide of arsenic and cobalt. It is
an important cobalt ore in, for
example, Sweden and Norway. BORNITE
Orthorhombic • h 3 • sg 5.08
This copper iron sulfide is a
coppery red color, tarnishing
IRON to shimmering purple and blue.
Cubic • h 4 ½ • sg 7.3–7.87 massive bornite It is an important copper ore.
Most native iron occurs at Earth’s core,
and near the surface it readily combines
with other elements.

BISMUTH
Trigonal
h 2–2 ½ • sg 9.7–9.83 GALENA
Native bismuth is Cubic • h 2 ½ • sg 7.58
relatively rare. Hardly Lead sulfide is one of the more
ever found as distinct abundant and widely distributed
crystals, it more often sulfide minerals. It is mined
has a granular or extensively as lead ore.
branching form.

mercury
globules in
rock cavities

common, massive
form of
chalcopyrite
MERCURY GREENOCKITE
Trigonal Hexagonal CHALCOPYRITE
h Liquid • sg 14.38 h 3–3 ½ • sg 4.82 Tetragonal
This is the only metal that is liquid at Named after Lord Greenock,
h 3 ½ – 4 • sg 4.35
normal temperatures. In liquid form, on whose Scottish property A sulfide of copper and
mercury appears as silvery globules. it was discovered in 1840, iron, chalcopyrite has a
this rare cadmium sulfide deep brassy yellow color. It
can be yellow, red, or orange. has significant value
as a copper ore.

usual, chalcopyrite crystals

crystalline
form of
sphalerite
ACANTHITE
Monoclinic
h 2–2 ½ • sg 7.22
A sulfide occurring
as dark, metallic, sometimes
SPHALERITE spiky crystals, acanthite is
SILVER Cubic • h 3 ½– 4 • sg 3.9– 4.1 the main ore of silver.
Cubic • h 2 ½–3 • sg 10.5 A sulfide of zinc with variable
Widely distributed but rarely found in iron content, sphalerite is
abundance, native silver occurs mainly as massive the most heavily mined
twisted wires, scales, and branching masses. sphalerite zinc-containing ore.
 SULFIDES MOLYBDENITE
Hexagonal
42 h 1–1 ½ • sg 4.62– 4.73
This molybdenum sulfide is
lead-gray in color. It has a greasy
feel, which is due to weak bonds
MINERALS • SULFIDES

within a layered atomic structure.

granite

ORPIMENT
Monoclinic
h 1 ½–2 • sg 3.49 REALGAR GLAUCODOT
Named from the Latin for “golden Monoclinic • h 1 ½–2 • sg 3.56 Orthorhombic • h 5 • sg 6.05
paint,” this arsenic sulfide occurs A bright orange-red sulfide of This sulfide of cobalt, iron, and arsenic thin hexagonal
as foliated, columnar masses arsenic, realgar was historically occurs as silver-white pyramidal crystals crystals in layers
around hot springs. used as a pigment. and brittle masses.

MARCASITE indigo-blue
Orthorhombic covellite
h 6–6 ½ • sg 4.89 slender prismatic
crystals
A sulfide of iron that is
lighter and more brittle than
pyrite, marcasite frequently
occurs as cockscombs and
spear-shaped “twins.”

COVELLITE
Hexagonal • h 1 ½–2 • sg 4.68
Although covellite is rare, it is a
widespread copper sulfide. Its shining
indigo-blue color makes it attractive
to mineral collectors.

HAUERITE
Cubic • h 4 • sg 3.46
Hauerite is a very rare sulfide
of manganese. The brown
octahedral crystals can form when
ARSENOPYRITE certain minerals are altered in
Monoclinic the caps of salt domes.
h 5 ½–6 • sg 6.07
Silvery-colored
arsenopyrite is a sulfide PENTLANDITE
of arsenic and iron. Cubic
With almost 50 percent STANNITE h 3 ½ – 4 • sg 4.6–5
arsenic content, it is a Tetragonal • h 4 • sg 4.3– 4.5 This nickel and iron
principal ore of arsenic Stannite is a sulfide of tin, sulfide is found in basic
which is poisonous copper, and iron mined for the igneous rocks. It is
to humans. tin. Its name comes from the an important source
Latin for tin, stannum. of nickel.

calcite
groundmass
PYRITE
Cubic
h 6–6 ½ • sg 4.8–5 MILLERITE
Nicknamed “fool’s gold” Trigonal
because of its light goldish h 3–3 ½ • sg 5.3–5.5
color, this iron sulfide is This sulfide of nickel occurs in
the most common of all limestones and ultramafic rocks.
sulfide minerals. It is sought after as a nickel ore.

PYRRHOTITE CHALCOCITE
Monoclinic • h 3 ½ – 4 ½ • sg 4.58– 4.65 Monoclinic • h 2 ½–3 • sg 5.5–5.8
An iron sulfide with variable iron content, Dark gray to black in color, this copper
pyrrhotite has a magnetism that increases sulfide has been mined for centuries.
as the iron content decreases. It is one of the most profitable copper ores.
 SU L F O SA LT S
Sulfosalts are a group of about 200 minerals,
43
structurally related to sulfides and with many
of the same properties. In these compounds

M IN ER A L S • SULFOSA LTS
sulfur is combined with a metallic element—
commonly silver, copper, lead, or iron—and a
semimetal, often antimony or arsenic. Sulfosalts
frequently occur in hydrothermal veins, usually
BISMUTHINITE
Orthorhombic • h 2 • sg 6.78 in small amounts.
This sulfide of bismuth is an important
ore. Much of the bismuth recovered is
used in medicines and cosmetics. PYRARGYRITE POLYBASITE BOULANGERITE
Trigonal Monoclinic Monoclinic
h 2 ½ • sg 5.85 h 21⁄2–3 • sg 6.1 h 2 ½–3 • sg 6.2
Also called ruby silver, Somewhat uncommon, A bluish-gray sulfide of lead
this sulfide of silver and polybasite is a sulfide of silver, and antimony, boulangerite
antimony is red-black, but copper, antimony, and arsenic. is one of the few sulfide
thin splinters appear It can yield worthwhile minerals to form fine,
deep ruby-red. quantities of silver locally. hairlike crystals.

striated prismatic
crystals
STEPHANITE
Orthorhombic
h 2–2 ½ • sg 6.26
Stephanite is an opaque
black sulfide of silver and
antimony. It can be an
important silver ore—for
example, in Nevada.

JAMESONITE
Monoclinic • h 2 ½ • sg 5.63
A sulfide of lead, iron, and antimony,
jamesonite’s dark gray crystals can be fine PROUSTITE
and hairlike or larger and prismatic. Trigonal • h 2–2 ½ • sg 5.55–5.64
This sulfide of silver and arsenic is also called
light ruby silver. The transparent crystals
are bright scarlet-red.
ZINKENITE
Hexagonal
h 3–3 ½ • sg 5.25–5.35 TETRAHEDRITE
Zinkenite is a sulfide of lead and Cubic
antimony. It occurs as steel-gray h 3– 4 ½ • sg 4.6–5.1
hairlike or needlelike crystals. This sulfide of copper, iron,
and antimony is named
after its tetrahedron-
shaped crystals
(crystals with four
triangular faces).

bright,
metallic
lustre TENNANTITE ENARGITE BOURNONITE
Cubic • h 3– 4 ½ • sg 4.59– 4.75 Orthorhombic • h 3 • sg 4.45 Orthorhombic • h 2 ½–3 • sg 5.83
radiating, Tennantite, a sulfide of copper, iron, A steel-gray colored sulfide of copper and Black or steel-gray bournonite is a
needlelike and arsenic, is dark gray or black. arsenic, enargite has a metallic luster. Crystals sulfide of lead, copper, and antimony.
crystals It is very similar to tetrahedrite. are usually small, tabular, or prismatic. Crystals are tabular to prismatic.
 OXIDES PEROVSKITE
Orthorhombic
44 Oxides are compounds of oxygen and h 5 ½ • sg 3.98–4.26
Discovered in Russia in
other elements. Many oxides are very 1839, this dark-colored
hard, some have a high specific gravity, oxide of calcium and titanium
MINER ALS • OXIDES

and a number are brightly colored and forms in igneous and

metamorphic rocks.
are sought after as gems. The group
includes the chief ores of iron, manganese,
aluminum, tin, and chromium. Oxides
can occur in hydrothermal veins, igneous
and metamorphic rocks, and also, because CUPRITE
they can be resistant to weathering and Cubic • h 3 ½ – 4 • sg 6.14
transport, in sands and gravels. Found in various shades of red, this
copper oxide forms near Earth’s surface
by the oxidation of copper minerals.
FRANKLINITE
Cubic
h 5 ½–6 • sg 5.07–5.22 ILMENITE
This black or brown zinc Trigonal
iron oxide is found in h 5–6 • sg 4.68–4.76
metamorphosed limestones, This iron titanium oxide is the principal ore of
notably those at Franklin, titanium, a high-strength, low-density metal
New Jersey. used in aircraft and rocket construction.

octahedral franklinite crystal

URANINITE striated
Cubic • h 5–6 • sg 10.63–10.95 crystal face
This highly radioactive, black- CASSITERITE
or brown-colored uranium oxide Tetragonal • h 6–7 • sg 6.98–7.01
is the main ore of uranium, which Almost the sole source of the world’s tin, this tin oxide
is used in nuclear reactors to is mainly found in hydrothermal veins and pegmatites
produce electricity, and and sometimes as small grains among river gravels.
in the construction of
nuclear weapons.

vitreous
luster

SAMARSKITE
Monoclinic
h 5–6 • sg 5–5.69
Minerals known as samarskite—a
radioactive oxide of various metals
including yttrium, iron, tantalum,
and niobium—occur in igneous
rocks and alluvial sands.

GAHNITE
Cubic • h 7 ½–8 • sg 4.62
A rare oxide of aluminum and
zinc found mainly in
metamorphic rocks, gahnite
can form dark green or blue
to black crystals.

CORUNDUM
Trigonal
h 9 • sg 3.98 – 4.1
Corundum is an aluminum
oxide, second only in
hardness to diamond. Ruby
(red) and sapphire (usually
blue) varieties are used
as gemstones.

CHROMITE
Cubic • h 5 ½ • sg 4.5 – 4.8
This iron chromium oxide is the only
important source of chromium, an element
used in making chrome and stainless steel.
 H Y DROX I DE S
Hydroxide minerals are compounds
45
of a metallic element and the hydroxyl
bright, metallic radical (OH). They are common

MINER ALS • HYDROXIDES

luster minerals and often form through
a chemical reaction between an
existing oxide and fluids rich in
HEMATITE
water, seeping through Earth’s
Trigonal • h 5–6 • sg 5.26 crust. Hydroxides tend to
Widespread and abundant, this occur in the altered parts
iron oxide is mined extensively of hydrothermal veins and
for iron. Different forms are
colored black, metallic gray, or in metamorphic rocks.
brownish red.
GIBBSITE
Monoclinic
h 2 ½–3 • sg 2.38 –2.42
Gibbsite is one of the STIBICONITE
essential aluminum Cubic • h 5 ½ –7 • sg 3.5 –5.5
hydroxides in the An uncommon hydroxide of antimony,
FERGUSONITE aluminum ore bauxite. stibiconite is white or yellowish brown,
Tetragonal It also occurs in and forms by the alteration of other
h 5 ½–6 ½ • sg 4.2–5.8 hydrothermal veins. antimony minerals, especially stibnite.
Fergusonite is the group
name for an oxide of
many metals, including
yttrium, lanthanum
niobium, and cerium.

prismatic
crystal

PYROLUSITE
Tetragonal
h 2–6 ½ LEPIDOCROCITE
sg 5.04–5.08 Orthorhombic • h 5
Pyrolusite is a
sg 4.05 –4.13
This relatively rare iron hydroxide
common manganese may occur with goethite. It is
oxide. It is the primary reddish brown and can form
ore of manganese, an irregular and fibrous shapes.
essential element in
steel production.
DIASPORE
Orthorhombic • h 6 ½–7 • sg 3.2 –3.5
Diaspore is an aluminum hydroxide which
is found in bauxite. It also occurs in marble
and altered igneous rocks.

massive romanÈchite
RUTILE
Tetragonal
h 6–6 ½ • sg 4.23 limonite
A source of titanium, this oxide ROMANÈCHITE
of titanium often forms impressive Monoclinic • h 5–6 • sg 6.45
displays of thin, translucent Dark and opaque, this oxide of
needles in quartz crystals. manganese with barium is usually
found in aggregates or in massive
forms. Crystals are rare.
botryoidal
goethite

GOETHITE
Orthorhombic • h 5–5 ½ • sg 4.27 – 4.29
Goethite is a common iron hydroxide. A
non-crystalline hydrated variety—limonite—
gives a yellow-brown color to soils and rocks.

BAUXITE
Amorphous mixture
h 1–3 • sg 2.3–2.7
CHRYSOBERYL ZINCITE The primary aluminum ore,
Orthorhombic • h 8 ½ • sg 3.75 Hexagonal • h 4 • sg 5.64–5.68 BRUCITE bauxite is not a single mineral,
Chrysoberyl is beryllium aluminum oxide. It Zincite is a rare oxide of zinc. Its Trigonal • h 2 ½–3 • sg 2.39 but rather an assemblage of
is a prized gemstone, known for its exceptional only significant deposit in the US Brucite is a magnesium hydroxide, and its color is white, aluminum hydroxides and
hardness and yellow-brown color. has been exhausted. gray, blue, or green. It occurs in metamorphic rocks. iron oxides.
 HALIDES
46 When metallic elements combine with halogen
elements, halides are formed. The halogens
are iodine, fluorine, chlorine, and bromine.
MINERALS • HALIDES

Halide minerals are commonly very soft and

of low specific gravity, often having crystals
classified in the cubic system. Many of these
minerals, such as halite and sylvite, form in
evaporite sequences by the drying out of
saline waters. Other halides—for example, yellow fluorite purple fluorite cubic
fluorite—occur in hydrothermal veins. crystal

FLUORITE
Cubic • h 4 • sg 3.18
This calcium fluoride
often forms transparent
to translucent crystals
of various colors. Large
quantities are used
for making
hydrofluoric acid.

SYLVITE
Cubic • h 2 • sg 1.99
Sylvite is a potassium chloride, a salt
similar to halite. It occurs with halite
in evaporite deposits. Sylvite is used
to make potash fertilizers.
granular
green carnallite
fluorite

CARNALLITE
Orthorhombic • h 2 ½ • sg 1.6
Carnallite is a hydrated chloride of
magnesium and potassium, formed by
the evaporation of saline water. It is
important in fertilizer manufacture.

vitreous
luster

transparent
cubic crystal

orange
halite

DIABOLEITE BOLEITE HALITE

Tetragonal • h 2 ½ • sg 5.42 Cubic • h 3–3 ½ • sg 5.05 Cubic • h 2 • sg 2.17
A lead copper chloride Deep blue boleite is a rare hydroxide Halite (common salt), a sodium
hydroxide with a light to dark of lead, silver, copper, and chlorine. chloride, occurs in extensive beds
blue color, diaboleite forms by It occurs where lead and copper formed by the evaporation of saline
the alteration of other minerals. deposits have been altered. water. It can be colored or colorless. halite crystals
 JARLITE
Monoclinic CA R BONAT E S
h 4 – 4 ½ • sg 3.78–3.93 Carbonate minerals are compounds of metallic
47
Usually white, jarlite is
found in igneous rocks. It or semimetallic elements combined with the
is a rare sodium strontium carbonate radical (CO3). More than 70 carbonate

MINER ALS • CARBONATES

magnesium aluminum minerals are known, but calcite, dolomite, and
fluoride hydroxide. siderite account for most of the carbonate
material in Earth’s crust. Carbonates usually
form as “good” crystals with regular shapes
chlorargyrite and no foreign substances enclosed within
crust them. Many carbonates are pale colored, but SMITHSONITE
some, such as rhodochrosite, smithsonite, and Trigonal
malachite, are brightly colored. h 4 – 4 ½ • sg 4.42– 4.44
A zinc carbonate found in the
upper oxidized zones of zinc
CHLORARGYRITE ore deposits, smithsonite is
Cubic • h 2 ½ • sg 5.55 also mined for zinc.
This silver chloride is typically
scaly, platelike, or, in masses,
resembling wax. It occurs where prismatic
silver deposits are altered. crystals

nailhead
spar

dark green,
tabular crystals
of atacamite
limestone
CALCITE groundmass
Trigonal
h 3 • sg 2.71 BARYTOCALCITE
Most calcite (a calcium carbonate), one of Monoclinic • h 4 • sg 3.66–3.71
the most abundant minerals, occurs in the This barium calcium carbonate is white
dogtooth rocks limestone and marble. It can also to yellowish, and often found in
spar form outstanding crystals. hydrothermal veins within limestone.

curved
crystal faces

DOLOMITE
Trigonal
h 3½ – 4
sg 2.84–2.86
Dolomite is a calcium
magnesium carbonate
widespread in altered
ATACAMITE limestones. Dolomite
Orthorhombic • h 3–3 ½ • sg 3.76 rock, formed exclusively
Green atacamite is a copper chloride of massive dolomite, is
hydroxide, occurring where copper deposits used as a building stone.
have been oxidized. It is a minor copper ore. TRONA
Monoclinic • h 2 ½ • sg 2.14
A hydrated sodium
carbonate, trona is gray,
yellowish, or brown. It forms
CALOMEL at Earth’s surface, especially
Tetragonal in saline desert environments.
h 1½–2 • sg 7.15
Rarely found, this mercury
chloride is a white to gray
or brown mineral. Its
color deepens when
exposured to light.

CRYOLITE WITHERITE MAGNESITE STRONTIANITE

Monoclinic Orthorhombic Trigonal • h 3 ½ – 4 ½ Orthorhombic • h 3 ½
h 2 ½ • sg 2.97 h 3–3 ½ • sg 4.29 sg 2.98–3.02 sg 3.74–3.78
A rare aluminum sodium This fairly uncommon A magnesium carbonate, magnesite This strontium carbonate is
fluoride, cryolite often has an carbonate of barium is usually usually occurs as white to brownish found in hydrothermal veins and
icelike appearance. It is found in white or gray, and occurs in dense masses. It is made into furnace limestones. The strontium is used
granitic pegmatites and granites. hydrothermal veins. bricks and magnesia cement. in sugar refining and fireworks.
 CA R BONAT E S
48 flos ferri
(“flowers of iron”)
aragonite

botryoidal siderite
ARAGONITE
Orthorhombic SIDERITE
h 3 ½ – 4 • sg 2.95 Trigonal
Aragonite is a calcium carbonate, h 3 ½– 4 ½ • sg 3.96
chemically identical to calcite, but A brown-colored carbonate of iron,
in a different crystal system and named from the Greek for iron, sideros,
much less common. siderite occurs in a variety of forms.
rhombohedral
siderite

short,
prismatic
crystals

PHOSGENITE ARTINITE
twinned Tetragonal • h 2–3 • sg 6.12–6.15 Monoclinic • h 2 ½ • sg 2.01–2.03
aragonite crystals
This rare lead carbonate chloride is A hydrated magnesium carbonate
formed close to the Earth’s surface by the hydroxide, artinite has a
reaction of lead-rich minerals with water. distinctive habit, with sprays
of white, needle-shaped crystals.
HYDROZINCITE It occurs in serpentine rocks.
Monoclinic
h 2–2 ½ • sg 3.5– 4
Hydrozincite, a zinc
carbonate hydroxide, is pale
gray, white, pink, or yellowish.
It fluoresces bluish white
under ultraviolet light.

patch of green
malachite
around margins

limonite matrix

AZURITE
Monoclinic • h 3 ½ – 4 • sg 3.77
Azurite is a hydrous copper carbonate. Its
rich blue color and frequent association botryoidal
with green malachite in hydrothermal habit
veins are distinctive features.

LEADHILLITE
Monoclinic • h 2 ½–3 • sg 6.55
This lead sulfate carbonate hydroxide
usually occurs as well-formed crystals in
the oxidized zones of lead deposits.
 BOR AT E S
cluster of Borates occur when metallic elements combine
49
twinned with the borate radical (BO3). There are more
crystals
than 100 borate minerals, the most common

MINER ALS • NITR ATES

cerussite
crystals
being borax, kernite, ulexite, and colemanite.
Borates tend to be pale colored, relatively soft,
and have low specific gravity. Many occur in
evaporite sequences where saline waters have
CERUSSITE dried out and minerals are then precipitated KERNITE
Orthorhombic • h 3–3 ½ • sg 6.53–6.57 among layers of sedimentary rocks. Monoclinic • h 2 ½ • sg 1.91
A carbonate of lead occurring where A colorless or white sodium
lead-bearing veins have been altered, cerussite borate hydrate, kernite has less
is the most common lead ore after galena. water than borax. The two
BORACITE minerals often occur together.
Orthorhombic
h 7–7 ½ • sg 2.91–3.1
rhombohedral Crystals of this magnesium
crystals borate chloride are pale
green or white, with a
vitreous luster and high
hardness. Boracite occurs
in salt deposits.

translucent,
prismatic
crystals

ANKERITE
Trigonal
h 3 ½ – 4 • sg 2.97
Ankerite is a carbonate of
calcium with lesser iron,
magnesium, and manganese.
It is sometimes found in
gold-bearing quartz veins.
COLEMANITE ULEXITE
Monoclinic • h 4 ½ • sg 2.42 Triclinic • h 2 ½ • sg 1.95
This hydrated calcium borate hydroxide forms when A hydrated sodium calcium borate
saline water evaporates. It was the main source of hydroxide, ulexite’s white, fibrous
boron until the discovery of kernite. crystals transmit light down their
length. It has uses similar to borax.

RHODOCHROSITE
Trigonal • h 3 ½ – 4 • sg 3.7
Fine crystals of this manganese carbonate,
in shades of rose pink, can be found in
South Africa, the US, and Peru. Banded
forms are used as gemstones.
aurichalcite
crystals
characteristic AURICHALCITE
green coloring Monoclinic • h 1–2 • sg 3.96
A blue- or green-colored zinc BORAX
copper carbonate hydroxide, Monoclinic • h 2–2½ • sg 1.71
aurichalcite forms in the A chalky-white sodium borate HOWLITE
oxidized zones of zinc and hydrate, borax has many Monoclinic • h 3 ½–6½ • sg 2.6
copper deposits. applications, including in Howlite is a calcium borosilicate
medicines, laundry detergents, hydroxide. It generally forms as
glasses, and textiles. chalky, rounded masses.
malachite on
chrysocolla
N I T R AT E S
Nitrates are a small group of
compounds where metallic
elements combine with the nitrate
radical (NO2). These minerals
associated
azurite are usually very soft and have
low specific gravity. Many
dissolve readily in water and they
MALACHITE only rarely occur as crystals. They
Monoclinic are generally confined to arid
h 3 ½ – 4 • sg 3.6–4.05 regions, forming coatings on the NITRATINE
This striking green copper carbonate Trigonal • h 1 ½–2 • sg 2.26
often occurs as botryoidal masses.
land surface that often cover wide This sodium nitrate typically occurs as crusts
It is used for ornamentation and botryoidal
areas. Commercially, nitrates can on the ground surface in arid regions, especially
as a source of copper. malachite be used as fertilizers or explosives. in Chile. It is white, gray, brown, or yellow.
 SU L FAT E S
50 Sulfates are composed of metals combined
with the sulfate radical (SO4). There are
about 200 sulfates and many are rare.
MINER ALS • SULFATES

Some, such as gypsum, form in evaporite

deposits where minerals are precipitated GYPSUM satin spar
from drying saline solutions. Others occur Monoclinic • h 2 • sg 2.31–2.32
A widespread mineral, gypsum, a hydrated
as weathering products, or as primary calcium sulfate, produces plaster of Paris
minerals in hydrothermal veins. Many are when heated and mixed with water. THENARDITE
economically important—barite is used Orthorhombic
to lubricate drills on oil rigs. h 2 ½–3 • sg 2.66
radiating crystalline A pale gray or brownish
gypsum
prismatic chalcanthite mineral, thenardite is a sodium
crystal ANGLESITE sulfate. It is found on lava flows
Orthorhombic and around salt lakes.
h 2 ½–3 • sg 6.32–6.39
galena This lead sulfate occurs
in a variety of colors and CHALCANTHITE
forms. It is an alteration Triclinic
product of galena, the h 2 ½ • sg 2.29
primary lead ore. Rich blue or green
chalcanthite is a hydrated
copper sulfate. It forms through
the oxidation of chalcopyrite
and other copper sulfates.

mass of needle-shaped
LINARITE brochantite crystals
radiating, hairlike Monoclinic
crystals h 2 ½ • sg 5.35
rock Bright blue linarite is
groundmass a hydrous copper lead
sulfate. It occurs in the
oxidation zones of
copper and lead ores.

GLAUBERITE
Monoclinic ALUNITE
h 2 ½–3 • sg 2.75–2.85 Trigonal
CYANOTRICHITE Glauberite is a sodium h 3 ½ – 4 • sg 2.6–2.9
Monoclinic • h 1–3 • sg 2.76 and calcium sulfate. A hydrous sulfate of potassium and
This hydrated copper aluminum sulfate is Colorless, gray, or aluminum, alunite may be found at
named after the Greek for “blue” and “hair,” yellowish, it forms where volcanic vents where rocks are
referring to its clusters of fine, blue crystals. saline water evaporates. altered by sulfur vapors.

C H ROM AT E S
red
crocoite Chromate minerals form when metallic elements
combine with the chromate radical (CrO4).
They are rare minerals—crocoite is the only
reasonably well-known chromate. Generally
brightly colored, they are sought after by
mineral collectors. Chromates often form
when hydrothermal veins are altered by fluids.
slender, elongate
crystals with
striations

orange
crocoite
CROCOITE
Monoclinic • h 2 ½–3 • sg 5.97–6.02
Orange or red lead chromate forms
in the oxidized zones of lead deposits.
Good specimens come from Australia.
 MELANTERITE
Monoclinic prismatic
h 2 • sg 1.89 crystals 51
White, green, or blue
melanterite is hydrated
iron sulfate. It is used for

MINER ALS • TUNGSTATES

purifying water supplies,
and as a fertilizer.

JAROSITE
Trigonal
h 2 ½–3 ½ • sg 2.9 –3.26
EPSOMITE
Orthorhombic Jarosite is a hydrous sulfate of iron and
h 2–2 ½ • sg 1.68 potassium, occurring as brown coatings
on pyrite and other iron minerals.
This hydrated magnesium
sulfate occurs in arid
regions and limestone cave
walls. It is the source of
the laxative Epsom salts.

CELESTINE
Orthorhombic • h 3–3 ½ • sg 3.96–3.98
This strontium sulfate is sought after not only
as the main source of strontium, but also for the
beautiful, transparent, pale-colored crystals.

COPIAPITE
Triclinic • h 2 ½–3 • sg 2.08–2.17
A yellow or green hydrated sulfate of
iron first described from Copiapó,
Chile, copiapite occurs where other
minerals are altered.
iron oxide
groundmass
BARITE
BROCHANTITE ANHYDRITE Orthorhombic
Monoclinic Orthorhombic h 3 • sg 4.5 POLYHALITE
h 3 ½ – 4 • sg 3.97 h 3–3 ½ • sg 2.98 The most common Triclinic • h 2 ½–3 ½ • sg 2.78
A copper sulfate A form of calcium sulfate, barium mineral, Polyhalite is a hydrated potassium
hydroxide, brochantite anhydrite occurs with gypsum but barium sulfate is calcium magnesium sulfate. Colorless,
forms emerald-green is less common. It alters to gypsum unusually heavy for a white, pink, or red, it is widespread
crystals, crusts, or masses. in humid conditions. pale-colored mineral. in many marine salt deposits.

M O LY B DAT E S T U NGSTAT E S HÜBNERITE quartz

Molybdates form when metals combine with Tungstate minerals are compounds with metallic Monoclinic hübnerite
h 4 – 4 ½ • sg 7.12–7.18 crystal
the molybdate radical (MoO4). These minerals elements joined to the tungstate radical (WO4). This manganese tungstate is a
are rare and tend to be dense and brightly These minerals are rare and usually have a high major source of tungsten,
colored. Molybdate minerals occur in mineral specific gravity. Some form as fine crystals. a metal used in steel alloys,
veins that have been altered by circulating water. Tungstates occur in hydrothermal veins and abrasives, and light bulbs.
Wulfenite is the best-known molybdate mineral. pegmatites—very coarse-grained granitic
It is prized for its fine crystals and brilliant rocks where minerals form from fluids
orange or yellow colors. permeating the rock.
SCHEELITE
Tetragonal
h 4 ½–5 • sg 6.1
thin, tabular Mined for tungsten, this
wulfenite calcium tungstate is found
crystal in hydrothermal veins,
metamorphic and igneous
rocks, and alluvial sands.

bipyramidal
scheelite crystal

FERBERITE
Monoclinic
h 4 – 4 ½ • sg 7.58
Opaque black
WULFENITE ferberite is found in
Tetragonal • h 2 ½–3 • sg 6.5–7.5 hydrothermal veins and
This lead molybdate is found in oxidized granitic pegmatites. It is
zones of lead and molybdenum deposits. an iron tungstate, mined
It is a minor source of molybdenum. as a source of tungsten.
 PHOSPH AT E S
52 When metals combine with the phosphate radical (PO4),
phosphate minerals are produced. These minerals form
a large group of more than 200—however, many are
MINER ALS • PHOSPHATES

very rare. Some minerals in this group are brightly

colored. They usually form by the alteration of sulfide TURQUOISE
minerals, but they are sometimes primary minerals. A Triclinic • h 5–6 • sg 2.6–2.8
number of phosphates are radioactive. A sought-after gem for thousands
of years, this hydrated phosphate of
copper and aluminum is found
HYDROXYLHERDERITE in altered igneous rocks.
Monoclinic • h 5–5 ½ • sg 2.95
Hydroxylherderite is a calcium
beryllium phosphate. It occurs
as pale yellow or greenish
crystals, with a glassy luster,
in granitic pegmatites.

PYROMORPHITE
aggregate Hexagonal • h 3 ½ – 4 • sg 7.04
of xenotime A lead phosphate chloride, variably
crystals colored greenish, orange, yellowish
or brownish, pyromorphite forms in
the oxidized zones of lead deposits.
prismatic
DUFRÉNITE apatite
Monoclinic crystal
h 3 ½ – 4 ½ • sg 3.10–3.34
This hydrated phosphate of iron
and calcium mainly occurs as green
to black masses or crusts in altered
veins and iron ores. XENOTIME-(Y)
Tetragonal
h 4 –5 • sg 4.4 –5.1
This widely distributed yttrium
phosphate is yellow-brown, gray,
or greenish, and forms in igneous
and metamorphic rocks. APATITE
Hexagonal
h 5 • sg 3.16–3.22
AUTUNITE Apatite is the group name used for
Orthorhombic three similar calcium phosphate
h 2–2 ½ • sg 3.05–3.2 minerals: fluorapatite, chlorapatite,
Radioactive autunite is a and hydroxylapatite.
lemon-yellow or pale
green hydrated phosphate
of calcium and uranium.
It occurs where uranium
minerals are altered.

tabular
torbernite crystal

TORBERNITE WAVELLITE
Tetragonal • h 2–2 ½ • sg 3.22 Orthorhombic • h 3 ½ – 4 • sg 2.36
This copper uranium phosphate hydrate Wavellite is a rare hydrated aluminum
is related to autunite and occurs in phosphate hydroxide. Colorless, gray, or
similar geological environments. It is greenish, glassy, needlelike crystals form
a radioactive mineral. radiating aggregates in altered rock.

sliced nodule

waxy luster

VARISCITE
Orthorhombic
h 3 ½ – 4 ½ • sg 2.56–2.61 TRIPLITE AMBLYGONITE
This hydrated aluminum Monoclinic • h 5–5 ½ • sg 3.5–3.9 Triclinic • h 5 ½–6 • sg 3.04–3.11
phosphate usually occurs as Triplite is a phosphate of manganese This rare lithium aluminum
green microcrystalline masses and fluorine, sometimes with iron. fluorophosphate mainly forms as masses,
in nodules, veins, or crusts. It forms in granitic pegmatites. but crystals occur in Zimbabwe and Brazil.
 VIVIANITE
Monoclinic • h 1 ½–2 • sg 2.67–2.69 VA N A DAT E S
Vivianite is a hydrated iron Vanadates are formed by the combination
53
phosphate. It commonly forms as
clusters of dark prismatic crystals of metallic elements and the vanadate
in altered iron deposits. radical (VO4). This group contains many

MINER ALS • ARSENATES

rare minerals, which may be dense and
brightly colored. Vanadates often form
when hydrothermal veins are altered by
radiating acicular permeating fluids. Most vanadates have
(needle-shaped)
crystals no commercial value; however, carnotite
is an important source of uranium.
powdery crust on sandstone
TYUYAMUNITE
CARNOTITE Orthorhombic
Monoclinic • h 2 • sg 4.75 h 1 ½–2 • sg 3.57–4.35
Generally occurring as powdery This rare hydrated vanadate of
yellow crusts in uranium calcium and uranium looks similar
libethenite deposits, radioactive carnotite to carnotite, and also occurs in
crystal is a hydrated vanadate of altered uranium deposits.
potassium and uranium.
LIBETHENITE
Orthorhombic
h 4 • sg 3.97
Libethenite is a light A R SE NAT E S
to dark green copper
phosphate hydroxide. Arsenates are mostly rare minerals composed of
It forms in the upper metallic elements and the arsenate radical (AsO3
oxidized zone of or AsO4). They generally have a low hardness.
copper deposits. ADAMITE
Many arsenates are brightly colored—adamite
Orthorhombic
is yellow or green, and clinoclase is green or blue. h 3 ½ • sg 4.32 – 4.48
This group of minerals form in a variety of This zinc arsenate hydroxide
MONAZITE geological situations, but many arsenates occur occurs in altered arsenic and
Monoclinic • h 5–5 ½ • sg 5–5.5 in altered metal deposits. zinc deposits, sometimes as
Phosphate minerals containing either exceptional crystals.
cerium, lanthanum, or neodymium are
all referred to as monazite. They are ERYTHRITE radiating clusters of
mined for the various elements. Monoclinic • h 1 ½–2 ½ • sg 3.06 clinoclase crystals
This hydrated cobalt arsenate forms
purple-pink crystals or coatings.
BRAZILIANITE Excellent examples occur in
Monoclinic • h 5 ½ • sg 2.98 Canada and Morocco.
First discovered in Brazil, this
sodium aluminum phosphate
hydroxide is yellow or
greenish and forms in cavities
in granitic pegmatites.

BAYLDONITE
Monoclinic
h 4 ½ • sg 5.24–5.65
This hydrous arsenate
of copper and lead is
usually found as green or
yellow crusts in altered
hydrothermal veins.
LAZULITE
Monoclinic
h 5 ½–6 • sg 3.12–3.24 olivenite OLIVENITE
A relatively rare, semiprecious crystals Monoclinic
blue gemstone, this magnesium h 3 • sg 4.46
bipyramidal aluminum phosphate hydroxide Olivenite is a copper
crystal occurs in metamorphic arsenate hydroxide. It
and igneous rocks. can be greenish, brownish, CLINOCLASE
yellow, or gray and occurs Monoclinic • h 2 ½–3 • sg 4.38
VANADINITE in altered copper deposits. Clinoclase is a dark blue-green copper
Hexagonal • h 2 ½–3 • sg 6.88 arsenate hydroxide that has a variety of
forms in altered copper sulfide deposits.
This relatively rare lead
vanadate chloride forms crystals quartz
in altered lead deposits. It is an
important source of vanadium,
used in steel alloys.

MIMETITE CHALCOPHYLLITE
Hexagonal • h 3 ½ – 4 • sg 7.24 Trigonal • h 2 • sg 2.67–2.69
Unusual, barrel-shaped crystals Bright blue-green chalcophyllite
characterize this lead arsenate chloride, is a hydrated copper aluminum
although other forms also exist. Mimetite arsenate sulfate. It forms in
occurs in altered lead deposits. oxidized copper deposits.
 SI L ICAT E S ANDRADITE
Cubic • h 6 ½–7 • sg 3.8–3.9
massive
dumortierite
54 Silicates are the most common and largest group Yellowish green, brown, or
black, andradite garnet is a
of minerals. The fundamental building blocks are calcium iron silicate. Cut gems
tetrahedra of silicon and oxygen (SiO4) together with are excellent at separating
MINER ALS • SILICATES

other elements. They are subdivided into six groups white light into colors.
based on the arrangement of the silica tetrahedra. Some
form as isolated tetrahedra (nesosilicates), others occur
in pairs (sorosilicates), and yet others have a three- DUMORTIERITE
dimensional network of tetrahedra (tectosilicates). Orthorhombic • h 7–8 • sg 3.21–3.41
Some silicates occur as chains of tetrahedra Dumortierite is a silicate of aluminum,
iron, and boron. It usually forms fibrous
(inosilicates), while others form as sheets aggregates of radiating crystals, but it
(phyllosilicates) or rings (cyclosilicates). EUCLASE can also be massive.
Monoclinic
N E SOSI L ICAT E S h 7 ½ • sg 2.99–3.1
Euclase is a beryllium
aluminum silicate hydroxide.
It may form white, colorless,
green, or blue prismatic,
striated crystals.

NORBERGITE
Orthorhombic
HUMITE h 6 – 6 ½ • sg 3.18 KYANITE
Orthorhombic • h 6 • sg 3.2–3.32 Norbergite mainly occurs as Triclinic • h 5 ½–7 • sg 3.53–3.67
A magnesium iron silicate fluorohydroxide, brownish yellow, white, or pink Kyanite is an aluminum silicate. Its
humite generally occurs as yellow to granular masses in metamorphic bladed crystals in schist and gneiss
orange granular masses in metamorphosed rocks. It is a magnesium form at high pressures in the Earth.
limestones and dolomites. silicate fluorohydroxide.

DATOLITE PYROPE
Monoclinic Cubic • h 7–7 ½ • sg 3.58
h 5–5 ½ • sg 2.96–3 Pyrope garnet is a dark red magnesium
Datolite is a hydrous calcium aluminum silicate. It forms at high pressures
boron silicate. Not very common, in metamorphic and some igneous rocks.
it is mainly found in veins or
cavities in igneous rocks. rhombic
crystal faces

ALMANDINE
Cubic • h 7–7 ½ • sg 4.32
The most common garnet,
pinkish red almandine is
an iron aluminum silicate.
It is widely used as a gem.

green coloring due vitreous luster

to vanadium

red coloring
due to iron

GROSSULAR
Cubic • h 6 ½–7 • sg 3.59
Grossular garnet is a silicate of
green calcium and aluminum that
grossular sometimes forms in marble. It
occurs in a wide range of colors. red grossular
 OLIVINE
Orthorhombic
pinkish brown
topaz
SOROSI L ICAT E S
h 7 • sg 3.27 – 4.39 55
Common in igneous rocks, nesosilicate
minerals varying in composition from magnesium
silicate to iron silicate are called olivine.

TOPAZ
Orthorhombic • h 8 • sg 3.4–3.6
Topaz is an aluminum silicate fluoride hydroxide.
Crystal size is usually small, but a giant crystal
weighing 596 lb (271 kg) was found in Brazil.
EPIDOTE
Monoclinic
h 6 • sg 3.38–3.49
Epidote is an abundant mineral.
Crystals of this hydrous calcium
aluminum iron silicate are prismatic
or tabular, green, and striated.
translucent
AXINITE
Triclinic
typical green color h 6 ½–7 • sg 3.25–3.28
Axinite is a hydrous
calcium iron manganese
wedge-shaped aluminum boron
crystal
silicate, with axe-head-
shaped crystals.

CHLORITOID
Monoclinic • h 6 ½ • sg 3.4–3.8
twinned TITANITE crystals Widespread in metamorphic and
titanite Monoclinic • h 5–5 ½ • sg 3.48–3.6 in matrix volcanic rocks, chloritoid is a dark green
crystals
Variable in color, titanite is a calcium or black hydrous aluminum silicate of
titanium silicate. It is excellent at dispersing iron, magnesium, and manganese.
light—even better than diamond.

ANDALUSITE prismatic
Orthorhombic crystal
h 6 ½–7 ½ • sg 3.13–3.21
Andalusite is an aluminum
silicate. It occurs mainly in
low-grade metamorphic rocks
as coarse prismatic crystals
with a square cross-section.

ZIRCON rounded
Tetragonal HEMIMORPHITE aggregates
h 7 ½ • sg 4.6 – 4.7 Orthorhombic
short, prismatic
Zircon, a zirconium silicate, h 4 ½–5 • sg 3.48
willemite crystal This hydrated silicate of zinc
is a gem used extensively
in jewelry. It is also the occurs in altered zinc deposits.
main source of the metal It is very variable in both its
zirconium, used in color and form.
nuclear reactors.

long, parallel, DANBURITE

fibrous crystals Orthorhombic
h 7–7 ½ • sg 2.93–3.02
Danburite is a calcium boron
silicate. Its variably colored
crystals resemble those of topaz,
but it can also be granular.

VESUVIANITE
WILLEMITE Tetragonal
Trigonal h 6 ½ • sg 3.32–3.43
h 5 ½ • sg 3.89 – 4.19 SILLIMANITE Vesuvianite, also known as
Willemite is a zinc silicate. White, green, Orthorhombic • h 6 ½–7 ½ • sg 3.23–3.27 idocrase, is a hydrous calcium
yellow, or reddish, and usually massive, Sillimanite is an aluminum silicate with long, slender sodium magnesium iron aluminum
it occurs in altered zinc deposits and crystals. Its composition is identical to that of andalusite, silicate, with fluorine. It occurs in
metamorphosed limestone. but it forms at higher temperatures and pressures. marble and igneous rocks.
 C YC LO SI L ICAT E S I NOSI L ICAT E S
ACTINOLITE
56 Monoclinic
BENITOITE h 5–6 • sg 3.03–3.24
Hexagonal Actinolite is a more iron-rich,
h 6–6 ½ • sg 3.64 –3.65 darker colored form of the
MINER ALS • SILICATES

This usually blue barium titanium amphibole tremolite. It is one

silicate occurs in serpentinite and of the asbestos minerals.
veins in schist. Gem-quality crystals
come from California.

six-sided crystal
TOURMALINE
Trigonal
h 7 • sg 2.9–3.1
Tourmaline is the name for
a group of 11 hydrous
boron silicate minerals with
the same crystal structure TREMOLITE
Monoclinic • h 5–6 • sg 2.99–3.03
but varying chemistry.
A widespread amphibole, this
hydrous silicate of calcium and
prismatic magnesium forms in metamorphic
crystal rocks. It has been used as asbestos.

aquamarine emerald
BERYL
Hexagonal
h 7 ½–8 • sg 2.63–2.92 vitreous luster
This beryllium aluminum
silicate is both a source of
beryllium and a gemstone.
Gem varieties include emerald
(green) and aquamarine
(greenish blue).

PECTOLITE NEPHRITE
Triclinic • h 4 ½–5 • sg 2.48 –2.9 Monoclinic • h 6 ½ • sg 2.99–3.24
This sodium calcium silicate hydroxide This very tough, cream to dark green
forms in cavities within basalt. It is common form of the amphiboles tremolite and
MORGANITE in Canada, the US, and England. actinolite is commonly known as jade.
Hexagonal
h 7 ½–8 • sg 2.63–2.92
Morganite is a pink variety of AEGIRITE
beryl, colored by additional Monoclinic • h 6 • sg 3.5–3.6
cesium or manganese. It This brown, green, or black
forms tabular crystals in SUGILITE pyroxene is a sodium iron silicate.
pegmatites. Hexagonal It forms in metamorphic and
h 6–6 ½ • sg 2.74–2.79 dark igneous rocks.
columnar, six-sided This rare silicate of
prismatic crystal potassium, sodium, iron,
and lithium occurs in
HELIODOR metamorphic rock.
Hexagonal HORNBLENDE
h 7 ½–8 • sg 2.63–2.92 Monoclinic • h 5–6 • sg 3–3.4
Named after the Greek for Common in igneous and
“sun,” heliodor is a yellow metamorphic rocks, this amphibole
variety of beryl. Fine examples mineral is a dark, hydrous silicate
come from Russia. of calcium, magnesium, iron,
and aluminum.

long,
prismatic
fibrous crystal
mass

RHODONITE
Triclinic
h 5 ½–6 ½ • sg 3.57–3.76
WOLLASTONITE Rose-red or pink rhodonite
Triclinic • h 4 ½–5 • sg 2.86–3.09 is a manganese calcium
rock This calcium silicate, found in marble and silicate, occurring as crystals,
groundmass other metamorphic rocks, is used in ceramics, masses, and grains. It is used
paints, and as a form of asbestos. in jewelry making.
 prismatic
slender diopside
prismatic crystal 57
crystal
quartz

MINER ALS • SILICATES

SPODUMENE DIOPSIDE
Monoclinic • h 6 ½–7 • sg 3.1–3.2 Monoclinic • h 5 ½–6 ½ • sg 3.22–3.38
richterite This pyroxene mineral is a lithium aluminum This pyroxene is a generally green-colored
crystal silicate. Some enormous crystals have been found; calcium magnesium silicate. It occurs in
the largest weighed almost 110 tons. metamorphic and igneous rocks.

PIGEONITE
Monoclinic
h 6 • sg 3.3–3.46
This brown to purplish
black uncommon
pyroxene is a magnesium
iron calcium silicate. It
RICHTERITE occurs in igneous rocks AUGITE ASTROPHYLLITE
Monoclinic and in meteorites. Monoclinic Triclinic • h 3 • sg 3.2–3.4
h 5–6 • sg 3.1 h 5 ½–6 • sg 3.19–3.56 This complex hydrous silicate
The amphibole richterite Augite is a common of potassium, sodium, iron,
is a hydrous silicate of pyroxene. This silicate and titanium, with fluorine,
sodium, calcium, and of calcium, magnesium, occurs in gneiss and cavities in
magnesium. It occurs in long prismatic and iron occurs in igneous igneous rocks.
metamorphosed limestones crystal and metamorphic rocks.
and igneous rocks.

jadeite

JADEITE
Monoclinic
h 6 • sg 3.25–3.35
One of two carvable materials
commonly referred to as
jade, this pyroxene mineral
is a sodium aluminum
iron silicate.

rock groundmass

long, striated
crystals

RIEBECKITE
Monoclinic • h 5–5½ • sg 3.26–3.44
This amphibole is a hydrous sodium
iron silicate, occurring in igneous
rocks. The variety crocidolite, or
“blue asbestos,” occurs in
metamorphosed ironstone.
 PH Y L LOSI L ICAT E S CLINOCHLORE
Monoclinic
58 h 2–2 ½ • sg 2.6–3.02
spherical mass This hydrous silicate of iron,
of radiating magnesium, and aluminum forms
crystals as green tabular crystals. It occurs
MINER ALS • SILICATES

in a variety of rock types.

tabular
crystal

radiating
crystal groups

PREHNITE OKENITE
Orthorhombic Triclinic • h 4 ½–5 • sg 2.28–2.33
h 6–6 ½ • sg 2.8 –2.95 This hydrated calcium silicate
Prehnite is a hydrous silicate has fibrous or bladelike crystals,
of calcium and aluminum. usually white or tinted blue or
It occurs in cavities in basalt. yellow. It occurs in basalt.
MUSCOVITE
prismatic PETALITE Monoclinic
crystal Monoclinic h 2 ½ • sg 2.77–2.88
h 6 ½ • sg 2.41–2.42 Muscovite, or white mica,
Petalite is a lithium is a potassium aluminum
aluminum silicate. aluminosilicate hydroxide, with
Crystals are usually fluorine. It is very common in
gray-white and in metamorphic rocks and granite.
aggregates. It is mined
for lithium.
CAVANSITE
Orthorhombic • h 3– 4 • sg 2.21–2.31
spherical crystal Cavansite is a hydrated calcium
aggregate vanadium silicate. It is blue
PHLOGOPITE or greenish blue, and
Monoclinic occurs in cavities
h 2–3 • sg 2.78–2.85 in basalt.
The colorless, yellow, or
brown mica phlogopite is
a potassium magnesium
aluminosilicate hydroxide.
tabular lepidolite
crystal

typical blue
coloring
SEPIOLITE
Orthorhombic • h 2 • sg 2–2.2 LEPIDOLITE
This pale-colored clay mineral, a Monoclinic • h 2 ½–3 ½ • sg 2.80–2.9
hydrous magnesium silicate, usually Lepidolite is the term used for mica minerals
occurs as earthy masses in altered rocks. that are potassium lithium aluminum
It is used for ornamental carving. aluminosilicate hydroxides, with fluorine.

prismatic crystal
T EC TOSI L ICAT E S CITRINE
Trigonal
h 7 • sg 2.65
Citrine is a yellow to
brownish variety of
quartz. It resembles
topaz and is often used
as a gemstone.

AMETHYST
MILKY QUARTZ Trigonal
SMOKY QUARTZ Trigonal h 7 • sg 2.65
Trigonal • h 7 • sg 2.65 ROSE QUARTZ h 7 • sg 2.65 Amethyst is a purple
Smoky quartz is a brown variety Trigonal • h 7 • sg 2.65 This very common, milky variety of quartz, prized
of quartz, a silicon dioxide. It Rose quartz is a prized, translucent pink white variety of quartz occurs since ancient times. It is
occurs in igneous rocks and variety of quartz. Crystals are very rare; in all types of rocks and in found in hydrothermal
hydrothermal veins. it usually has a massive habit. hydrothermal veins. veins and in lava cavities.
 slender crystal ZINNWALDITE
Monoclinic • h 3 ½– 4
sg 2.9–3.1 59
This brown, gray, or green mica is
potassium lithium iron aluminum
aluminosilicate hydroxide, with

MINER ALS • SILICATES

fluorine.

CHRYSOCOLLA
Orthorhombic
h 2 ½–3 ½ • sg 1.93 –2.4
This blue or greenish blue
hydrated silicate of copper
and aluminum forms in
altered copper deposits.
Crystals do not occur.

VERMICULITE
Monoclinic
h 1 ½ • sg 2.4–2.7
This green or yellow clay
mineral often occurs where
micas have been altered. It is a
hydrated silicate of magnesium,
iron, and aluminum.

GLAUCONITE
Monoclinic • h 2 • sg 2.4 –2.95
A mica, glauconite is a potassium
sodium magnesium aluminum iron
aluminosilicate hydroxide. It occurs
in marine sedimentary rocks.

tabular biotite
crystal

CHRYSOTILE BIOTITE
Monoclinic • h 2 ½ • sg 2.53 Monoclinic • h 2 ½–3 • sg 3.3
Chrysotile is a hydrous silicate of Biotite, or black mica, is a potassium
magnesium, forming fibrous, silky, white iron magnesium aluminosilicate
crystals in serpentinite rock. It is the most hydroxide, with fluorine. It is abundant
abundant of the asbestos minerals. in igneous and metamorphic rocks.

ALLOPHANE
Amorphous TALC
PYROPHYLLITE h 3 • sg 2.8 Triclinic
Triclinic A clay mineral, h 1 • sg 2.58–2.83
h 1–2 • sg 2.65–2.9 this aluminosilicate The softest mineral, white, gray, or
This aluminum silicate hydroxide, hydrate is an alteration greenish talc is magnesium silicate
of variable form and color, occurs product of feldspars hydroxide. Its many uses include
in low-grade metamorphic rocks. and other minerals. It toiletries, paint, and ceramics.
It has good insulating properties. forms crusty masses.

vitreous prismatic
luster crystals

AGATE
ROCK CRYSTAL Trigonal
Trigonal h 7 • sg 2.6
h 7 • sg 2.65 JASPER Forming in cavities in lavas,
Rock crystal is a transparent, Trigonal • h 7 • sg 2.6 white agate is a type of chalcedony.
colorless variety of quartz, which Jasper is a variety of chalcedony, or quartz It is characterized by
is widely used as an ornament or microcrystalline quartz, and is used for jewelry. It vein concentric color bands
a gemstone. is opaque, and usually colored red by impurities. caused by impurities.
 T EC TOSI L ICAT E S
60
alternating
bands of
different colors
MINER ALS • SILICATES

CARNELIAN BLOODSTONE
Trigonal Trigonal • h 7 • sg 2.7
h 7 • sg 2.7 Bloodstone is a variety of
Carnelian is a variety of chalcedony chalcedony colored dark green by
colored red to orange by iron oxide. The traces of iron silicates. Flecks of red
CHALCEDONY finest quality carnelian comes from India. jasper throughout resemble blood.
Trigonal
h 7 • sg 2.65 CHRYSOPRASE
Chalcedony is microcrystalline Trigonal • h 7 • sg 2.7
quartz, a silicon dioxide. Pure
chalcedony is white. It forms in veins Chrysoprase is a variety of
and cavities in many rock types. chalcedony containing nickel, which
imparts a green color. It is the most
valuable of the chalcedony varieties.
ONYX
Trigonal
h 7 • sg 2.7 precious
opal
Onyx is a parallel-banded gemstone
variety of chalcedony. It is not particularly
common; notable locations are India
and South America.

OPAL
Amorphous
h 5 ½–6 ½ • sg 1.9–2.3
Opal is a hydrated silicon dioxide,
occurring as nodules, encrustations,
or masses in most rock types.
Impurities impart a variety of
colors. Opal is treasured
as a gemstone.

streaks of
yellow opal ironstone matrix
CANCRINITE
Hexagonal
yellow cancrinite h 5–6 • sg 2.42–2.51
The feldspathoid
prismatic crystal cancrinite is a
variously colored
hydrated sodium
calcium aluminosilicate
carbonate sulfate.

MICROCLINE ANORTHITE
SCAPOLITE Triclinic Triclinic HEULANDITE
Tetragonal h 6–6 ½ • sg 2.54–2.57 h 6–6 ½ • sg 2.74 –2.76 Monoclinic
h 5 ½–6 • sg 2.5–2.78 A very common alkali This plagioclase h 3–3 ½ • sg 2.2
The name scapolite feldspar, this potassium feldspar is a calcium This zeolite is a hydrated
encompasses a series of aluminosilicate is usually aluminosilicate. It sodium calcium
complex sodium calcium white or pinkish. The forms pale-colored aluminosilicate. It is
silicates, occurring mainly green variety is crystals, grains, used in petroleum refining
in metamorphic rocks. called amazonstone. or masses. as a molecular sieve.
 SCOLECITE ANDESINE
Monoclinic Triclinic
h 5 –5 ½ • sg 2.25 –2.29 h 6–6 ½ • sg 2.66–2.68 61
This zeolite is a hydrated A plagioclase feldspar,
calcium aluminosilicate. andesine is a gray or
Usually colorless or white, white sodium calcium

MINER ALS • SILICATES

it is common in igneous aluminosilicate. It is
and metamorphic rocks. widespread in igneous rocks.

massive
sodalite NATROLITE
Orthorhombic
h 5–5 ½ • sg 2.2–2.26 HYALOPHANE
One of the most Monoclinic
widespread zeolites, h 6–6 ½ • sg 2.81
this hydrated sodium A relatively rare barium
aluminosilicate occurs in feldspar, this potassium
long, slender, cavities in basalts, and in barium aluminosilicate
needlelike hydrothermal veins. is colorless, white,
crystals yellow, or pink.
HARMOTOME
SODALITE STILBITE Monoclinic ANALCIME
Cubic Monoclinic h 4 –5 • sg 2.41–2.47 Triclinic
h 5 ½–6 • sg 2.27 –2.33 h 3 ½– 4 • sg 2.19 This widespread zeolite is h 5–5 ½ • sg 2.24–2.29
The feldspathoid mineral This widespread zeolite is a a pale hydrated barium This pale zeolite, a hydrated
sodalite is a sodium aluminum hydrated sodium calcium potassium calcium potassium sodium sodium aluminosilicate, occurs
silicate chloride. Rare crystals aluminosilicate. It forms sheaflike aluminosilicate. It occurs in igneous, metamorphic,
have been found in Canada. crystals in a variety of rock types. in hydrothermal veins and and some sedimentary rocks.
volcanic rocks.
lazurite crystal

calcite
ALBITE ANORTHOCLASE LAZURITE
Triclinic Triclinic Cubic
h 6–6 ½ • sg 2.6–2.65 h 6–6 ½ • sg 2.57–2.6 h 5–5 ½ • sg 2.38 –2.45
A plagioclase feldspar, albite An alkali feldspar, anorthoclase The intense blue feldspathoid
is a pale-colored sodium is a sodium potassium mineral lazurite is a sodium calcium
aluminosilicate. It is aluminosilicate occurring as aluminosilicate sulfate. It is the
an abundant mineral. prismatic or tabular crystals. main mineral in lapis lazuli gems.
short, prismatic
orthoclase crystal THOMSONITE
LAUMONTITE POLLUCITE Orthorhombic
Monoclinic Cubic h 5 –5 ½ • sg 2.23 –2.29
h 3½– 4 • sg 2.23–2.41 h 6 ½ • sg 2.9 A pale-colored zeolite,
A widespread zeolite, A rare zeolite, this complex thomsonite is a hydrated
laumontite is a hydrated calcium hydrated aluminosilicate of aluminosilicate of sodium
aluminosilicate. It occurs in cesium and sodium commonly and calcium, widespread
igneous, metamorphic, and contains other elements, such as in cavities in basalts.
sedimentary rocks. calcium. It is a source of cesium.
ORTHOCLASE
Monoclinic • h 6 • sg 2.55–2.63 CHABAZITE
Orthoclase, an alkali feldspar, is a Trigonal
potassium aluminosilicate. It is a major h 4 • sg 2.05 –2.2
component of many igneous and This common zeolite is a
metamorphic rocks. hydrated sodium calcium
aluminosilicate. Crystals
hairlike tufts of are colorless, white,
mesolite crystals yellow, or pink.

MESOLITE
Orthorhombic • h 5 • sg 2.26
This white or colorless zeolite
occurs in igneous and
metamorphic rocks. It is a
hydrated sodium calcium
aluminosilicate.

HAÜYNE
Cubic • h 5 ½–6 • sg 2.44–2.5
A feldspathoid mineral, haüyne is a pseudocubic
sodium potassium calcium aluminosilicate, rhombohedral
with sulfate and chlorine. It occurs mainly crystal
in silica-deficient volcanic rocks.
62 RO C K S
The rocks that form Earth’s crust are composed of different combinations of minerals. Thought of as
MINERALS, ROCKS, AND FOSSILS • ROCKS

the epitome of strength and solidity, they are actually in a state of constant change, with rock being
destroyed and reformed over immense periods of time. Rocks are classified into three main groups,
depending on how they form.
The world’s oldest known rocks, from Canada’s Sedimentary rocks are created on Earth’s surface.
Northwest Territories, have existed for about 4 Their key feature is their layers, or strata, built
billion years, but most rocks are much younger. up over long periods of time. Some sedimentary
The chalk cliffs facing the English Channel date rocks—such as sandstone and shale—form when
back to the Cretaceous Period, which ended 66 existing rocks are eroded, releasing particles that
million years ago (see timeline, below), and the are then washed or blown away, producing rock
European Alps are younger still. Even in the elsewhere. Others, such as rock salt and rock
Igneous rocks can be formed by the Grand Canyon, the oldest rocks date back 2 gypsum, are created when salt water evaporates,
cooling of magma deep underground, like billion years—less than half the lifetime of the leaving behind its dissolved minerals, known as
this granite, or by volcanic eruption.
planet as a whole. The reason for this is that Earth evaporites. Sedimentary rock can also have a
is tectonically active, with new rock being formed biological origin: chalk is formed from the
by the planet’s internal heat. At the same time, microscopic skeletons of marine organisms, and
existing rocks are broken down, in an endless coal is derived from the remains of plants,
cycle that started when Earth’s crust first solidified. compressed over millions of years.
Metamorphism takes place deep beneath
rock groups Earth’s surface, when rocks are altered by heat,
Geologists classify rocks into three groups— pressure, or both. For example, marble is created
Sedimentary rocks such as sandstone igneous, sedimentary, and metamorphic— when limestone is heated by lava or by magma.
are formed by the erosion of existing rocks
and the deposition of their fragments. reflecting the different ways in which they are Recrystallization of the limestone may remove
formed. Igneous rocks are created by lava from any bedding in the original rock, enabling some
volcanic eruptions on Earth’s surface and from marbles to be cut without splitting apart, a
the cooling of molten magma underground. The quality prized in sculpture. Metamorphism can
heat in lava and magma rises from the mantle take place over very wide areas. If temperature
beneath Earth’s crust. The commonest kind, a and pressure are sufficiently high, all rocks can
black volcanic rock called basalt, forms most of be metamorphosed and ultimately melted,
the sea floor. Igneous activity can also create forming magma and completing the rock cycle.
plutonic rock, which cools and solidifies beneath
Metamorphic rocks such as schist form GRAND CANYON, ARIZONA
when changes in pressure, temperature, or the surface, often in huge masses called batholiths. This view of the Grand Canyon shows the almost horizontal
both alter existing rocks. This is how most of the world’s granite originates. layering of sedimentary rocks and the effects of river erosion.

GEOLOGICAL TIMELINE
2.58 MYA

years ago
11,700
4,600

4,000

2,500

201
541

485

444

419

359

299

252

145

5.3
66

56

34

23

Hadean Archean Proterozoic Phanerozoic Eon

Era

Paleozoic Mesozoic Cenozoic

Carboniferous

Quaternary
Cretaceous
Ordovician
Cambrian

Paleogene
Devonian

Neogene

Period
Permian
Silurian

Triassic

Jurassic

Trilobites
were a group of
marine animals
that became extinct
at the end of the
Pleistocene

Geologists divide Earth’s history into periods of time. The transition

Oligocene
Paleocene

Holocene
Miocene
Pliocene

Epoch

Paleozoic Era.
Eocene

Dalmanites caudatus from one to the next is defined by a global event verified by fossil evidence.
was a common Eons begin with a major change, such as the evolution of multicellular
trilobite during the organisms that ushered in the Phanerozoic. Epochs are divided by events
Silurian Period. such as the last ice age, after which the Holocene begins. MYA: million years ago
 IGN EOU S ROC K S RHYOLITE
This fine-grained,
64 Rocks that solidify from a molten state are called igneous rocks and pale-colored lava
contains much quartz,
are broadly divided into extrusive rocks and intrusive rocks. Extrusive mica, and feldspar. It
rocks form from lava on Earth’s surface, while intrusive rocks form frequently includes
ROCKS • IGNEOUS ROCKS

underground, from magma. Lava and magma are rich in silica and visible phenocrysts
metallic elements. As they cool, minerals such as feldspars, micas, (larger crystals).
amphiboles, and pyroxenes form. Different combinations of these
minerals make up many of the igneous rocks.
vesicle
(gas bubble cavity)
BASALT VESICULAR BASALT
A dark, fine-grained Dominated by plagioclase
volcanic rock, basalt feldspars, pyroxenes, and
is the most common olivine, this lava is
rock forming the dark-colored, with
oceanic crust. numerous vesicles BANDED
(former gas cavities). RHYOLITE
Similar in composition
to granite, rhyolite also
has tiny crystals because
dark, fine-grained rock it forms from rapidly
cooled lava. Banding
shows the direction
of lava flow.
PAHOEHOE
Common in Hawaii,
pahoehoe is named after
the Hawaiian word hoe,
meaning “to swirl.”
This basaltic lava
is also known as
ropy lava.

PORPHYRITIC
AMYGDALOIDAL BASALT BASALT
Amygdaloidal means almond-shaped and This dark-colored
refers to the gas bubble cavities found in rock has large crystals,
basaltic lavas, which are typically infilled typically of olivine or
with secondary minerals such as plagioclase feldspars, set
zeolites, carbonates, and agate. in a fine-grained matrix.

PUMICE PÉLÉ’S HAIR

Formed from frothy lava, pumice Named after a Hawaiian goddess,
contains small crystals of feldspar in a and composed of numerous
very fine-grained matrix. It is of such slender, brown, glassy strands,
low density that it floats on water. Pélé’s hair is formed by lava
sprays being blown in the wind.

LITHIC TUFF
This rock contains
fragments of
previously formed
rock in a very fine-grained
IGNIMBRITE matrix. Usually pale, lithic
A very fine-grained, tuff is formed by violent
pale-colored volcanic volcanic eruptions.
tuff, ignimbrite often
shows banding caused
by the flow of lava
when molten.

SPINDLE BOMB BREADCRUST BOMB

Molten, low-viscosity basaltic lava blasted through the This variety of volcanic bomb is characterized
air can assume aerodynamic forms such as this one. by a cracked crust, due to continued expansion
These masses cool to form “bombs.” of the interior after the exterior has solidified.
 AGGLOMERATE ANDESITE
Composed of relatively large Named after the Andes
rock fragments set in a finer mountains, andesite is 65
matrix, agglomerate forms common in many of
after volcanic explosions. the world’s subduction-
PORPHYRITIC related volcanic
ANDESITE arcs—chains of islands
Commonly composed of and mountains formed
plagioclase feldspars, by tectonic plate
pyroxenes, and amphiboles, movement. It is
porphyritic andesite typically about
contains larger crystals set 60 percent silica.
in its fine-grained matrix.

PITCHSTONE
This dense volcanic DACITE AMYGDALOIDAL
A pale- to medium- ANDESITE
rock has variable composition
and color, and a pitchlike, colored rock with very Brown, gray, purple,
resinous luster. fine grains, dacite is or red, this is a
mainly made of plagioclase fine-grained volcanic
feldspars and quartz, with rock containing
PORPHYRITIC pyroxenes, biotite mica, infilled gas bubbles
TRACHYTE and hornblende. called amygdales.
This rock has a complex
mineralogy of alkali
feldspars, quartz, micas,
pyroxenes, and hornblende.
Its fine-grained matrix
contains larger crystals.

SPILITE
This fine-grained,
brownish volcanic
rock contains augite
and plagioclase
feldspars. It forms
by the alteration
of basalt lava in
contact with
sea water.
TRACHYTE
Trachytes are a group of fine-
grained volcanic rocks containing
alkali feldspars and dark-colored
mafic minerals such as biotite,
hornblende, and pyroxenes. curved fracture

OBSIDIAN
Typically very dark, obsidian
is formed by the rapid cooling of
highly viscous rhyolitic lava
before individual minerals have
had time to form crystals.
This gives it a glassy texture.
Obsidian has been used as a
cutting tool since ancient times.
pale patch of
devitrified glass
RHOMB PORPHYRY
This rock is characterized by large crystals of feldspars with a
SNOWFLAKE OBSIDIAN rhomb-shaped cross-section set in its dark, fine-grained matrix.
The “snowflakes” in this glassy, black Unlike all the extrusive rocks on this spread, rhomb porphyry is
volcanic rock, with a high silica content, an intrusive rock—formed from magma underground—like
are areas of devitrified (crystalline) glass. all the rocks on the following spread.
 IGN EOU S ROC K S
66
DIORITE
Composed of plagioclase feldspars,
with amphiboles and pyroxenes,
diorite contains little or no quartz.
It is a coarse-grained plutonic rock.

GRANODIORITE
This is probably the most
common intrusive igneous
rock in the continental
crust. It is over 65 percent
plagioclase feldspar.

pale plagioclase
feldspar

LARVIKITE
A form of syenite, larvikite
is blue-black in color
and composed of large
amounts of feldspar that
can have a shimmering
blue schiller effect.

NEPHELINE SYENITE
Coarse-grained, pale-colored,
and composed of feldspars, SYENITE LAMPROPHYRE
micas, and hornblende, this
rock also contains nepheline Gray or pinkish, syenite is a Lamprophyre’s matrix
but usually no quartz. plutonic rock that occurs in is medium-grained and
large intrusions. Coarse- studded with distinct
grained, it contains feldspars, crystals of micas and
micas, and hornblende, and amphiboles. It forms
little or no quartz. in dykes and sills.

GRANITE
Coarse-grained and of varying color,
light plagioclase granite contains more than ten percent
feldspar quartz. Attractive when polished, it is
often used on the facade of buildings.
GABBRO
This dark, plutonic rock black tourmaline
contains plagioclase
feldspars, pyroxenes, and
olivine. Coarse-grained,
it forms from the slow
cooling of basaltic
magma at depth.

OLIVINE GABBRO
Gabbro is a dark, coarse-grained
rock with much pyroxene and
plagioclase feldspar. Olivine occurs LAYERED GABBRO PORPHYRITIC MICROGRANITE
in significant amounts in this This type of gabbro shows banding Larger crystals are set into the matrix of this
variety of gabbro. caused by the settling of minerals of medium-grained rock, which is composed
different density in the magma. mainly of quartz, micas, and feldspars.

white microgranite
MICROGRANITE
DOLERITE BOJITE This medium-grained
Typically occurring in sills and A dark-colored rock, granite—often with a
dykes, dolerite is a dark, medium- bojite is a general term for porphyritic texture—
grained rock composed of hornblende-rich gabbro. occurs in sills and dykes,
plagioclase feldspars, pyroxenes, It is coarse-grained and which are sheets of
and iron oxides. forms from magma. intrusive igneous rock.
 black biotite crystal of
mica red garnet
67

gray quartz

ROCKS • IGNEOUS ROCKS

GARNET PERIDOTITE DUNITE
Peridotite has a composition similar to that Composed almost entirely of olivine,
of Earth’s upper mantle. This dense, this rock is dark green or brown,
greenish variety is made of dark minerals with a medium-grained texture. It
including garnets, olivine, and pyroxenes. usually includes a little chromite.

green olivine crystals

PERIDOTITE
A dark, dense rock composed
mainly of olivine and pyroxenes,
peridotite is coarse-grained and
forms slowly at great depth.

pink orthoclase
feldspar KIMBERLITE
Dark-colored and coarse-
grained, kimberlite has a very
low silica content. Of variable
composition, it is the world’s
main source of diamonds.

ADAMELLITE GRAPHIC GRANITE

This type of granite forms at depth This coarse-grained rock contains quartz
with crystals of quartz, micas, and and feldspars, whose intergrowth results
feldspars. One-third to two-thirds of in a graphic texture, vaguely resembling
the total feldspar content is plagioclase. runic writing. It also contains micas.

FELSITE
dark Fine-grained felsite forms in
hornblende sheetlike intrusions, sills, and
dykes. Pale in color, it is
composed mainly of feldspars
and quartz.

PEGMATITE
Pegmatites are very PORPHYRITIC HORNBLENDE
coarse-grained rocks, GRANITE GRANITE
formed from residual liquid A pale-colored rock Granite typically contains ANORTHOSITE
magma after most of a composed of feldspars, quartz, feldspars, and Pale-colored, anorthosite is
granite intrusion has cooled quartz, and micas, micas. This variety also composed mainly of large
and crystallized. Some porphyritic granite has contains hornblende, a crystals of plagioclase
pegmatites are an important large, well-formed crystals member of the amphibole feldspars. It can also
source of gemstones. set into the matrix. mineral group. contain olivine and augite.
 M E TA MOR PH IC RO C K S
68 When existing rocks are subjected to heat or pressure or both
within the Earth, they are changed into different assemblages of
minerals. Contact metamorphism occurs when intense, localized
ROCKS • META MOR PHIC ROCKS

heat radiating from a body of igneous rock causes the surrounding

rocks to recrystallize. Regional metamorphism takes place over
wide areas, often at considerable depth, as a result of intense
heat and pressure. Movements of the Earth’s crust can result HALLEFLINTA
in rocks being pulverized by dynamic metamorphism. Originally volcanic tuff, rhyolite,
GRANULITE SCHIST or quartz porphyry, halleflinta
streaked Formed at very high Schist is characterized by is fine-grained, pale-colored,
texture temperatures and pressures, its parallel planes of similarly and rich in quartz. It is
granulite is dark, coarse- oriented minerals within a type of hornfels.
grained, and rich in pyroxenes, the rock. The corrugated
garnets, micas, and feldspars. pattern seen on the specimen
is called crenulation.

MYLONITE SPOTTED SLATE

When rocks are pulverized deep within a thrust fault This is a dark, fine-grained
zone, the rock dust and fragments produced form a rock characterized by black
fine-grained rock called mylonite. spots (porphyroblasts) of
minerals, such as cordierite
and andalusite.

KYANITE SCHIST
Mainly composed of feldspars,
MUSCOVITE SCHIST micas, and quartz, this schist
also contains crystals of the
A typical schist with pale, glittery blue mineral kyanite.
muscovite mica, this rock also
contains quartz and feldspars.

GARNET SCHIST BIOTITE SCHIST

SLATE
The red garnets in this variety of Formed at relatively high pressures
schist indicate that it developed and temperatures, biotite schist Dark and very fine-grained, slate
at relatively high temperatures contains feldspars, quartz, and is a compact rock with parallel cleavage
and pressures deep within the much dark-colored biotite mica. planes. It is formed by low-pressure
continental crust. metamorphism.

pink
calcite
tubular structure

PHYLLITE
Phyllite is formed at lower temperatures
and pressures than schist, but higher than
slate. It is fine-grained and splits into
slabs with a characteristic sheen.

SKARN
Formed when rocks rich in carbonate
minerals are altered by high-
temperature contact metamorphism,
skarn contains minerals rich in calcium,
magnesium, and iron.

FULGURITE METAQUARTZITE
When lightning strikes in deserts or on With a high percentage of quartz, this rock is
beaches, sand can melt, forming small, harder than most metamorphic rocks. It is formed
tube-shaped structures, called sand fulgurites. from sandstone altered at high temperature.
 bladed crystal 69

ROCKS • META MOR PHIC ROCKS

CORDIERITE HORNFELS
Dark and splintery, cordierite
hornfels is produced by the heat
of nearby igneous intrusions. It GNEISS AUGEN GNEISS
is fine- to medium-grained. A coarse-grained rock formed at very Gneiss contains quartz,
high temperatures and pressures, feldspars, and micas,
gneiss is recognized by its alternating often in parallel bands.
dark and light crystalline bands. Augen gneiss has crystals
GARNET HORNFELS resembling lens-shaped
Hornfels is a tough, dark, eyes (augen in German).
flinty rock formed by the heat
from an igneous intrusion.
Reddish garnets occur in
this particular variety.
CHIASTOLITE HORNFELS
Hornfels forms at very high PYROXENE HORNFELS
temperatures near magma intrusions. Fine- to medium-grained,
Bladed crystals of pale chiastolite give tough, and flinty, this variety
this particular variety its name. of hornfels contains quartz,
micas, and pyroxenes. It forms
close to igneous intrusions.
FOLDED GNEISS
Gneiss becomes plastic and folded at great
AMPHIBOLITE depth. The dark bands are hornblende-rich
Formed by moderate heat and and the paler ones are of quartz and feldspars.
varying pressure deep in the
crust, this coarse-grained rock ECLOGITE
has an abundance of hornblende Composed of two main minerals, green
and plagioclase feldspar, as omphacite pyroxene and red garnet,
well as other minerals. eclogite is coarse-grained and forms at
very high temperatures and pressures.

marble
fragment

marble MIGMATITE GRANULAR GNEISS

breccia Formed at the highest temperatures This gneiss has a granular texture
and pressures, coarse-grained migmatite with equal-sized grains. It has dark
contains bands of dark basaltic minerals and bands of hornblende and biotite mica,
pale granitic minerals. It is often folded. and pale bands of quartz and feldspars.

gray marble
MARBLE
Formed by contact or regional
metamorphism, marble is rich in
calcite, often with colorful veins of other
minerals. It is prized as a carving material.

SERPENTINITE
Commonly banded, speckled, or streaked,
serpentinite is a dense, but soft, metamorphic
rock derived from peridotite. It is found in
green marble convergence zones between tectonic plates.
 S E D I M E N TA RY RO C K S
70 Sedimentary rocks are generally characterized by
stratification, or bedding, and they can contain fossils.
These rocks are divided into three groups according
ROCKS • SEDIMENTARY ROCKS

to their origin. Clastic rocks are made up of fragments

of rocks and minerals eroded from previously formed
rocks; organic rocks are derived from the remains
of plants and animals; chemical rocks result from
the precipitation of chemicals. MILLET-SEED SANDSTONE GREENSAND
The medium-sized grains of this rock are Colored greenish by the silicate
often coated with reddish iron oxides. mineral glauconite, greensand is
The well-rounded, equal-sized quartz a quartz-rich sandstone formed
grains have been shaped by the wind. in the sea.
MICACEOUS
SANDSTONE
Rich in quartz, LIMONITIC SANDSTONE
this sandstone also This rock is colored red-brown
contains glittery flakes or yellowish by the iron oxide
of mica. It usually has mineral limonite, which coats its
medium-sized grains. medium quartz grains.

iron oxide
gives red TRAVERTINE
color A pale-colored and often layered rock,
travertine is virtually pure calcite. It is
formed around hot springs and volcanic vents.

sandstone

SANDSTONE
ROCK SALT ROCK GYPSUM This rock typically occurs as layers of sand
Composed of halite, rock salt is often A crystalline rock formed when grains, held together by various mineral
colored by inclusions of iron oxides and salt water evaporates, rock cements that impart different colors.
clay minerals. It is soluble in water, with a gypsum is pale colored, often Sandstones are quartz-rich.
low hardness and a salty taste. fibrous, and very soft. It occurs
with other evaporite minerals.
red sandstone

BOULDER CLAY CLAYSTONE

Gray to brownish in color, boulder clay, or till, Of varying color, this very fine-grained
has a very fine-grained clay matrix with glacially rock is composed mainly of silicate clay quartz grains colored
derived angular and rounded rock fragments. minerals such as kaolinite—mostly by iron oxides
derived from the weathering of feldspars.

bands of hematite and chert

OOLITIC IRONSTONE
This rock is composed of
small, rounded grains (ooliths)
of iron minerals, such as
siderite, cemented by other
iron minerals, as well as
by calcite and quartz.

LOESS
A clay with very fine,
BANDED IRONSTONE dustlike grains lifted by
This rock, deposited in marine or fresh water, the wind from dry land
has alternating bands of black hematite and surfaces, loess is crumbly
red chert. It is one of the best ores of iron. and lacks obvious layering.
 71

ROCKS • SEDIMENTARY ROCKS

gray
orthoquartzite
TUFA
This porous rock is formed by ORTHOQUARTZITE MANGANESE NODULE
the precipitation of carbonate Rarely containing fossils, this rock Rich in valuable metals such as
minerals from lower ambient is composed almost entirely of silica- pink manganese, iron, nickel, cobalt, and
temperature bodies of water cemented quartz grains. Orthoquartzite orthoquartzite copper, these nodules are rounded,
than travertine, including is sometimes known as quartz arenite. black concretions that form on the
those in cave systems. deep ocean floor.
layered structure rounded
nodule

lignite jet
LIGNITE
Lignite is brown coal with a lower carbon content
than bituminous coal. Jet is a black, lustrous form
of lignite, and will take a high polish.

ANTHRACITE
The purest form of coal, which is derived from
plant material, anthracite is black, with a vitreous
luster. It has curved edges when broken.

BITUMINOUS COAL
This sedimentary
rock has a lower carbon
content than anthracite.
Brittle and dull, it is the
most abundant type of coal.

internal
radiating
structure

pale calcite
filling cracks
(septa)

SEPTARIAN
NODULE
This type of concretion occurs
in sedimentary rocks as individual, PYRITE NODULE
rounded masses. Internally, These spherical nodules occur in shale
cracks known as septa are infilled and clay, and are composed entirely
by quartz or calcite. of radiating crystals of pyrite.
 S E D I M E N TA RY RO C K S pisolith cemented
by calcite
72 fossilized crinoid,
or sea lily, stalk

CRINOIDAL
ROCKS • SEDIMENTARY ROCKS

LIMESTONE
Crinoids are echinoderms
that are attached to the
seabed by a flexible stalk.
Crinoidal limestone is a mass
of broken stalks cemented
by hardened lime mud.

FRESHWATER LIMESTONE
This limestone is a pale, calcite-rich
NUMMULITIC rock with some quartz and clay.
LIMESTONE It contains fossils of freshwater
A marine foraminiferan, organisms, which indicate the
Nummulites is the main fossil environment of deposition.
in this rock. The cement is PISOLITIC LIMESTONE
calcite, originally lime mud.
This rock is composed of pisoliths—
pea-sized grains slightly larger than
CORAL LIMESTONE ooliths, often flattened, and loosely
cemented by calcite.
This rock is a
mass of fossilized
corals cemented by OOLITIC LIMESTONE
fine-grained calcite. This limestone is composed of
It is gray to white or ooliths—small, rounded,
brownish in color. concentrically banded grains
rolled by seabed currents and
cemented by carbonate mud.

BRYOZOAN LIMESTONE
LIMESTONE BRECCIA
This is a gray or Often forming below
reddish organic limestone cliffs, this
limestone, which breccia comprises large,
contains fossils angular fragments
of bryozoans in a of limestone and other
matrix of hardened, rocks, cemented
calcite-rich mud. by calcite.
FELDSPATHIC
GRITSTONE
Coarse-grained and pale- to
dark-colored, this gritstone
contains much quartz and up
to 25 percent feldspar.

QUARTZ GRITSTONE
This gritstone is composed
of quartz with some
feldspar and mica, all
of coarse grain size.
DOLOMITE
This yellowish-brown or gray brachiopod fossil
rock contains a high percentage
ARKOSE of dolomite (calcium magnesium
GRAYWACKE Variable in color and carbonate). It is also called
This dark rock contains quartz, rock medium-grained, arkose is dolostone, to distinguish it
fragments, and feldspars, set in a mass of finer a sandstone with a high from this mineral.
clay and chlorite. It forms in marine basins. percentage of feldspar.
FOSSILIFEROUS
SHALE
Fine-grained marine
sedimentary rocks such as shale
often contain large numbers
of well-preserved fossils.

SHALE
This fine-grained, layered rock
varies in composition, usually
containing silt, clay minerals,
organic materials, iron oxides,
and minute crystals of minerals
such as pyrite and gypsum.
 POLYGENETIC
CONGLOMERATE rounded
quartz pebble
A coarse-grained sedimentary rock,
polygenetic conglomerate has many
73
different, rounded rock and mineral
fragments in a finer grained matrix.

ROCKS • SEDIMENTARY ROCKS

sandstone
matrix

iron oxide QUARTZ

gives red color CONGLOMERATE
Varying in color, this rock typically
has dirty white, pebble-sized,
rounded quartz fragments set
red in a finer, darker matrix.
chalk

BRECCIA
Containing large,
angular fragments
of rocks and minerals
set in a finer matrix
of sand or silt, breccia
rarely forms in layers.
CHALK
Pure calcite, chalk is very fine-
grained and powdery. It is composed SILTSTONE
of minute fossilized organisms, This dark-colored rock comprises particles,
including coccoliths and radiolarians. white chalk mainly of quartz, which are smaller
than fine sand but larger than clay
grains. It can also contain organic
MARL material and calcite. CHERT
Intermediate between clay A very fine-grained form
and limestone, marl is a of silica, chert occurs as
fine-grained, calcite-rich, layered bands and nodules in rocks
rock. Chlorite and glauconite can such as limestone. It is
give it a green coloring. usually gray in color.

MUDSTONE FLINT
Composed of clay, with Usually found as nodules
fine-grained quartz and in chalk, flint is very hard,
feldspars, mudstone lacks black, compact silica. It breaks
the bedding found in shale. leaving sharp, curved edges.
74 FOSSILS
Fossils are evidence of past life that has been buried and preserved
MINERALS, ROCKS, AND FOSSILS • FOSSILS

in the rocks of the Earth’s crust. They give scientists important clues
about how life has evolved; they can also be used to date rocks,
and create a timeline of the events that have shaped our world.
Life has existed on Earth for at least 3.7 billion minerals, which literally turns them to stone.
years. Initially, living things were small and Once formed, many fossils are destroyed by heat,
soft-bodied, and left few obvious traces of their pressure, or geological movements while still
existence. But during the past billion years, deep underground. But if a fossil survives all
life gradually changed. Organisms evolved hard this, uplift may eventually bring it back to the
body parts, which—given enough time—could surface, where erosion can release it from its
fossilize. The significance of this change is hard parent rock (see panel, below). There, it has to
Over time the tissues of this plant to overstate. It turned the world’s sedimentary be found before it eventually breaks apart.
have decayed. Only the outline remains, rocks into a global data bank, teeming with an These body fossils can be breathtaking
covered by a thin film of carbon.
incredible array of fossilized species, arranged objects, particularly when they are complete
in the exact order in which they appeared. skeletons several yards long. However, they are
These fossils show the path evolution has not the only kind of fossilized remains. Rocks
followed. They also highlight mass extinctions— can also yield trace fossils, which are fossilized
when enormous numbers of species have been footprints, burrows, or other signs of animal
wiped out in a relatively short space of time. activity. Trace fossils provide indirect but
fascinating evidence of how animals lived: for
dea d an d bu r i ed example, dinosaur footprints can show how fast
An insect became stuck in resin Fossilization is a lottery, and only a small fraction they moved, how they interacted in herds, and
oozing from a tree. The resin turned into
amber, perfectly preserving the creature. of living organisms end up being preserved. On even how they put on weight as they grew.
land, it is usually triggered by chance events— Much farther back in time, rocks sometimes
for example, when animals are overwhelmed by contain chemical fossils—ancient carbon-based
landslides or flash floods, or when they drown compounds that have been produced by
in lakes. Marine animals have a much better biological processes. Although unspectacular,
chance of being fossilized, because sediment these chemical smudges are key evidence in
routinely accumulates over their dead remains. the hunt for Earth’s earliest living things.
Fine sediment can preserve soft bodies, but the
best fossils are left by animals with hard body
This fish skeleton has been fossilized SUDDEN DEATH
in shale. Here all the pores in the original parts, such as shells or bones. After burial, Trilobites from the Late Ordovician Period are fossilized together,
skeleton have been infilled by minerals. the remains are slowly infiltrated by dissolved suggesting the sudden burial of these animals by sediment.

D E B AT E
INDEX FOSSILS HOW FOSSILS FORM
skeleton layers of
The geological timescale was largely sediment builds compressed compacted
decaying body over skeleton bones by sediment sediment
established using fossils. Species which
lived across a wide geographical range,
but only existed for a short time, are rock exposed by
retreating sea fossilized fish
known as index fossils. They can be skeleton
used to identify particular strata and
link them from place to place—the
presence of the same index fossil in
different places shows that the strata
were laid down at the same time. Index
fossils therefore help geologists to date
rocks and build a relative time sequence. A dead fish lies on the The fish skeleton has been More sediment has Millions of years later
Mesozoic ammonites (a group of extinct seabed, where its flesh covered by sediment. To accumulated on the seabed the seabed sediment, now
marine mollusks) are among the best may rot or be eaten. To be become fossilized, the bones and compressed the lower turned to rock, has been
preserved, it must be buried must undergo a chemical layers. The sediment is exposed by retreating seas.
index fossils—an ammonite zone may rapidly. As the mud turns change, called permineralization, compressed and the fossil Weathering has further
be as little as a million years. into shale, this change will where the pores in the bone are may become flattened, denuded the rock around
compress and flatten the body. replaced by minerals. distorted, or destroyed. the fossilized fish skeleton.
 FOSSIL PLANTS branch
76 Plants are among the first organisms to
appear in the fossil record. Algae are found CLADOXYLON
in rocks from the Precambrian. Vascular STEMS
Cladoxylon scoparium
FOSSILS • PLANTS

plants (plants with tissues for conducting

water and nutrients) evolved in the Silurian Fossilized in Devonian
and Carboniferous rocks,
Period, and by Carboniferous times, Earth Cladoxylon was a low-
was colored green by vast, coal-forming growing plant, with a tough
swamp forests. Flowering plants developed central stem and leafless,
light-absorbing branches.
later, in the Mesozoic Era. EARLY LAND PLANT
Cooksonia hemisphaerica
Found in Silurian and Devonian
rocks, Cooksonia was one of the CALAMOPHYTON STEMS tough stem
earliest vascular plants. It had a stiff Calamophyton primaevum
stem and leafless branches. A primitive, leafless plant, probably
related to ferns, Calamophyton is
found in Devonian and Early
SEED FERN LEAF Carboniferous rocks.
Alethopteris serlii
A seed fern from
Carboniferous and Permian
strata, Alethopteris had
compound pinnate fronds,
consisting of thick, strongly
veined leaflets.

HORSETAIL FOLIAGE
CYCLOPTERIS LEAFLETS SEED FERN SEEDS Asterophyllites equisetiformis
Cyclopteris orbicularis Trigonocarpus adamsi Found in Carboniferous and
Oval leaflets from a seed fern called The name Trigonocarpus is given to Permian strata, Asterophyllites
Neuropteris are given the scientific fossil seeds from seed ferns found had needle-shaped leaves and
name Cyclopteris. Its fossils belong in Carboniferous strata. Each a structure similar to that
to Carboniferous strata. seed has three ribs. of modern horsetails.
CLIMBING HORSETAIL
Sphenophyllum emarginatum
Found fossilized in rocks of
Carboniferous to Permian age, LEPIDODENDRON
this horsetail had wedge- ROOT
shaped leaves and long, soft Stigmaria ficoides
stems adapted for climbing. Occurring in Carboniferous
to Permian rocks, Stigmaria
is the name for the fossil
SIGILLARIA STEM roots of the club moss
Sigillaria aeveolaris relative, Lepidodendron.
Found in Carboniferous and Permian
rocks, Sigillaria was a giant relative
of clubmosses that grew to over vertical rib
100 ft (30 m). It had a narrow stem,
and its leaves grew in clumps.

PERMIAN FERN SALVINIA

Oligocarpia gothanii RHIZOME
This ground-hugging fern Salvinia formosa
is found in Carboniferous and A floating water fern
Permian strata. It occupied from the tropics, Salvinia
a wetland habitat. is found fossilized in
rocks from Cretaceous
to recent strata.

CRETACEOUS
FERN
Weichselia reticulata
Found fossilized in
Cretaceous strata,
Weichselia was similar
to modern bracken and
had fronds that were
divided twice.

SEED FERN LEAFLETS

pinnate Dicrodium sp.
(featherlike) A seed fern from the
leaf Triassic Period, Dicrodium had
pinnate leaves and its fronds
were about 3 in (7.5 cm) long.
 PALEOZOIC
CONIFER
Lebachia piniformis 77
A cone-bearing plant
from Carboniferous
and Permian strata,

FOSSILS • PLANTS
Lebachia is an ancestor
of modern conifers.

CONIFER SEED CONES

Taxodium dubium COAST REDWOOD CONE CRETACEOUS CONIFER
Found in Jurassic strata, Taxodium Sequoia dakotensis Glyptostrobus sp.
is related to modern cypress trees. Cones of the giant evergreen This conifer grew in swamps
It grew in damp habitats and tree Sequoia have been found in during the Cretaceous Period
section had needlelike leaves. Cretaceous and recent rocks. and into the Cenozoic Era.
through Some living members of Sequoia Glyptostrobus was an important
cone are more than 2,000 years old. coal-forming tree.
JURASSIC CONIFER
Araucaria mirabilis
This extinct species of monkey puzzle SUBFOSSIL
tree, A. mirabilis, bore characteristic TREE RESIN
female cones with spirally Kauri pine amber
arranged scales attached Amber is the hardened
to a central axis. resin from pine trees,
such as Kauri pines. First
occurring in the Early
Cretaceous, it often
contains fossils of insects
that perished on the
seed fragrant, sticky resin.

CARBONIFEROUS GIGANTOPTERID LEAVES

GYMNOSPERM Gigantopteris nicotianaefolia
Cordaites sp. A flowerless plant from Permian
An ancestor of the conifers, Cordaites times, this species was so named
grew during the Carboniferous and because its leaves resembled
Permian Periods. It was a tree-sized those of tobacco plants.
plant, which reproduced by seed.

PERMIAN
GINKGO LEAVES
Psygmophyllum
multipartitum
Still found in China, ginkgos
first appeared in the Permian
Period. The fan-shaped leaves
can be identified in fossils of
Psygmophyllum, a precursor TRIASSIC GINKGO
of the modern-day ginkgo. Baiera munsteriana
Growing up to 6 in (15 cm) in
length, the fan-shaped leaves of
Baiera were split into separate
ribs. In living ginkgos, the leaves
are almost entire.

OAK TREE TRUNK

Quercus sp.
growth rings The well-known oak genus, Quercus, first
occurs as fossils in Cretaceous strata. There
are over 500 species of oak living today.

STEMLESS PALM FRUIT MAGNOLIA LEAF

Nypa burtinii Magnolia longipetiolata
central axis Fossils of Nypa date from One of the earliest genera of
the Eocene Period onwards. This flowering plants, magnolias first
palm holds its woody seeds in a appeared in the Cretaceous Period.
10 in (25 cm) ball-shaped fruit. Early insects fed on their nectar.
 FOSSI L I N V E RT E BR AT E S ARCHAEOCYATHID
Metaldetes taylori
corallite
78 Invertebrates, animals without a solid internal skeleton, are These reef-building
organisms are known only
among the most common fossils found. They first appeared from the Cambrian Period.
in the Precambrian, but it is only in Early Cambrian times Metaldetes had a cuplike
FOSSILS • IN VERTEBR ATES

that complex invertebrates such as trilobites become numerous structure, not unlike
in the fossil record. Fossils of invertebrates such as arthropods, that of a coral.
mollusks, brachiopods, echinoderms, and corals are especially
common, because they had hard external structures and lived
in the sea, where most fossil-bearing rocks form.

STROMATOPOROID
tubelike sheetlike Stromatopora concentrica
compartment structure Found in rocks ranging CALCAREOUS SPONGE
from the Ordovician to Peronidella pistilliformis
the Permian, often in reef Characterized by needlelike
limestone, the fossil sponges spicules (in the form of
in this group were made of calcite) fused together,
porous, calcium-rich tubes. Peronidella is found in Triassic
and Cretaceous rocks.

TREPOSTOME BRYOZOAN
Diplotrypa sp.
A bryozoan from Ordovician strata,
Diplotrypa was a small invertebrate,
not unlike coral, that lived in
dome-shaped colonies.

CHEILOSTOME BRYOZOAN
Biflustra sp.
Found in Cenozoic rocks, the Biflustra genus is extant.
These bryozoans have minute compartments, which house
zooids—the soft-bodied individuals of a colony.

LACE CORAL BRANCHING BRYOZOAN

Schizoretepora notopachys Constellaria sp.
A lace coral, Schizoretepora is A bryozoan that built branching
found in Eocene to Pleistocene colonies on the seabed,
strata. It lived on rocky seabeds. Constellaria occurs in
Ordovician strata.
SERPULID WORMS
Rotularia bognoriensis SPRIGGINA
Found in Jurassic to Eocene rocks, Rotularia is a Spriggina floundersi
genus of serpulid worm. Like all serpulids, each A very early fossil found in Ediacaran
worm protected its soft body by producing rocks, Spriggina had a long, wormlike thecae housed
coiled tubes made of calcium carbonate. body. Its classification is uncertain. soft-bodied individuals
of the colony
network
of branches

single, curved
branch

SPIRAL GRAPTOLITE
Monograptus convolutus
“TUNING FORK” GRAPTOLITE BRANCHING GRAPTOLITE A single branch, with thecae
Didymograptus murchisoni Rhabdinopora socialis (cuplike structures) on one side,
A graptolite (extinct colonial invertebrate) with two Until recently, this graptolite was called characterizes Monograptus, which
stipes (branches), Didymograptus is found in Ordovician Dictyonema. It had numerous thin, radiating is found in Early Silurian strata.
rocks. It grew from 3/4 –23½ in (2–60 cm) in length. stipes, and is from Ordovician strata. M. convolutus had unusual coiling.
 TABULATE CORAL
Catenipora sp.
A simple, tabulate coral 79
with a chainlike structure,
Catenipora lived in warm,
shallow seas during
Ordovician and
Silurian times.

chainlike
colonial
structure
SCLERACTINIAN CORAL
Meandrina sp.
Shaped like the human brain, this
colonial coral has ridges and valleys
on its surface. First found in Eocene
rocks, Meandrina still survives today.

RUGOSE CORAL
Goniophyllum pyramidale
A solitary coral occurring in
Silurian rocks, Goniophyllum
has a cone-shaped structure
in which the polyp lived.
thick
corallite wall

CAMBRIAN
TRILOBITE
Paradoxides bohemicus
Some Paradoxides
trilobites grew to nearly
3¼ ft (1 m) in length.
This species had long
spines on its body and is
from Cambrian strata.

SILURIAN TRILOBITE
Dalmanites caudatus
Common in the Silurian
Period, Dalmanites had a DEVONIAN ORDOVICIAN
segmented thorax and ENROLLED TRILOBITE TRILOBITE
a pointed tail spine. Phacops sp. Eodalmanitina macrophtalma
Characterized by compound eyes, Phacops is An Ordovician trilobite, Eodalmanitina
found in Devonian strata. Trilobites could had large, crescent-shaped eyes. Its
roll up, like many modern arthropods. thorax was composed of 11 segments.

pincers

HORSESHOE
CRAB RELATIVE LOBSTER
Euproops rotundatus Eryma leptodactylina CRAB COCKROACH RELATIVE
A Carboniferous relative of A fossil lobster from rocks of Avitelmessus grapsoideus Archimylacris eggintoni
horseshoe crabs, Euproops had Jurassic and Cretaceous times, Covered with many spines, this crab is A relative of cockroaches from the
a crescent-shaped headshield Eryma was 2¼ in (6 cm) long, from the Cretaceous Period. It grew Carboniferous Period, Archimylacris had
and long tail spine. and similar to modern species. to 10 in (25 cm) in width. hind wings with a distinctive vein pattern.
 FOSSI L I N V E RT E BR AT E S SPIRIFERID BRACHIOPOD
Spiriferina walcotti
80 A common brachiopod from Triassic and
Jurassic strata, Spiriferina had a rounded
shell up to 1¼ in (3 cm) wide, with
clearly visible growth lines.
FOSSILS • IN VERTEBR ATES

growth line

ARTICULATE BRACHIOPOD RHYNCHONELLID

Leptaena rhomboidalis BRACHIOPOD
A brachiopod from Ordovician, Homeorhynchia acuta
Silurian, and Devonian strata, Leptaena Found in Early Jurassic strata,
grew to about 2 in (5 cm) in width. Its Homeorhynchia was a small brachiopod
shell had concentric and radial ribs. that grew to about 3/8 in (1 cm) in width.

SWAMP CLAM
Carbonicola pseudorobusta
Occurring in nonmarine rocks
of the Carboniferous Period,
Carbonicola had a tapered shell.
Its fossils have been used in the
relative dating of these rocks.

DEVIL’S TOENAIL
Gryphaea arcuata
This fossil oyster
occurs in Triassic
and Jurassic rocks.
Gryphaea had one
large, hooked valve
CLAM MUSSEL RELATIVE and a smaller, flat one.
Crassatella lamellosa Ambonychia sp.
ribbed valve A small bivalve from Cretaceous to Found in Ordovician strata,
SCALLOP
Pecten maximus Miocene times, Crassatella had marked Ambonychia, an early bivalve
Found in Paleogene to recent rocks, Pecten concentric growth lines on its shell. mollusk, grew up to 21/2 in
is a genus of bivalve mollusks. P. maximus (6 cm) wide. Both valves had
is extant and swims by flapping its valves. radial ribs on the surface. NAUTILOID
Vestinautilus
cariniferous
DEVONIAN This early Nautilus
GASTROPOD relative had very open
Murchisonia bilineata coiling and little
A gastropod mollusk shell ornamentation.
from Silurian to Permian ROSTROCONCH Vestinautilus is found in
strata, Murchisonia grew Conocardium sp. Carboniferous rocks.
up to 2 in (5 cm) tall, Occurring in Devonian
with ridges around and Carboniferous strata,
its whorls. Conocardium resembled
clams, but its shell had
no functional hinge.
JURASSIC GASTROPOD
Pleurotomaria anglica simple rib
Found in Jurassic and Cretaceous rocks, this
whorl with gastropod mollusk had a broad shell with a
ridges combination of radial and spiral patterns.

CARBONIFEROUS DEVONIAN AMMONOID

AMMONOID Soliclymenia paradoxa
Goniatites crenistria An Early Devonian ammonoid, Soliclymenia
An ammonoid mollusk from had thin ribs across the shell. Some species
Devonian and Carboniferous had an unusual, triangular shell.
rocks, Goniatites had angular
sutures where the chamber
walls joined the shell.
AMMONITE
TRIASSIC AMMONOID Mortoniceras rostratum
Ceratites nodosus This ammonite mollusk
Found in Triassic strata, Ceratites from the Cretaceous
shell was an ammonoid mollusk. Its Period grew up to 4 in
aperture heavily ornamented shell had (10 cm) in diameter, with
open coiling and strong ribs. ribs across the shell.
 DEVONIAN CRINOID
Cupressocrinites crassus
Up to 11/4 in (3 cm) in 81
diameter, this crinoid from
Devonian strata had a tall,
five-sided cup at the

FOSSILS • IN VERTEBR ATES

end of its stem.
BELEMNITE respiratory
Pachyteuthis abbreviata flexible arm
structure
A squid relative from the
Jurassic, Pachyteuthis had
a calcite guard. It was
about 4 in (10 cm) long.
JURASSIC CRINOID
Pentacrinites sp.
Named for its five-sided ossicles
(stem segments), Pentacrinites densely
CYSTOID packed
Pseudocrinites bifasciatus grew to more than 31/4 ft (1 m) branches
Characterized by in height. It is often found
rhomboid-shaped respiratory attached to fossil wood.
structures, Pseudocrinites lived stem
in Silurian and Devonian
times. It was attached to the BRITTLESTAR
seabed by its stem. Lapworthura miltoni
An early fossil brittlestar,
tubercle Lapworthura is from
where spine Ordovician and Silurian
was attached strata. It grew to 4 in (10 cm)
in diameter. This species
had five relatively short,
thick arms.

STARFISH
Tropidaster pectinatus
This extinct starfish from the Early
Jurassic was about 1 in (2.5 cm)
wide. It had five thick arms.

BLASTOID
Pentremites pyriformis
This echinoderm from
the blastoid group lived
during Carboniferous
times. It had long,
armlike structures that
were used for feeding.

SEA URCHIN HEART URCHIN

Hemicidaris intermedia Lovenia sp.
This common sea urchin from A heart-shaped, burrowing
Jurassic strata grew to about sea urchin, Lovenia is known
11/2 in (4 cm) in diameter. Its from the Palaeocene and still
many bumps (tubercles) exists today. It is up to 2 in
supported stout spines. (5 cm) in diameter.
 FOSSI L V E RT E BR AT E S ZENASPID FISH
Zenaspis sp.
82 Fossilized vertebrate remains are not as common as those Found in Devonian
strata, Zenaspis had a
of invertebrates, since many vertebrates lived on land, massive headshield. Up
where fewer fossils form, and they evolved much later to 10 in (25 cm) long,
FOSSILS • VERTEBR ATES

than the invertebrates. Fishes were the earliest vertebrates its body was covered
to evolve, some dating back to Cambrian times. Their in bony scales.
rapid evolution in the Silurian and Devonian led to
amphibians, which first appeared in the Devonian Period. finlike structure
Dinosaurs flourished during the Mesozoic Era, toward aided movement
the end of which mammals began to diversify.

EARLY FISHLIKE
VERTEBRATE
Loganellia sp.
A primitive, jawless, flattened
“fish,” Loganellia was covered
with toothlike scales. Up
to 43/4 in (12 cm) long, it is
found in Devonian rocks.

LOBE-FINNED FISH PSAMMOSTEID

Eusthenopteron foordi FISH
The bones in the heavily built Drepanaspis sp. PLACODERM
fins of this Late Devonian A jawless, primitive fish, Bothriolepis canadensis
fish were similar to those in Drepanaspis had a flattened A Devonian placoderm
the limbs of land-dwelling headshield. It is found only (an extinct group of
vertebrates (tetrapods). in Devonian strata. jawless fishes), Bothriolepis
had large head- and
SHARK TOOTH trunk-shields and
Otodus sokolovi spinelike pectoral fins.
The serrated edges of
the teeth of this Cenozoic
shark could easily cut
through flesh.
eye socket

SHOAL OF DACE
Leuciscus pachecoi fine vertebral
Found in Miocene strata, column
extinct species of Leuciscus
STINGRAY or dace resembled modern
Heliobatis radians bony fishes. L. pachecoi grew
Found in Eocene strata, to 2½ in (6 cm) in length.
Heliobatis was a freshwater
stingray that grew to
about 12 in (30 cm) in
length and had a skeleton PRIMITIVE FROG
of cartilage. Rana pueyoi
Dating back to Miocene times,
Rana is a genus of frogs. R. pueyoi grew
to 6 in (15 cm) in length and shared
features such as long hind limbs
with modern frogs. SKULL OF
LARGE, PREDATORY
BONY FISH
Xiphactinus sp.
long, sharp A bony fish from Late Cretaceous
tooth strata, Xiphactinus was a
marine predator with
a muscular body and
large front teeth.

DIPLOCAULID
AMPHIBIAN
Diplocaulus magnicornis
A salamanderlike amphibian
from the Permian Period,
Diplocaulus had protrusions on
the sides of its skull. It grew
up to 3¼ ft (1 m) in length.
 DICYNODONT SKULL
Pelanomodon sp.
This tuskless herbivore was a 83
dicynodont—a member of a group
of mammal relatives that lived in
the Permian and Triassic Periods.

FOSSILS • VERTEBR ATES

DIMETRODON SKULL
Dimetrodon loomisi
Noted for a saillike structure on its back,
Dimetrodon was an early relative of mammals eye socket
from the Permian Period. A high skull and
short snout translated into a powerful bite.

PLESIOSAUR MARINE TURTLE

FLIPPER SKULL
Cryptoclidus eurymerus Puppigerus crassicostata
Growing up to 26 ft (8 m) Fossilized marine turtles are
in length, Cryptoclidus was found in rocks ranging from the
a long-necked plesiosaur Mesozoic Era to Recent times.
from the Jurassic Period. Puppigerus had a heavy shell and
is found in Eocene strata.
GIANT MONITOR CYNODONT SKULL
LIZARD VERTEBRA Cynognathus crateronotus
Varanus priscus A carnivore with a stout skull and large
A huge monitor lizard, Varanus canine teeth, Cynognathus belonged to
priscus grew to 23 ft (7 m) a group of mammal precursors called
in length. It is found in cynodonts. It is found in Triassic strata.
rocks of Pleistocene age.
EARLIEST BIRD
Archaeopteryx GIANT GROUNDBIRD SKULL
lithographica Phorusrhacos inflatus
Archaeopteryx was A carnivore up to 81/4 ft (2.5 m) tall,
thought to be Phorusrhacos was a flightless bird with a
the earliest bird. powerful beak. It is found in Miocene rocks.
However, recent finds
in Jurassic rocks in
China are challenging
that view.

EARLY HORSE TEETH

Protorohippus sp.
A dog-sized, many-toed SABRE-TOOTHED CAT SKULL
ancestor of the modern Smilodon sp.
horse, Protorohippus is Large, curved canine teeth were typical
found in Eocene strata. It of Smilodon, which was the size of a tiger
had low-crowned molars. and lived in the Pleistocene Period.

vertebrae EARLY ELEPHANT JAW

Phiomia serridens
Found in Eocene to Oligocene
strata, Phiomia was 81/4 ft (2.5 m)
tall. It had tusks in its upper
jaw, and a small trunk.

low, sloping
forehead

ANTHROPOID
SKULL
Proconsul africanus NOTOUNGULATE
The first fossil anthropoid Toxodon platensis
(apelike primate) to be Growing to about 8¾ ft (2.7 m), Toxodon had a
found in Africa, Proconsul sturdy body with a hippopotamuslike head. It
comes from Miocene strata. lived from Pliocene to Pleistocene times.
 FOSSI L V E RT E BR AT E S: DI NOSAU R S PLATEOSAURUS
SKULL
84 Plateosaurus sp.
A bulky plant-eater
from the Late Triassic,
Plateosaurus grew to
FOSSILS • VERTEBR ATES

about 26 ft (8 m) in
length and had a
very small head.

bony spike
COELOPHYSIS
DIPLODOCUS BRACHIOSAURUS SKELETON
TAIL VERTEBRA THIGH BONE Coelophysis bauri
Diplodocus longus Brachiosaurus sp. Fossils of Coelophysis are found
Found in Jurassic strata, A huge, plant-eating in Triassic rocks. Only 10 ft
Diplodocus was a giant dinosaur that grew to 82 ft (3 m) long, this carnivore had
plant-eater that grew up (25 m) in length, Brachiosaurus a birdlike skeleton.
to 89 ft (27 m) in length. Its lived in the Jurassic and
tail was long and whiplike. Cretaceous Periods.

skull
PROCERATOSAURUS PARTIAL SKULL
Proceratosaurus bradleyi
A carnivore from Middle Jurassic strata
in Gloucestershire, England, Proceratosaurus
had a bony crest on its head. long tail
used for
balance

MEGALOSAURUS
SACRAL VERTEBRAE
Megalosaurus bucklandi
Found in Middle Jurassic strata, Megalosaurus
was 30 ft (9 m) long, with a large head and
strong hind limbs. It was carnivorous.

COMPSOGNATHUS
SKELETON
Compsognathus longipes
An active predator, Compsognathus
could probably move at speed. It was
only 5 ft (1.5 m) long and is found long hind legs
in Late Jurassic rocks. for running
fast

GALLIMIMUS SKULL
Gallimimus bullatus
Growing up to 20 ft (6 m) long,
Gallimimus had a birdlike, small brain cavity
beaked skull, and long
neck and legs.

strong,
serrated teeth
ALBERTOSAURUS
SKULL
Albertosaurus sp.
A predator and close
relative of Tyrannosaurus DASPLETOSAURUS JAW
rex, Albertosaurus grew Daspletosaurus torosus
to 26 ft (8 m) in length This Cretaceous dinosaur had massive hind legs and
and is found in Late small arms, and it grew to a length of 30 ft (9 m). It had
Cretaceous rocks. a powerful jaw with the formidable teeth of a carnivore.
 85

SCELIDOSAURUS FOOT

FOSSILS • VERTEBR ATES

Scelidosaurus harrisonii
Found in Early Jurassic rocks,
Scelidosaurus grew to 13 ft (4 m) STEGOSAURUS PLATE
in length and was covered with Stegosaurus sp.
sharp, bony knobs. It had A Late Jurassic plant-eater, ANKYLOSAURUS SKULL EUOPLOCEPHALUS TAIL CLUB
long toes and blunt claws. Stegosaurus grew up to 30 ft (9 m) Ankylosaurus magniventris Euoplocephalus tutus
in length. It had two rows of huge Found in Cretaceous rocks, Ankylosaurus Growing to 23 ft (7 m) in length,
bony plates running along its back. was a heavily armored plant-eater. It Euoplocephalus lived in the Late
grew to about 20 ft (6 m) in length. Cretaceous. A bony club at the tip of
its tail was probably used for defense.

PARASAUROLOPHUS SKULL
Parasaurolophus walkeri
A Cretaceous herbivore,
Parasaurolophus had a long, curved,
hollow crest on its skull. This may
have been used to make deep,
resonating calls.

HYPSILOPHODON TOE
Hypsilophodon foxii
Dating back to the Cretaceous
Period, Hypsilophodon was a
swift-moving herbivore. It grew
up to 7½ ft (2.3 m) in length.

PACHYCEPHALOSAURUS SKULL
Pachycephalosaurus wyomingensis
Living at the end of the Cretaceous
Period, Pachycephalosaurus had a thick,
domed skull and grew up to
16 ft (5 m) in length.
STEGOCERAS SKULL
Stegoceras validum
Found in Cretaceous strata,
large nasal cavity Stegoceras grew to 6½ ft (2 m) in
length. Its small, serrated teeth
indicate it was probably a herbivore.

TRICERATOPS SKULL
Triceratops prorsus
cheek horn A massive horned and plated
skull characterized Triceratops,
a plant-eater from Late
Cretaceous strata.

STYRACOSAURUS
SKULL
Styracosaurus albertensis
Although similar to PSITTACOSAURUS SKELETON
Triceratops, Styracosaurus Psittacosaurus sp.
had slender horns along the One of the earliest horned dinosaurs, Psittacosaurus toothless
back of its skull. It is found is found in Cretaceous strata. It was a plant-eater beak
in Late Cretaceous rocks. and grew to 6½ ft (2 m) in length.
86 EUOPLOCEPHALUS
Euoplocephalus belonged to a family of dinosaurs called the
ankylosaurids, which were characterized by an armored
F O S S I L S • V E RT E B R AT E S

head and bony plating on the body. This dinosaur grew to

about 20 ft (6 m) in length and weighed about two tons. Its
tail, body, and neck were covered by plates and bands of tough,
leathery skin set with bony studs. Two rows of larger spikes ran
along its back. Even its eyes were protected by bony eyelids. Fused
bony knobs on the end of the long tail formed a tail club, which
the dinosaur could swing at attacking predators. Euoplocephalus was a size 20 ft (6 m)
herbivore—its beaked mouth was ideally suited for grazing on vegetation in date Late Cretaceous
the thick forests of the Late Cretaceous. It may even have used the blunt hoofs distribution N. America
group Ankylosaurids
at the end of each toe to dig in the ground for roots and tubers. Euoplocephalus
was probably a solitary dinosaur, though juveniles may have lived in herds.

ARMORED HEAD
The head had a massive skull, with short shoulder blade
protective spikes at the back and a beaked
mouth. The name Euoplocephalus means
“well-armored head.”
WALKING TANK
NECK VERTEBRAE Euoplocephalus had a wide, low-
Although the head was relatively small slung body, supported by short,
and the neck short, the neck vertebrae had stout limbs. In cross-section it
to be strong to support the head’s weighty, would have been almost round.
studded armor plating. Its small head was protected at
the back by spikes, and its beaked
mouth would have had small, ribbed
teeth adapted for munching plants.

STUDDED PLATES
One of the most important broad, rounded
ribcage
features of Euoplocephalus
was its armor plating. This
was made of tough plates of
skin studded with ridged
ovals of bone.

FRONT FOOT
This dinosaur’s limbs were short, stocky,
and strong. The front feet had short,
sturdy toes, which helped to support
TAIL CLUB the considerable body weight.
The massive tail club was made of
fused bones—two large and several
smaller bones. This weapon was
probably used in defense.
TAIL VERTEBRAE
About halfway along the tail, the typical tail vertebrae— in life, bony studs were
covered with scaly horn
armed with spikes—gave way to a fused, bony structure.
This rigid structure would have supported the club at the
end of the tail. The tail would have been quite muscular. elbow joint
 bony studs and
spines along back

head spike

massive leg bones

supported the weight
of the armored body

long, convoluted nasal

passages inside the skull
suggest this dinosaur had
a keen sense of smell
hind feet had three
toes, each tipped
with a blunt hoof
Micros
life
copic
Despite their tiny size, microscopic organisms dominate life on
Earth. They were the first living things to evolve, and they underpin
all the world’s ecosystems, capturing and releasing nutrients that
other forms of life need to survive. From the simplest bacteria to
the most complex protists, they make up a constellation of varied
and versatile life-forms that largely goes unseen.

90 94
ARCHAEA AND PROTISTS
BACTERIA Among the most numerous
Life at its most basic, living things, protists are
archaea and bacteria are also some of the most
single-celled organisms diverse. Some have no
without nuclei. Most live fixed shape, but many
as individuals, though have elaborate mineral
some form colonies such skeletons or shells. They
as filaments and chains. are typically single-celled.
90 A RC H A E A A N D
BAC T E R I A
MICROSCOPIC LIFE • ARCHAEA AND BACTERIA

An alien visiting Earth might conclude that its true masters

were the archaea and the bacteria. They outnumber, and are
more diverse than, the more complex life-forms known as the
eukaryotes, and thrive in every nook and cranny of the planet.
DOMAIN ARCHAEA The archaea and bacteria are single-celled organisms
PHYLA 12 and were the first forms of life on Earth. They form
ORDERS 18 two of life’s three fundamental divisions, along with
FAMILIES 28
SPECIES Probably millions
the eukaryotes. While all living cells possess DNA,
eukaryote cells contain it within a membrane-bound
nucleus, and most also have energy-generating
DOMAIN BACTERIA
PHYLA About 50
mitochondria. Archaea and bacteria have neither
ORDERS 180 a nucleus nor mitochondria. They are only distantly
FAMILIES 430 related to each other, having evolved from separate,
SPECIES Probably millions as yet unknown genetic origins. Archaea cells are
contained by chemically unique membranes overlain
by a tough outer cell wall, and within the cell their
DNA is often protein-covered. Bacteria cells have very
different physical and chemical makeups, particularly
in their cell wall. Their different properties make the
typical archaea especially suited to harsh habitats, while
bacteria are ubiquitous and thrive in every environment.

sm a llest living th ings

Hydrothermal vents at the bottom of All archaea and bacteria are tiny, their size measured
oceans are home to various thermophile
archaea living at high temperatures. in microns or micrometers (µm); 1 µm equals one-
thousandth of a millimeter. A human hair is about
80 µm thick; most archaea and bacteria are just 1–10 µm
in length.Yet they survive in almost every corner of
the biosphere, from the outer atmosphere to deep
within Earth’s crust, and from ocean depths to inside
the human body. Some archaea and bacteria thrive in
boiling water or freezing ice, survive radiation, and even
live off poisonous gases and corrosive acids. Most get
nourishment from dead material, others from infesting
Staphylococcus bacteria are common living bodies. While some make food in darkness, by
agents of food poisoning in humans, using the energy in minerals, others use photosynthesis,
forming microscopic clusters in food. using the energy in light to convert carbon dioxide and
water into food and oxygen. Although notorious for
D E B AT E causing infectious diseases, bacteria are actually essential
CAULDRON OF LIFE
to human health, with humans depending on their gut
Many archaea are adapted for bacteria to help break down food and to manufacture
extremes; in hot water their fragile essential nutrients. For almost 4 billion years, archaea
DNA is lagged by protective protein.
Similiar protein braces the longer
and bacteria have profoundly influenced Earth’s climate,
chromosomes of eukaryotes (fungi, rock formation, and the evolution of other forms of life.
plants, and animals). So perhaps what
began as insulation in primeval pools
evolved into a “scaffold” for the extra MICROSCOPIC CYANOBACTERIA COLONY
DNA needed for more complex life. Although bacteria have single cells, the cells of some, such as
cyanobacteria, can join together in spectacular, long filaments.
 A RC H A E A protein- BAC T E R I A
92 making
METHANOCOCCOIDES
granules
BURTONII
This methane producer was
discovered living at the
bottom of Ace Lake in
Antarctica, where there
is no oxygen and the
average temperature
is 33 ºF (0.6 ºC).

1.2 μm
1–4 μm 2–3 μm 1–2 μm

flexible cell wall

ACETOBACTER BACILLUS BACILLUS
ACETI SUBTILIS THURINGIENSIS
Used to make vinegar, this Up to one billion of This bacterium produces
species is also a common these bacteria can be found toxic crystals insoluble in
contaminant wherever in a single gram of soil. the human gut but lethal
alcohol is brewed, causing When food is scarce or the to insects, and is therefore
discoloration and souring of environment becomes harsh it commercially marketed as
beer in particular. can live as an inactive spore. an insecticide.

0.5–15 μm DEINOCOCCUS BORDETELLA

RADIODURANS PERTUSSIS
This species is This species causes
known as the whooping cough,
STAPHYLOTHERMUS world’s toughest or pertussis,
MARINUS bacterium. It was a respiratory
Discovered in a hydrothermal discovered in meat infection marked by
vent on the sea floor, this that had been severe, spasmodic
SULFOLOBUS species grows best at exposed to doses coughing episodes,
ACIDOCALDARIUS 184–198 ºF (85–92 ºC). It of radiation in and breakdown of
Like many archaea, forms grapelike clusters, and an experiment. blood cells. 0.25 μm
this thermophile has a can grow comparatively large.
heat-resistant cell wall. 1.62 μm
It thrives in the high
temperatures of hot ESCHERICHIA COLI
springs in Yellowstone An organism much observed
National Park. and used by scientists, the
rod-shaped E. coli lives
1–5 μm harmlessly in the gut,
although virulent strains
are a major cause of
food poisoning.

1 μm 1.5–4.5 μm 1–3 μm

DESULFUROCOCCUS
MOBILIS
This anaerobe thrives where BACTEROIDES FRAGILIS
oxygen is limited or absent Forming a large part of the
and uses sulfur-containing human gut’s flora, this
80 μm compounds in place bacterium is usually harmless
of oxygen. An extreme to its host. But it can also
THERMOPROTEUS TENAX thermophile, it grows cause disease, invading tissues
This species forms rod-shaped cells of best at 184 ºF (85 ºC). to form pus-filled abscesses.
varying lengths but constant diameter.
Its heat-resistant outer envelope is
ancient in evolutionary terms.
individual rod-shaped
E. coli bacterium
0.8–
2 μm

8 μm

NITROBACTER SP.
PYROCOCCUS FURIOSUS This genus of bacteria plays a
METHANOSPIRILLUM HUNGATEI Pyrococcus means “fireberry,” role in the nitrogen cycle by
Discovered in human sewage, this archaean reflecting this archaean’s shape oxidizing nitrites into nitrates.
generates large amounts of methane during and its tolerance of extreme This helps purify water and
sewage disposal. Each cell is confined temperatures: it grows best is central to fertilizing the
within a hollow sheath. at 212 ºF (100 ºC). 0.6–4 μm oceans and the soil.
 4–8 μm 2–3 μm NOSTOC SP.
cell dividing This cyanobacteria forms
gelatinous filaments in 93
colonies that are hardy
enough to survive from
the poles to the tropics.

MICROSCOPIC LIFE • ARCHAEA AND BACTERIA

3–7 μm
3–8 μm 1–3 μm

DNA
CLOSTRIDIUM CLOSTRIDIUM TETANI SALMONELLA SHIGELLA
BOTULINUM A soil organism that can ENTERICA DYSENTERIAE
This bacterium thrives where grow in dead tissue in Belonging to the same family This gut bacteria produces
oxygen is limited or absent. wounds or burns, C. tetani as E. coli, some subspecies shiga toxin that can cause
Living in soil, it produces produces tetanospasmin, of Salmonella bacteria cause epidemic dysentery. As few
the neurotoxins that cause the neurotoxin that gastroenteritis; others as ten bacteria are needed
botulism, but which also have causes tetanus. cause typhoid fever. to cause an infection.
medical and cosmetic uses.

STREPTOCOCCUS 0.9 μm 1.5–6 μm 1 μm

PNEUMONIAE
This bacterium, present
inside humans, can cause
pneumonia. In children
and the elderly, it is the
leading cause of invasive
infection that moves into LACTOBACILLUS ACIDOPHILUS STAPHYLOCOCCUS EPIDERMIDIS pigmented
all parts of the body. Found in the intestine and vagina, this This spherical bacterium can be found membranes for
photosynthesis
bacterium has nutritional and anti- on the skin as part of normal flora, but
microbial properties. It is used in can cause infections in immune-
probiotic drinks and supplements. compromised patients.

1.5–2 μm
flagellum helps 0.4–0.5 μm
propel organism

1–3 μm

VIBRIO CHOLERAE FUSOBACTERIUM PSYCHROBACTER

Highly mobile curved rods NUCLEATUM URATIVORANS
with a single flagellum at one This species makes its home in This species is a
end, this species secretes a the human mouth, and forms a psychrophile, or cryophile,
potent enterotoxin that major component of plaque. It which means it thrives at
causes cholera. can also cause premature birth. very low temperatures,
thanks to natural antifreeze
molecules in its cytoplasm
NITROSOSPIRA SP. within the cell membrane.
Filling a vital ecological 1 μm
niche, this nitrifying soil
numerous rods bacteria oxidizes ammonia cell wall
of bacteria to form nitrites as part of
grouped together the nitrogen cycle.

cell contents,
or cytoplasm

1–3 μm

ENTEROCOCCUS
FAECALIS
Normally a harmless YERSINIA PESTIS
inhabitant of the human This species is the agent that causes
digestive tract and vagina, bubonic plague. Transmitted to
this species can invade humans via fleas that live on rats,
wounds and is resistant to it causes up to 3,000 cases
many antibiotics. 0.5–1 μm worldwide each year.
94 P RO T I S T S
From tiny amoebas to giant kelp, the protists defy simple
MICROSCOPIC LIFE • PROTISTS

description, yet this informal grouping of eukaryotes included the

first life-forms to evolve that were more complex than the archaea
and bacteria. It still produces most of Earth’s food and oxygen.

DOMAIN EUKARYOTA Protists are mainly single-celled creatures that, unlike

KINGDOM PROTISTA archaea and bacteria, have cell nuclei. Their basic cell
CLADES AND DIVISIONS More than 9 nature separates them from the higher eukaryotes—
FAMILIES About 841
SPECIES About 73,500
plants, fungi, and animals—that later emerged from
them. The protists include an incredible range of
organisms with diverse lifestyles and ecological niches.
Most are microscopic, ranging in size from 10 to
100 µm, and some are tiny enough to infest red blood
cells. Others, however, are multicellular organisms such
as kelp, a seaweed that grows up to tens of yards in
length, or the strange, funguslike slime molds, which
form creeping puddles of slime that are essentially
one giant cell. Typical protists include amoebas, which
In some slime molds, which are a move around and capture food particles with their
group of amoebas, thousands of individual pseudopods (cell extensions), or plankton drifting
nuclei coexist within one giant cell.
in the sea, such as the beautiful diatoms, which have
intricate silicon-based skeletons.

a h i dden r ea lm of li fe
Protists are among the most numerous creatures
on Earth. Vast numbers of them live in the oceans
and rivers, in sediments at the bottom of seas and
lakes, and in the soil, while many spend all or part
Many single-celled protists have of their life cycles as parasites inside other organisms.
extraordinary shapes, such as these
grapplelike dinoflagellates. They play a vital role in the planet’s ecosphere, in
particular as primary photosynthesizers, using the
energy in light to convert carbon dioxide and water
into food, while releasing oxygen into the atmosphere.
They also act as predators and recyclers of matter. A
few species are well known because they cause major
diseases: the parasitic Plasmodium causes malaria, one
of the biggest killers of humans; another parasite,
Trypanosoma brucei, causes sleeping sickness. Also
Some protists cause devastating
disease, such as this Giardia, which infects familiar are the dinoflagellates, a group of plankton
intestines of humans and other animals. organisms that can cause “red tides”—huge blooms
of toxic creatures that kill fishes and poison humans.
D E B AT E The taxonomy of the protists is complex, as they
M U LT I P L E K I N G D O M S don’t form a natural kingdom. However, molecular
and genetic analysis has allowed most protists to
The kingdom Protista includes many
organisms that are not classified
be placed in a few large clades, groups that each share
as fungi, plants, or animals. And it a common ancestor, such as the Amoebozoa, Rhizaria,
includes many groups, ranging from and Alveolata. The red algae, green algae, and the
single-celled amoebas to multicellular stoneworts each have their own division.
seaweeds, that are not closely related
to one another. Many scientists think
that Protista should be split into more SMALL WONDERS
than one kingdom. This polarized light micrograph reveals an Arcella shelled
amoeba and a Micrasterias desmid in all their beauty.
p ro t i s t g ro u p s
The taxonomic groupings for the
protists, which can be difficult to
order, are new and evolving and
can be subject to argument. The
latest thinking is set out here.
a m o e ba s a n d
r e l at i v e s

96
f l ag e l l at e s

97
rhizarians

98
a lv e o l at e s

100
h e t e ro kon t s

101
red a lga e

103
green a lga e

105
s t on e wo rt s a n d
r e l at i v e s

105
96 A M O E B A S A N D R E L AT I V E S
Two of the protist clades, the aggregate into a tiny creeping slug. This then DOMAIN EUKARYOTA
Amoebozoa and the Opisthokonta, sprouts upward a number of stalks, which burst
P ROT I STS • A M O E B A S A N D R E L AT I V E S

KINGDOM PROTISTA

have evolved different ways of moving to scatter spores. Each spore hatches into a new CLADES 2

around and obtaining food. amoeba ready to hunt on new ground. FAMILIES About 50
SPECIES About 4,000

Amoebas of the clade Amoebozoa can change a n i m a l- f u n g u s o r i g i n s

shape by oozing projections, called pseudopods, Most protists of the clade Opisthokonta D E B AT E
of their single cells. They use these “false feet” have developed a single whiplike flagellum for LIFE’S FIRST BRANCHES?
to creep forward and hunt smaller organisms, propulsion in open water. Early in the history
enveloping victims within a sac of fluid and of life, some of these protists may have given One theory of evolution says that the
digesting them while still alive. Some amoebas are rise to animals, and the single flagellum is still eukaryotes divided into two branches:
giant cells, and are just visible to the naked eye. seen in the tail of animal sperm today. Others, unikonts (cells with one flagellum),
such as opisthokonts, and bikonts
A few are gut parasites, causing amoebic dysentery called nucleariids, lost their flagellum and actually (cells with two). Unikonts evolved into
in humans. One group, the slime molds, have reverted to the amoebalike state. The nucleariids animals and fungi, while bikonts gave
a remarkable strategy for avoiding starvation. may be closely related to fungi, as fungi also rise to plants. However, the DNA
When prey runs low their cells are attracted to lack flagella and fertilize one another without evidence for the theory is equivocal.
one another by a chemical distress signal, and the involvement of free-swimming sperm.

15–50 μm 49–53 μm 100–130 μm pseudopod

ingested
alga
DYSENTERY AMOEBA ARCELLA ARCELLA VULGARIS
Entamoeba histolytica BATHYSTOMA Mainly found in stagnant water
This parasitic amoeba lives in This amoeba has a circular shell and soil, this amoeba has a convex
the human intestine and can pitted with pores, domed on one shell, with a hole that allows
cause amoebic dysentery. It can side but sometimes with angular pseudopods to emerge.
contain up to eight nuclei. facets that develop into spines.
one of two nuclei
90–110 μm
PROTACANTHAMOEBA
CALEDONICA ARCELLA DISCOIDES
Originally identified in This amoeba with two nuclei
a Scottish estuary, close has a yellow-brown shell with
relations of this amoeba have a hole on one side through
been found in the liver of which pseudopods emerge.
tench in the Czech Republic.
10–16 μm
19–40 μm

NAEGLERIA
FOWLERI
This so-called “brain- 120–150 μm
eating amoeba” lives in
warm, fresh water.
It enters the human CENTROPYXIS ACULEATA
body through the nose,
causing severe brain This amoeba lives on algae in lakes
damage, which is fatal in and marshes. It uses sand and the
98 percent of cases. cell walls of some algae to make a
shell with four to six spines.

pseudopod
180–230 μm 1.2–2.2 μm
4∕5 in
2 cm DIFFLUGIA SCALY NUCLEARIID
PROTEIFORMIS Pompholyxophrys ovuligera
STEMONITIS SP. This pond-ooze amoeba Once assigned to the
This “chocolate tube” or “pipe cleaner” constructs a shell by heliozoans clade, this
slime mold begins as a mass of cell sticking together minuscule flagellate of the Opisthokonta
content with many nuclei from which grains of sand and the cell is covered in hollow
numerous spore-bearing stalks sprout up. walls of some algae. scales or beads.
F L AG E L L AT E S 97
The whiplike propulsion of single- photosynthesize; but in darkness their light- DOMAIN EUKARYOTA
celled organisms has evolved in several absorbing cell organelles, known as chloroplasts,

P ROT I STS • F L AG E L L AT E S
KINGDOM PROTISTA

groups of protists, some unrelated. shrivel and they revert to being a predator. CLADE EXCAVATA
FAMILIES 40
But it predominates in the Excavata. SPECIES About 2,500
li v i ng i nsi de an i m a ls
Flagellates are powerful swimming microbes Many flagellates of this group lack the ordinary
that move by the whiplike action of one or cellular features that would enable them to use D E B AT E
more external threads called flagella. Many are oxygen in their respiration and survive inside the EVOLUTION DOWN-SIZING
predators, preying on smaller organisms such as oxygen-poor environment of animal guts. Many
bacteria. However, unlike changeable amoebas, are extraordinarily specialized, subsisting on the The flagellates inside termite guts lack
these flagellates have a fixed, rigid shape and partially-digested food inside the abdomens of mitochondria, the cell organelles most
direct food into a cellular “mouth” at the base of insects, but causing their host no harm. Others other protists use to respire with
oxygen. Some think these flagellates
the flagella. Remarkably, however, some, such as are parasites that cause devastating disease, are primitive protists. Others say they
Euglena, have a degree of behavioral versatility including in humans. One notorious group, the are more advanced organisms, and
that provides them with plant- and animal-type trypanosomes, are transmitted by biting insects lost their mitochondria by evolving in
nutrition, depending upon circumstance. When and are responsible for sleeping sickness and an oxygen-poor environment.
exposed to bright light these organisms can leishmaniasis in the tropics.

35–55 μm 50 μm 10 μm
chloroplast SLENDER
EUGLENA
Euglena gracilis
This species, with
a single external chloroplast
flagellum, can
aggregate with
others to form a
gelatinous slime.
EUGLENA
MUTABILIS
50 μm VANED FLAGELLATE
Normally long and
wormlike, this species Jakoba bahamensis
can also change shape. four flagella This flagellate lives associated
GREEN EUGLENA Some species of the with marine sediment,
Euglena viridis Euglena genus have feeding on detritus. One of its
A common species often forming external flagella, two flagella may be concealed
blooms, this free-swimming others do not. 16 μm within its feeding groove.
flagellate has a flexible cell wall.

flagellum

125–130 μm

EUGLENA SPIROGYRA
This green or yellow-brown
species of the Euglena genus has
two flagella. The coat of its cell
TERMITE FLAGELLATE body is ornamented with
Trichomonoides trypanoides rows of wartlike bumps.
Living in the guts of termites,
where it helps digest wood,
this species has four flagella 18–29 μm
at one end and a fifth running
along its body.
cyst covers
cell body
10–20 μm

9–21 μm
GIARDIA LAMBLIA one of two nuclei
Also known as Giardia intestinalis,
this intestinal parasite causes the
disease giardiasis. It makes a
dormant cyst before entering its LEISHMANIA TROPICA TRYPANOSOMA BRUCEI
host, and has paired nuclei. flagellum for Spread by sandflies, this parasitic This organism causes sleeping
movement species causes cutaneous sickness, or trypanosomiasis,
leishmaniasis, a skin disease which is transmitted between
affecting more than 1 million vertebrate hosts by the bite
people a year worldwide. of the tsetse fly.
98 RHIZARIANS
The clade Rhizaria includes two phyla photosynthetic partners make as they drift in DOMAIN EUKARYOTA
that are among the most beautiful sunlit tropical oceans. Their silica dependence
PROTISTS • RHIZARIANS

KINGDOM PROTISTA

of all of the miniature protists: binds radiolarians to the sea, but their allies, the CLADE RHIZARIA

the radiolarians and the forams. cercozoans, have also exploited soil and fresh water. FAMILIES 108

Cercozoans typically retain the long pseudopods, SPECIES About 14,000

The unique, intricately sculptured shells of but have shelled, and shell-less forms, and some
both phyla make them distinctive members of have flagella, depending on habitat demands. D E B AT E
the microscopic world, and some have left an The forams, or foraminiferans, have flourished CONVERGING GIANTS
impressive fossil record. Most radiolarians build in the oceans for hundreds of millions of years and
a glassy shell with silica, stripping the ocean surface their calcium carbonate shells litter the seabed to The pseudopods of radiolarians and
of this otherwise abundant mineral when they form layers of chalky sediment. Foram shells are forams enmesh to form nets around
their cells. This perhaps supports the
bloom in large numbers. Their body radiates long so distinctive, even when fossilized, that geologists
idea that both forms descend from a
pseudopods that poke through their shell like can use them to date deposits and identify hidden common ancestor. But it is possible
the rays of the sun, supplemented in some by reservoirs of oil. When alive, each shell harbors that this net evolved independently
hardened spines. Radiolarians use their pseudopods a tiny amoeba that hunts with pseudopods—like in the two groups, perhaps a result
for catching food, but some also harbor living the radiolarians—although some are big enough of both having large single cells.
algae, gaining nourishment from the sugars their to catch animal larvae.

LATTICE-SHELLED ¼ in/6 mm 110 μm

RADIOLARIAN
This radiolarian species of
plankton belongs to the
Spumellaria order, most
of which have silica 350 µm
latticework skeletons.

100 µm

STAR RADIOLARIAN SPHERICAL RADIOLARIAN DIDYMOCYRTIS

Astrolithium sp. Physematium sp. TETRATHALAMUS
In this radiolarian, the bases of The cell surface of this species holds A living representative
radial spine of silica the radial spines are fused into a photosynthesizing organisms in a of an ancient fossil
star-shaped mass, or central body. mesh of thin pseudopods. lineage, this radiolarian
is named for its twin
shells, which sometimes
SPONGASTERICUS have hemispherical or
QUADRICORNIS conical caps.
One of the radiolarians used by
geologists to help date strata by stiffened
the presence of its fossils, this pseudopod
species’ name means
“four-cornered.”

150–302 μm 200–289 μm

70–120 μm

SPINY
RADIOLARIAN
SUN RADIOLARIAN Lithomelissa setosa THREE-ARMED
Heliodiscus sp. This species displays the RADIOLARIAN
Sun radiolarians send greatest seasonal variation Euchitonia elegans
119–191 μm out pseudopods through of any radiolarian, making This three-armed radiolarian
the many pores in their it a useful seasonal forms a particularly spiny
silica skeleton to indicator in fossil and shell. Stiffened pseudopods
pores on surface catch food. sedimentary records. emerge from shell pores.
 20–32 μm 60–90 μm SPHAGNUM CERCOZOAN
Archerella flavum
This shelled amoeba lives in 99
sphagnum wetlands. Its fossils
are used as indicators of
shell past climate changes.

PROTISTS • RHIZARIANS
45–77 μm

protective lorica

captured
cyanobacterium ELEGANT CERCOZOAN
Clathrulina elegans
nucleus This species spends most of its life
as an amoeba inside a perforated
organic capsule, or lorica.
OVAL-GREEN
CERCOZOAN rear flagellum
Paulinella chromatophora
This freshwater shelled HELKESIMASTIX LONG-
amoeba contains living SP. FLAGELLATED
cyanobacteria that can Using its flagellum, this CERCOZOAN
photosynthesize, providing species glides about Cercomonas longicauda
the amoeba with food. preying on bacteria and This species moves
even other species of using its two flagella,
the same genus. one of which trails
behind the body, while
oval cell the other lies ahead.
6–7 μm 40–150 µm

14 μm
pseudopod flagellum surface scales

open “mouth” 18–36 μm

SCALY CERCOZOAN
Euglypha sp.
This amoeba constructs
35–102 μm 60–200 μm
oval-shaped shells from
thin scales of silica, which flagellum lies
fossilize well, making them ahead of body
objects of study for
micropaleontologists.

chloroplast threadlike pseudopods

radiate from cell
GYMNOCHLORA cell wall encrusted
STELLATA by protective layer
Unlike many cercozoans, this of sand grains
species contains chloroplasts,
which enable it to gain food
by photosynthesizing in
brightly lit waters.

pseudopod OVAL-SHELLED YELLOW-SHELLED

CERCOZOAN CERCOZOAN
Trinema sp. Cyphoderia ampulla
This shell of this amoeba is This amoeba has a colorless or
open at one end, creating yellow shell composed of
a type of mouth. disk-shaped scales.

20 μm 11∕2 in/38 mm 35–50 μm

MARINE-GREEN GIANT CERCOZOAN SAND FORAM

CERCOZOAN Gromia sphaerica Lithocolla globosa
Chlorarachnion reptans This species of cercozoan lives This species of foram appears like
This species contains characteristic at the bottom of the ocean at a radiating sun. It builds up a
cell organelles, which are tiny the boundary between sea protective layer of sand grains
relics of a green alga it ingested water and sediment. It feeds and other granules on the outside
in its early evolution. on organic detritus. of its cell wall.
100 A LV E O L AT E S
Alveolates are united by possession of Most ciliates feed on smaller organisms. Their DOMAIN EUKARYOTA
the same anatomical quirk—a fringe of smooth graceful motion is due to the coordinated
PROT IST S • A LV EOL AT ES

KINGDOM PROTISTA

tiny sacs around the cell called alveoli, beating of countless tiny hairs called cilia, which CLADE ALVEOLATA

after which the clade is named. can cover their entire single-celled body. They FAMILIES 222

also use these cilia to waft their food into a SPECIES About 20,000

Three groups of superficially different but furrow that approximates to a mouth. Ciliates
still single-celled protists are included in the are virtually ubiquitous; some even live in the D E B AT E
alveolates: the dinoflagellates, the ciliates, and stomachs of mammalian herbivores, where they PROTOZOANS
the apicomplexans. Predatory dinoflagellates help in digesting the tough cellulose in plants.
live in the ocean, most using two whiplike In contrast, all apicomplexans are parasites. Early classifications placed food-
flagella that emerge from grooves in their cell They are named for an arrangement of structures, eating microbes into different classes
of the animal phylum Protozoa.
coat at right angles to one another to swim. Some called the apical complex, which helps them
Some microbes, such as the ciliates
can discharge stinging barbs to immobilize their penetrate the living cells of animals, from which and the dinoflagellates, are now
prey. Others release poisons, and sudden blooms they derive their nourishment. Infamous among grouped together in modern protist
of dinoflagellates are responsible for toxic red tides the apicomplexans are the malaria parasites, taxonomy, although the exact nature
in parts of the world. A few are bioluminescent, which penetrate the red blood cells of animals of their relationships is debatable.
sparkling with light when disturbed at night. for food, destroying the cells in the process.

STALKED CILIATE
Vorticella sp.
This organism has an inverted
bell-shaped body on a stalk,
which is known to coil like a
spring when stimulated.

50–160 µm 35–90 µm 40–110 µm 50–130 µm

6 µm

SOIL CILIATE POND CILIATE GUT CILIATE TOXOPLASMA GONDII

1∕10 in/2–3 mm Colpoda inflata Colpoda cucullus Balantidium coli Passing between cats and other
This normally kidney-shaped Usually found in fresh water This is the only ciliate mammals (including humans),
TRUMPET CILIATE species plays an important among decaying plants, this known to parasitize this apicomplexan causes the
Stentor muelleri role in soil ecology but is ciliate has food-containing humans. Infection can cause disease toxoplasmosis, which
This algae-feeding species, with a horn-shaped vulnerable to pesticides. organelles called vacuoles intestinal ulcers, or serious is dangerous to the unborn
cell body, is large for a single-celled organism. within its cell. intestinal infections. child in pregnancy.

225 µm 19–17 µm 4–6 µm 9–14 µm

38–50 µm

CHAIN-FORMING MAHOGANY-TIDE CRYPTOSPORIDIUM PLASMODIUM

THREE-HORNED DINOFLAGELLATE DINOFLAGELLATE PARVUM FALCIPARUM
DINOFLAGELLATE Gymnodinium catenatum Karlodinium veneficum This apicomplexan can The most deadly of the
Tripos muelleri This species of This dinoflagellate plankton can cause cryptosporidiosis, malaria-causing Plasmodium
Formerly known as Ceratium dinoflagellate forms long, undergo a population explosion a diarrhoeal disease species, this apicomplexan
tripos and among the most swimming chains of up leading to “mahogany tides” transmitted by ingestion of parasite is responsible for the
distinctive dinoflagellates, to 32 cells. that are lethal to fishes. its spores via feces, usually deaths of over 1 million
blooms of this plankton can in contaminated water. people a year worldwide.
cause dangerous red tides. organelle
200–2,000 µm
10–100 µm 20–40 µm
40–74 µm 11–24 µm

AKASHIWO
SANGUINEA
A large pentagonal species
responsible for several recorded
gas bag instances of harmful algal
blooms, this dinoflagellate can
both photosynthesize and prey
on other plankton.
GYMNODINIUM SP. KARENIA BREVIS
Producing a neurotoxin, Formerly known as SEASPARKLE AMPHIDINIUM CARTERAE
blooms of this dinoflagellate, Gymnodinium brevis, this Noctiluca scintillans This plankton-forming species
found in both fresh and salt dinoflagellate plankton is This bioluminescent, plankton- causes ciguatera, a fish poisoning
water, cause red tides and a major cause of red tides forming species has a gas bag allowing that can cause human fatalities if
shellfish poisoning. in the Gulf of Mexico. it to float just below the surface. contaminated fishes are consumed.
H E T E RO KO N T S 101
Heterokonts include some types of their pigments absorb energy from sunlight to DOMAIN EUKARYOTA
algae, which are photosynthesizing make food. Diatoms contain green chlorophyll—

PROTISTS • HETEROKONTS
KINGDOM PROTISTA

protists growing in or near water that as do plants—but also a brown pigment called CLADE HETEROKONTOPHYTA

do not have true leaves or roots. fucoxanthin, which serves to broaden the FAMILIES 177

spectrum of light that can be used, making SPECIES About 20,000

Heterokonts are mostly defined as having two their photosynthesis significantly more efficient.
different types of flagella on the sperm they use Brown algae occupy coastal habitats around the D E B AT E
in reproduction. One flagellum is covered in world. They also make use of fucoxanthin, and FROM ALGA TO MOLD
tiny, bristly hairs, or mastigonemes, and the other have evolved into complex multicellular seaweeds
is smooth and whiplike. Heterokonts include that superficially resemble a plant. Rather than Water molds grow and feed like mold,
diatoms and brown algae, as well as water molds. true roots and leaves, their body consists of but unlike true molds, which are fungi,
have plantlike cell walls and heterokont
Diatoms are single-celled algae with finely a holdfast for clasping to rock and a creeping
flagella, while some also cause plant
sculptured silica walls divided into two parts thallus that lacks the internal veins of a true leaf. diseases. DNA analysis links them with
called valves. They are the dominant eukaryotic Nevertheless, some brown seaweeds, such as diatoms and brown algae, so perhaps
phytoplankton—plankton communities of kelps, can grow to extraordinary lengths and they evolved from algae that abandoned
small organisms that drift in open waters and make extensive underwater forests on some their chloroplasts and turned parasitic.
engage in photosynthesis. Here, near the surface, offshore coastlines.

80–125 µm 50–80 µm 25–200 µm

1∕10 in/0.3 mm 60–240 µm

EPIPHYTIC DIATOM BARREL DIATOM SADDLE DIATOM COLONIAL DIATOM GYROSIGMA SP.
Biddulphia sp. Biddulphia pulchella Campylodiscus sp. Isthmia nervosa The name of this diatom refers
The cause of brown film on fish This image clearly displays the different This genus of diatom shows a groove This species grows on other to the sigmoidal curve of its
tanks, in the wild this species parts of a diatom: the two valves bound between the tubular lips, known as algae, specifically seaweeds, cell, meaning that it follows
grows on seaweed and rocks. together by a narrow girdle. canals, around the edge of its valves. forming branching colonies. a very slight S-shape.

STEPHANODISCUS SP.
12–20 µm disk-shaped girdle
With its disk shape, areolae
(open circles), and a ring of
spines, this diatom occurs
individually or in chains.

125 μm 18–90 µm
spine
POND DIATOM
SLIMY DIATOM
Pinnularia sp.
Lyrella lyra Two chloroplasts are visible in
This species secretes a gluey this pen-shaped diatom, found
slime from its central groove in ponds and wet ground.
to help the cells to glide
along their host’s surface. areola

10–100 µm

200–1,000 µm

GROOVED DIATOM
URCHIN HELIOZOAN Diploneis sp.
Actinosphaerium sp. The valves of this species of
This diatom resembles a sea diatom appear as two heavily
urchin. It moves by shifting pronounced lips, known as
cell contents into its canals, either side of a slitlike
thin pseudopods. groove known as a raphe.
 H E T E ROKON TS
102
20–220 μm THREE-CORNERED
DIATOM
Actinoptychus sp.
This diatom can be found
in the neritic zone on the
shallow continental shelf.

pores allow intake

of minerals for use
in photosynthesis
1∕32 in
0.9 mm

SPIDER’S-WEB DIATOM
Arachnoidiscus sp.
Strong radial ribs and web-patterned
valves are features of this disk-shaped
diatom. It can grow very large.
top valve

220 μm 20–220 μm

protective cell wall

140 μm 140 μm
TRICERATIUM FAVUS
The heavily silicified cell
walls of this species can
leave a trail of silica in the
water as it moves. This
SUN-RAY DIATOM FIVE-ARMED DIATOM TRICERATIUM SP. can show how seawater
Actinoptychus heliopelta Actinoptychus sp. More than 400 species of penetrates into freshwater
A distinctive pattern of alternating A five-arm appearance is this genus of marine diatom environments.
raised and depressed sectors on produced by the alternating are known. Diatoms of this
its top and bottom valves sectors on the valves of this genus are often, but not geometric cell wall
characterize this diatom species. species of the Actinoptychus genus. always, three sided.

fronds are leaflike

structures used in
photosynthesis
air bladder on
frond helps keep
seaweed bouyant

11–125 μm 44–82 μm

DAPPLED DIATOM
BOAT-SHAPED DIATOM Stictodiscus sp.
Navicula sp. Seen under a scanning electron
This species belongs to the microscope, this diatom shows
largest genus of diatoms, which pores dappled on its valve surface,
has thousands of species. above a surrounding girdle.

shrublike
fronds
6½ ft
2m

13 ft 6½–11 ft 3¼–10 ft 12–39 in

4m 2–3.5 m 1–3 m 30–100 cm
23½ in
60 cm
SUGAR KELP CUVIE WIREWEED SEA OAK
THONGWEED TOOTHED WRACK Saccharina latifolia Laminaria hyperborea Sargassum muticum Halidrys siliquosa
Himanthalia elongata Fucus serratus Found on rocky shores This alga is commercially Originally from Japan, A large brown seaweed,
During its reproductive phase, This robust, shrubby, and at depths of important in iodine this invasive brown common in rock pools
this brown seaweed of the olive-brown seaweed 26–100 ft (8–30 m) in production. It grows seaweed has now spread in Europe, this species
northern hemisphere produces grows on lower northern seas, this brown at depths of 26–100 ft to Europe. It can grow as has a distinctive zigzag
long straplike fronds, which shores all around the seaweed is also often (8–30 m), mainly in the much as 4 in (10 cm) stem and air-filled pods
are types of divided leaves. North Atlantic. known as sugar wrack. northern hemisphere. in a day. known as air bladders.
RED ALGAE 103
Although some are microscopic r e d f o r s u rv i va l DOMAIN EUKARYOTA
organisms, the most familiar red algae Like land plants, most algae use green chlorophyll

PROTISTS • RED ALGAE

KINGDOM PROTISTA

are multicellular seaweeds, which are to trap the energy of sunlight and manufacture DIVISION RHODOPHYTA

among the largest of the protists. They food in the process of photosynthesis. However, the FAMILIES 92
SPECIES About 6,500
have evolved into an extraordinary cells of red algae also contain a red pigment, called
phycoerythrin, which masks the green color of the
diversity of forms, ranging from rock- chlorophyll and gives them their red color—and
hugging encrustations that look more D E B AT E
their name. This pigment allows them to continue C L O S E R E L AT I V E S ?
like lichens to bristly or leafy clumps. photosynthesizing even at depths in the water
where only a little blue light can penetrate. This Red algae were previously grouped with
The great majority of the 6,500 or so species of red green algae, but it is now recognized
allows red algae to live much deeper in the oceans
algae are marine, but there are a few freshwater than brown or green seaweeds, and some have that green algae are two separate and
species. Unlike brown and green algae, they do not very diverse groups. Some species
been reported growing as deep as 650 ft (200 m). may be close to red algae, but others
produce flagellated sperm and rely instead on water Some red algae become calcified, forming are closer to land plants. Red algae
currents to transport male sex cells to the female encrustations on rocks or hard, upright, antler- branched off the taxonomic tree before
organs. The cells then merge, but subsequent like forms. And, in spite of their name, their the land plants began to evolve.
development is highly variable between species. pigments can make red algae appear olive or gray.

20 in 2∕5–6 in
50 cm 1–15 cm

16 in
40 cm
31∕4 in
8 cm

DULSE CORAL WEED AGARDHIELLA SUBULATA SCHMITZIA HISCOCKIANA

Palmaria palmata Corallina officinalis Originally from the western Found in areas swept almost bare by
In N. Atlantic coastal communities, This red seaweed is common in Atlantic, Caribbean, and Gulf of the tide, this red seaweed is fleshy and
this edible seaweed is a traditional rock pools worldwide, forming Mexico, this fleshy red alga has gelatinous, with flattened branches
source of protein and vitamins. branching, featherlike fronds. invaded some parts of Europe. and fingerlike projections.

12 in 12 in
30 cm 30 cm EYELASH WEED
MAERL Calliblepharis
Phymatolithon calcareum ciliata
This species, an encrusting This northern
coralline alga of the British hemisphere seaweed
Isles, is commercially dredged has flat fronds,
23∕4 in and ground up as a calcium-
7 cm fringed with
brittle rich soil additive. numerous
branch BLACK “branchlets.”
CARRAGEEN
Furcellaria
lumbricalis
12 in
30 cm This northern
hemisphere seaweed
has cylindrical,
brown-black fronds
that branch into masses
of fleshy fingers.
tubular frond
fanlike branch

frond

61⁄2 in
4–12 in 17 cm 8¾ in
10–30 cm 22 cm
MASTOCARPUS
GRACILARIA GRACILARIA STELLATUS
FOLIIFERA BURSA-PASTORIS This red seaweed has IRISH MOSS
Rising as slender dull purple A long, thin red seaweed with conspicuous papillae, Chondrus crispus
stems, this red seaweed branches forked or alternating branches, which are reproductive Also known as carrageen moss, this
sparsely to give straplike fronds. It this species is found from bodies, on the fronds. seaweed of the British Isles is
grows abundantly in shallow southern England to the Pacific It is found in the important as a source of carrageen,
lagoons worldwide. and the Caribbean. North Atlantic. which is a gelling agent.
 RED ALGAE 6 in
15 cm
104
8 in
20 cm
PROTISTS • RED ALGAE

flat, leaflike
structure

PTEROCLADIELLA CAPILLACEA
With feathery branches that taper BUSH WEED
toward the tip, this red seaweed, found Ahnfeltia sp.
in pools worldwide, often develops a A source of the gelatinous
Christmas-treelike appearance. substance agar, used in petri
dishes for cell culture, this
northern-hemisphere seaweed
forms dense tufts of fronds.
1∕10 in
2.5 mm
61⁄2 in
17 cm
blades on
MELANAMANSIA frond
FIMBRIFOLIA 2¾–8¾ in
Found in N. America and 7–22 cm
Australia, this seaweed grows
on sediment-covered reefs at
depths of up to 180 ft (55 m).

CHONDRIA DASYPHYLLA
SPOROLITHON This species, found worldwide,
PTYCHOIDES has feathery fronds ending in
This red algae forms a crust using clublike branchlets that sprout
calcareous deposits in its cell walls. spore-bearing branchlets and
It is found worldwide in rock urn-shaped fruit bodies.
pools and on current-swept
4–81⁄2 in LAURENCIA OBTUSA
rocky seabeds. 10–21 cm
This tropical species is a source
12 in of halogenated terpenoids—
30 cm chemical defenses against grazing
crabs and urchins—which also
make useful antifouling agents.

8 in
LEATHER WEED
20 cm Ptilophora leliaertii
First described in 2004
after its discovery on a
reef off the coast of
South Africa, this red
seaweed has feathery 12 in
CERAMIUM VIRGATUM compound branches. 30 cm
This small red seaweed grows 14 in
worldwide on rocks and other 35 cm
seaweeds. From a tiny holdfast
it grows into filamentlike SEA BEECH
fronds with forked tips. Delesseria sanguinea
Known for its characteristic beechlike
1∕10–3∕5 in
leaves, this red seaweed grows in the
2–15 mm understory of kelp forests in Europe.
GELIDIELLA ACEROSA
An important source of
agar, used in the food and
pharmaceutical industries,
this red seaweed, found
in India, has a slender,
cylindrical shoot, known
as a stolon.

stolon 31∕4 in ¾–4 in

BROAD WEED 8.5 cm 2–10 cm
GELIDIUM PUSILLUM Lenormandiopsis nozawae
Growing worldwide from an On both sides of this
extensive creeping base that broad-bladed seaweed, POLYSIPHONIA LANOSA
incorporates shells and small found in temperate areas, Growing on other seaweeds in tufts like
snails, this turf-forming are clusters of spore-bearing pom-poms in the northern hemisphere,
red seaweed has flattened, bodies, themselves home this red alga has branching filaments
leaflike fronds. to tiny, parasitic algae. made up of long, tubular cells.
 GREEN ALGAE 105
Green algae are a loose taxonomic that sometimes clog ponds. Some reproduce DOMAIN EUKARYOTA
aggregation of very diverse species. asexually by splitting, budding, or producing

PROTISTS • STONEWORTS AND REL ATIVES

KINGDOM PROTISTA
Some live in freshwater ponds and motile spores, but sexual reproduction is DIVISION CHLOROPHYTA

streams, some form green mats over common in larger species. These typically FAMILIES 127
SPECIES 4,300
damp, shady rocks or tree trunks, produce sperm with two identical flagella,
and others grow as leafy seaweeds, but also have spore-producing stages in their
attached to rocks in shallow sea water. life cycle. The cells of green algae use the 5–8 μm
same chlorophyll pigments as land plants for
Many green algae are microscopic and free- photosynthesis, and they are sometimes included
floating; others are multicellular with variably with them in an informal grouping called the
complex structures, including the blanketweeds Viridiplantae (green plants).

4¾–231⁄2 in 2–16 in 61⁄2–16 ft

12–60 cm 5–40 cm 2–5 m

SEA LETTUCE SEA GRAPE

Ulva lactuca Caulerpa lentillifera
A popular food This edible seaweed
worldwide, this produces many upright VOLVOX AUREUS
green seaweed has stalks from a long, Just visible to the naked eye,
broad, crumpled GREEN SEA FINGERS creeping runner. In the spherical colonies of this
fronds, anchored to Codium fragile Philippines, the freshwater alga consist of
rock by a holdfast. It This tubular green alga occurs succulent stalks are thousands of microscopic
can also form floating worldwide, in seashore rock pools and harvested and individuals. Threadlike flagellae
communities. coastal waters to 6½ ft (2 m) deep. eaten raw in salads. spin the colony through the water.

S T O N E WO RT S A N D R E L AT I V E S
Although related to green algae, the However, the stoneworts and brittleworts DOMAIN EUKARYOTA
stoneworts and their relatives have are the best known group of streptophytes, KINGDOM PROTISTA
more complex structures and more and are often regarded as “honorary plants.” DIVISION STREPTOPHYTA

advanced chemistry in their cells, They grow in the mud of shallow, freshwater FAMILIES 16
SPECIES 2,700
suggesting that they are the ancestors or brackish habitats, rooted by cell filaments,
of the true plants. called rhizoids, like the roots of simple plants. prong
Their branched form and reproductive structures
The Streptophyta (sometimes called Charophyta) are also reminiscent of true plants. In fact, some
includes desmids, which are classed as “conjugating classification systems regard the Streptophyta
350 μm
green algae.” These typically have a single cell, as an “infra-kingdom” that also includes all of
divided into two symmetrical semicells. the land plants.

egg-bearing
capsule CLOSTERIUM SP.
32–70 μm This crescent-shaped
long stem is desmid, found
plantlike stem made of worldwide, displays
12–231⁄2 in elongated cells semicells, each
30–60 cm
housing a chloroplast,
4–20 in
10–50 cm joined at the isthmus,
where the nucleus
is situated. cell
divide

100–460 μm
semicell
NITELLA TRANSLUCENS COMMON STONEWORT PENIUM SP. MICRASTERIAS SP.
This delicate “plant” with translucent Chara vulgaris This desmid, found in N. America, This desmid, found in temperate
green leaves grows in the clear water Also known as skunkweed, this is symmetrically divided into areas, has multiple spined arms,
of ponds, streams, and fens around streptophyte of the northern cylindrical semicells with blunt, tipped with prongs that allow
W. and S. Europe. hemisphere emits an unpleasant smell. oval ends and girdle bands. the semicells to interlock.
plants

108 110
MOSSES LIVERWORTS
Widespread in damp, Related to mosses,
cool, and shaded places, liverworts also have no
mosses are non-flowering flowers, and spread by
plants which have wiry shedding spores instead
stems and small, spirally of making seeds. They
arranged leaves. Many are typically small and
mosses grow on bare ribbonlike, or straggling,
rocks and on trees. with tiny leaves.
 By harnessing the energy in sunlight, and using it to grow,
green plants play a central role in life on Earth. They generate food
for animals and other organisms, and often form their habitats as
well. Some plants are small and simple, but they also include giant
conifers, and a dazzling wealth of flowering species that have evolved
a remarkable array of shapes and strategies for survival.

111 112 116 118 122

HORNWORTS • FERNS AND CYCADS, CONIFERS FLOWERING
LYCOPHYTES RELATIVES GINKGOS, AND Although far fewer in PLANTS
Hornworts are small plants, Ferns are the largest plants GNETOPHYTES species than flowering This is by far the largest
related to mosses, which that reproduce with spores These plants do not have plants, conifers dominate group of living plants,
produce spores from an instead of seeds. Many are flowers, but they do form the landscape in some making up the bulk of the
elongated, hornlike structure. low-growing, but some seeds. Before flowering parts of the world. All of world’s vegetation. They
Lycophytes were previously form trees, which have plants evolved, cycads them are trees or shrubs, all grow flowers—often
thought to belong with the trunks made of fibrous formed an important part and they typically form inconspicuously—and
ferns, and include clubmosses. roots rather than wood. of the world’s vegetation. seeds in woody cones. spread by forming seeds.
108 MOSSES
Mosses are nonflowering plants that typically form mats or cushion-shaped
PLANTS • MOSSES

clumps. Despite their small size, they are remarkably resilient plants. They
are able to grow in a wide range of habitats, from woodland to deserts, and
are found on every continent, including Antarctica.

DIVISION BRYOPHYTA Mosses have thin leaves that usually spiral Because moss spores are so small and light,
CLASSES 8 around slender, wiry stems, and they reproduce they are carried large distances by the slightest
ORDERS 30 by scattering spores. Like liverworts, they need breeze. This makes mosses very good at
FAMILIES 110
SPECIES About 10,000
moist conditions to grow. They can be extremely dispersing and enables them to colonize all
abundant in habitats that are always damp, and kinds of microenvironments, from crevices
some species—particularly sphagnum mosses— in the bark of trees to damp walls and roofs.
form extensive soggy blankets, dominating
the ground in cold parts of the world. Others, r e l at i v e s a n d l o o k- a l i k e s
however, are able to remain dormant during The division name Bryophyta is sometimes used
droughts. They look gray and lifeless, but become to encompass the liverworts and hornworts
green again within minutes if it rains. (see pp.110–111), but these are now regarded as
Mosses have a two-stage life cycle—a separate, but closely related, taxonomic divisions.
feature shared by all other plants. In mosses, Clubmosses are more complex than true mosses,
the dominant stage is the gametophyte, which and belong in the division Lycophyta (see p.111).
produces male and female cells. After the Other moss look-alikes include reindeer moss—
female cells are fertilized, they produce the actually a form of lichen—and Spanish moss,
spore-forming stages or sporophytes, which a flowering plant in the bromeliad family, that
remain attached to the parent plant. Most grows festooned on trees.
moss sporophytes have a capsule on the end
of a long stalk. Once ripe—a process that can
In the far north sphagnum mosses form DEEP COVER
raised bogs—here interspersed with gray take many months—the capsule splits open, The cool, moist climate in New Zealand’s Fjordland National
patches of reindeer moss lichen. releasing up to 50 million spores into the air. Park enables a wide variety of mosses and liverworts to thrive.
 1 1/4 in/3 cm 1 1/4 in/3 cm 3/4 in/2 cm VELVET FEATHER-MOSS spore capsule
Brachythecium velutinum with a conical lid
f: Brachytheciaceae 109
With its much-branched stems,
this abundant moss forms
creeping mats on dead wood and 4 in

PLANTS • MOSSES
in poorly drained grass. It has 10 cm
a worldwide range.

BLACK ROCK MOSS FIRE MOSS COMMON

Andreaea rupestris Ceratodon purpureus POCKET MOSS
f: Andreaeaceae f: Ditrichaceae Fissidens taxifolius
Widespread on mountains, Found worldwide, particularly f: Fissidentaceae
this dark-colored moss grows on burned or disturbed ground, This widespread moss has
on bare rock. Unlike most this low-growing moss is also short, spreading stems
mosses, its capsules release common on roofs and walls. It bearing two ranks of pointed
their spores through four produces a dense carpet of leaves. It grows in shaded
microscopic slits. spore capsules in spring. ground and on rocks.

6 in/15 cm 2 in/5 cm

2 in/5 cm

WHITE FORK MOSS MOUNTAIN FORK MOSS COMMON

Leucobryum glaucum Dicranum montanum TAMARISK MOSS
f: Dicranaceae f: Dicranaceae Thuidium tamariscinum
Growing in large, neatly A moss that forms compact f:Thuidiaceae 32 in
rounded cushions, this feathery mounds, this lowland Its finely divided “fronds” make 80 cm
woodland moss has a species has narrow leaves that this woodland moss resemble
characteristic gray-green curl up when dry, sometimes a miniature fern. It grows on
hue, turning almost white breaking off to form rotting wood and rocks in
in dry weather. new plants. Europe and N. Asia.

1½ in
1 1/4 in/3 cm 4 cm
PURPLE FORK MOSS
Grimmia pulvinata
f: Grimmiaceae
1 1/4 in 4 in
Growing on rocks, roofs, 3 cm 10 cm
and walls, this widespread
moss has leaves that end CYPRESS-LEAVED OSTRICH-PLUME GREATER WATER-MOSS
in long, silvery hairs. Its PLAIT MOSS FEATHER MOSS Fontinalis antipyretica
spore capsules grow on Hypnum cupressiforme Ptilium crista-castrensis f: Fontinalaceae
curved stalks. f: Hypnaceae f: Hypnaceae A submerged freshwater
SWAN’S NECK THYME MOSS This highly variable Found mainly in northern species, greater water-moss
Mnium hornum mat-forming moss has forests, this moss has trails from rocks in slow-
spore capsule is f: Mniaceae crowded, overlapping leaves. feather-shaped, symmetrically flowing rivers and streams.
horizontal when ripe Found worldwide, it is branched stems, often It has three rows of keeled,
Brilliant green in spring, this is
a common woodland moss in common on rocks and walls, forming extensive patches dark green leaves.
Europe and N. America. Each spore and beneath trees. below spruces and pines.
capsule has a curved stalk
resembling a swan’s neck. 16 in ½ in
40 cm 1.5 cm

10 in
25 cm
BLUNT-LEAVED COMMON BONFIRE MOSS
BOG MOSS HAIR-CAP MOSS Funaria hygrometrica
Sphagnum palustre Polytrichum commune f: Funariaceae
CAPE f: Sphagnaceae f: Polytrichaceae One of the world’s most
THREAD-MOSS Like its relatives, this This tall, tussock-forming widespread mosses, this
Orthodontium lineare peat-forming moss grows moss is common in wet mat-forming species is
f: Bryaceae on wet ground, and holds large moorland throughout the particularly common on
⅜ in/1 cm
This moss from the southern hemisphere was amounts of water. Each stem northern hemisphere. Its stems disturbed ground and sites
introduced to Europe in the early 20th century ends in a flat-topped rosette are stiff and unbranched, with of recent fires. Its ripe spore
and has become an invasive species. of small branches. narrow, pointed leaves. capsules have orange stalks.
110 L I V E RW O R T S
Found mainly in damp, shaded habitats, liverworts are thought to be the simplest of all the existing
PLANTS • LIVERWORTS

groups of land plants. They come in two distinct forms: some are flat and ribbonlike; others are more
like mosses, with slender stems flanked by tiny leaves.
DIVISION MARCHANTIOPHYTA Ribbonlike liverworts look unlike any other gradually fragment as they grow, forming
CLASSES 3 plants. Instead of stems and leaves, they have multiple plants. In addition, some species have
ORDERS 16
FAMILIES 88
a flat body, or thallus, that repeatedly forks as cell clusters, called gemmae, in depressions
SPECIES About 7,500 it grows. Many species have a glistening upper on their upper surface. Raindrops scatter
surface with deeply divided lobes. Medieval these gemmae, which grow into new plants.
herbalists noted this liverlike shape, giving Liverworts also reproduce by scattering
D E B AT E the plants their common name. Leafy liverworts spores, microscopic cells that form new plants.
P L A N T S A PA RT look quite different, with trailing or spreading Spore formation requires damp conditions,
Liverworts have unique features that
stems. They usually have two ranks of leaves, because a liverwort’s female cells must first
distinguish them from mosses and with a third row of smaller leaves on their be fertilized by swimming male sperm. Sperm
hornworts, although they were once underside. Some kinds form loose mats in damp cells often reach egg cells with the help of
all grouped together as bryophytes. grass, and others grow on rocks. They are much raindrops, which splash them from one plant
They are the only land plants without more numerous than ribbonlike liverworts, to another. Many liverworts produce their
stomata, or breathing pores, in their
sporophyte (spore-producing) stage.
especially in the tropics, where many species sperm or egg cells in structures that look
Also, their hairlike “roots” are live epiphytically on rainforest trees. like tiny parasols. After fertilization, the
single-celled. They are therefore developed spores are scattered into the air.
classed in their own division, spr ea di ng t h ei r k i n d
Marchantiophyta. Unlike most flowering plants, liverworts have
no preset limits to their growth. They can female reproductive
structure with
form patches up to several yards across, which star-shaped rays

GREATER CRESCENT-CUP LIVERWORT

WHIPWORT gemma cup Lunularia cruciata
Bazzania trilobata f: Lunulariaceae
f: Lepidoziaceae Common in gardens and greenhouses,
A native of damp woodland, this ribbon-shaped liverwort is bright
this mound-forming green, with distinctive reproductive
liverwort has main leaves structures, gemma cups, that look
that overlap and curve like tiny fingernails.
downward, giving its stems
a caterpillarlike shape.

two ranks of OVERLEAF PELLIA

main leaves Pellia epiphylla
f: Pelliaceae
Growing on wet peat and rocks, this
ribbon-shaped liverwort often forms
a tufted mat. It produces black spore
capsules on slender white stalks.

glossy thallus
GREATER FEATHERWORT EVEN SCALEWORT COMMON LIVERWORT
Plagiochila asplenioides Radula complanata Marchantia polymorpha
f: Plagiochilaceae f: Radulaceae f: Marchantiaceae
A delicate liverwort with translucent leaves, Ranging in color from light green In spring and summer this ribbon-shaped
this species forms spreading, mosslike tufts. to brown, this scaly-leaved liverwort liverwort produces conspicuous spore-forming
It grows on shaded rocks and soil, particularly creeps over flat, unshaded surfaces as male reproductive structures that look like tiny parasols. It is
on chalk and limestone. varied as tree trunks and coastal rocks. structures widespread in damp habitats and gardens.
H O R N WO RT S 111

Hornworts are the third division in the in diameter. In their sporophytes, the spores are DIVISION ANTHOCEROTOPHYTA

PL A NTS • LYCOPHY TES

informal grouping called bryophytes. released from a characteristic, elongated hornlike CLASSES 2

They are small, easily overlooked, leafy structure, which gives them their name. Some ORDERS 5
FAMILIES 5
plants found in damp places worldwide. species grow as tiny weeds on the soil of gardens SPECIES About 220
or cultivated fields. Other, rather larger, tropical
Like mosses and liverworts, hornworts have and subtropical species grow mainly on the bark
two generations in their life cycle: a vegetative of trees. These hornworts do not produce hornlike
sporophyte
gametophyte and a spore-bearing sporophyte. flowers, unlike the small order of flowering
The gametophytes form flattened, leafy lobes, water plants that, confusingly, are also
rather like some liverworts, up to 2 in (5 cm) called hornworts (see p.124).

leafy
gametophyte
DENDROCEROS FIELD HORNWORT CAROLINA HORNWORT
Dendroceros sp. Anthoceros agrestis Phaeoceros carolinianus
f: Dendrocerotaceae f: Anthocerotaceae f: Notothyladaceae
The ribbonlike lobes of dendroceros gametophytes Damp stubble fields, trampled ground, and the sides Growing in similar places to field hornwort, this species
spread across damp rocks or tree bark in tropical and of ditches are home to this inconspicuous hornwort has separate male and female gametophytes. Raindrops
subtropical forests. The hornlike sporophyte with distinctive, frilly leaf lobes. It is found across carry sex cells from leaf pits on males to fertilize
grows from this, up to 2 in (5 cm) tall. temperate Europe and N. America. females, allowing sporophytes to develop.

LY C O P H Y T E S
Although their taxonomy is still under nutrients around their roots, stems, and leaves. DIVISION LYCOPHYTA

review, lycophytes include clubmosses This allows lycophytes to grow taller and survive CLASSES 1

and quillworts, plants which are closely in drier habitats than bryophytes. Fossil lycophytes ORDERS 3
FAMILIES 3
related to, but distinct from, ferns. date back 425 million years. They were dominant SPECIES About 170
plants during the Carboniferous era. Some then
Lycophytes are the first plants in this book that were tree-sized, unlike their smaller modern
have a vascular system, a network of conducting relatives. Lycophytes spread by spores, produced in
tissues that transport water, minerals, and sporangia (spore cases) at the base of simple leaves.

STAGHORN CLUBMOSS
Lycopodiella cernua
f: Lycopodiaceae
Lycopodiella species are
generally more delicate than
other clubmosses. Looking
like a miniature pine tree,
staghorn clubmoss grows
in bogs throughout
the tropics.

IBERIAN QUILLWORT INTERRUPTED CLUBMOSS HICKEY’S TREE-CLUBMOSS

Isoetes longissimi Lycopodium annotinum Lycopodium hickeyi
f: Isoetaceae f: Lycopodiaceae f: Lycopodiaceae
Quillworts grow in slow-moving This clubmoss of northern moors Native to hardwood forests and scrub
streams and clear ponds. They have and mountains produces spores in eastern N. America, this species
densely tufted, hollow leaves, with spore from conelike clusters at the spreads by underground stems which
cases hidden in their swollen bases. tip of its upright stems. produce treelike, fertile fronds.
112 F E R N S A N D R E L AT I V E S
Most ferns can be recognized by their graceful fronds, which unroll as they grow. Together
PL A NTS • FER NS AND REL ATIVES

with horsetails and whisk ferns, they make up a diverse and ancient group of nonflowering
plants that reproduce by means of spores. Ferns grow in a wide variety of habitats,
although the majority thrive where there is moisture and shade.

DIVISION POLYPODIOPHYTA Some ferns are small and easily overlooked, but the Tiny, flat, and paper-thin, this eventually produces
CLASSES 1 group also includes large, treelike species that can a new sporophyte—the spore-forming plant we
ORDERS 11 be over 50 ft (15 m) high. Many are compact plants, recognize as a fern.
FAMILIES 40
SPECIES About 12,000
with a single cluster of fronds, but others have
creeping stems that sprout fronds as they spread. f e r n r e l at i v e s
Bracken fern—one of the most widespread Fern taxonomy is in a state of flux, but all the
D E B AT E creeping species—is particularly vigorous. Over scientific evidence suggests that ferns are more
LIVING FOSSILS many years, it can produce clumps, or clones, closely related to seed plants (gymnosperms and
over ½ mile (800 m) across. Ferns also include angiosperms) than to the lycophytes with which
Horsetails have been around for over
300 million years. One of the largest
species that live in fresh water and a wide variety they were once grouped.
extinct genera, called Calamites, of epiphytic species, which live on other plants. The fern division also includes horsetails, which
grew up to 65 ft (20 m) tall, and had Fern fronds are often finely divided, and are upright plants with hollow, cylindrical stems,
massive ridged stems over 231 ⁄ 2 in usually develop from tightly coiled buds, known surrounded by whorls of slender branches. Silica
(60 cm) across. The 20 or so species as crosiers or fiddleheads. Spores are formed granules give them a rough texture, and they were
of horsetails that survive today are
much smaller plants, growing in water,
on the underside of the fronds, in structures once used for scouring pots and pans. Adder’s
woodland, and disturbed ground. that look like raised dots or lines. In many fern tongues and whisk ferns are distinctive groups,
They look very different from ferns, species, all the fronds produce spores, but some with twig-shaped stems or a single, divided
producing their spores in conelike have two kinds of fronds: fertile fronds form frond, that are also classed within the ferns.
structures at the tips of some of spores, while sterile fronds are primarily adapted
their stems, but most botanists now
agree that they should be classified
to collecting sunlight so the fern can grow.
CHARACTERISTIC COIL
as an order within the ferns. When a fern spore germinates, it develops into The delicate tips of young fern fronds are protected inside tight
an intermediate stage known as a gametophyte. coils. These unwind as they stretch upward toward the light.
HOR SE TA I L S 32 in/80 cm W H I SK F E R NS , MO ON WORT S ,
A N D A DDE R’ S TONGU E S 113
COMMON
COMMON HORSETAIL MOONWORT
Equisetum arvense

PL A NTS • FER NS AND REL ATIVES

Botrychium lunaria
f: Equisetaceae f: Ophioglossaceae
Widespread in the northern hemisphere, Found worldwide in
this horsetail can be a troublesome weed. temperate regions,
Its black underground rhizomes sprout moonwort has a single
hollow shoots, bearing symmetrical vegetative frond and a
rings of bright green branches. fertile frond topped
by a cluster of
spore capsules.
T RU E F E R NS 23½ in
60 cm
12 in
30 cm

slender, equally 8 in
spaced fronds 20 cm
4 in/10 cm WHISK FERN ADDER’S TONGUE FERN
Psilotum nudum Ophioglossum vulgatum
f: Psilotaceae f: Ophioglossaceae
A primitive relative of true This unusual fern has a single,
ferns, this largely tropical plant oval frond, clasping a slender,
has leafless, brushlike stems, spore-producing stalk. It grows lobed
bearing berrylike capsules that in grassy places throughout the vegetative
produce its spores. northern hemisphere. frond

4 in/15 cm

PILLWORT creeping WATER CLOVER

Pilularia globulifera stem Marsilea quadrifolia
f: Marsileaceae f: Marsileaceae
This marshland fern from W. Europe With its four-lobed fronds, this clump-
forms clumps resembling tufts of grass. forming aquatic fern looks deceptively like
Its spores develop at ground level, in a flowering plant. It is widely distributed
green capsules shaped like pills. across the northern hemisphere. ½ in 3 1/4 ft
1.5 cm 1m

5 ft ¾ in MOSQUITO FERN UMBRELLA FERN

1.5 m 2 cm
Azolla filiculoides Sticherus cunninghamii
f: Salviniaceae f: Gleicheniaceae
ROYAL FERN FLOATING FERN A floating plant with This distinctive fern from New
Osmunda regalis Salvinia natans mat-forming fronds, Zealand has a pencil-thin stem
f: Osmundaceae f: Salviniaceae mosquito fern spreads topped by a horizontal crown
Often cultivated, this Often forming a dense rapidly across lakes and of narrow, radiating fronds. It
stately fern from the carpet, this water fern has ponds. It is widespread in spreads by creeping rhizomes.
northern hemisphere has a small, oval leaves covered warm parts of the world.
rosette of spreading fronds with water-repellent 33 ft
with several narrower, hairs. It is common 10 m
spore-bearing fronds throughout the tropics.
in the center.
SILVER TREE FERN
Alsophila dealbata
f: Cyatheaceae
This tree fern gets its name
from its fronds, which have
a silvery underside. Native
to New Zealand, it grows
in open forest and scrub.
20 ft
6m

BLACK TREE FERN SOFT TREE FERN

TASMANIAN Sphaeropteris medullaris Alsophila smithii
TREE FERN 59 ft
f: Cyatheaceae 26 ft
f: Cyatheaceae
Dicksonia antarctica 18 m From New Zealand, this 8m Native to New Zealand and
fronds are evergreen
f: Dicksoniaceae tall and slender tree fern sub-Antarctic islands, this
in mild climates This stout-stemmed tree fern is has a sooty-black trunk is the most southerly tree
widespread in Tasmania and S.E. and dark leaf stalks, fern. It often has a collar
Australia. It needs shady conditions, contrasting with its bright of dead fronds hanging
and usually grows among trees. green fronds. beneath its crown.
 T RU E F E R NS BLACK
MAIDENHAIR FERN
12 in
30 cm
114 Adiantum capillus-veneris
f: Pteridaceae
Found in crevices on
limestone, this widespread
PL A NTS • FER NS AND REL ATIVES

long fern has bright green,

terminal translucent leaflets that
leaflet contrast with its slender
black stems.

20 in
50 cm
23½ in
60 cm
3¼ ft
1m
10 in
purple stem 25 cm
and veins
SILVER
LIP FERN
SILVER BRAKE PAINTED BRAKE CLIFF BRAKE
Cheilanthes argentea
Pteris argyraea Pteris tricolor Pellaea viridis f: Pteridaceae
f: Pteridaceae f: Pteridaceae f: Pteridaceae From E. Asia, this
This shade-loving fern is From Malaysia, painted brake has an Native to South Africa, this drought- small, evergreen fern
named for its distinctive unusual range of colors: its metallic- resistant fern has shining green fronds with has wedge-shaped
fronds, which have a looking, copper-colored young fronds wiry, black stems. It grows in open fronds with black
silvery white central turn green as they mature. woodland on mountains. veins and distinctive
stripe. Originally from silvery markings on
S.E. Asia, it is widely their underside.
grown as a houseplant. LADDER BRAKE 3¼ ft
Pteris vittata 1m
f: Pteridaceae
Widespread in warm regions, this fern
has upright or arching fronds with
narrow, linear leaflets. It usually grows
16 in on limestone and in alkaline soils.
40 cm

6 ½ ft
2m

30 in
75 cm

NORTHERN BEECH FERN MOUNTAIN FERN

Phegopteris connectilis Oreopteris limbosperma BRACKEN FERN purplish-black
f:Thelypteridaceae f:Thelypteridaceae Pteridium aquilinum stalk
Found as far north as Greenland, this Found in damp habitats on acid soil, f: Dennstaedtiaceae
compact fern grows in a variety of this clump-forming European fern Found on every continent except Antarctica,
habitats from woodland gives off a characteristic lemony bracken spreads vigorously by growing
to rocky tundra. smell if its fronds are bruised. underground rhizomes. It often dies back
in winter, sprouting new fronds in spring.

20 in 6 ft 30 in
50 cm 1.8 m 75 cm
SQUIRREL’S EUROPEAN DEER
FOOT FERN CHAIN FERN FERN
Davallia trichomanoides Woodwardia radicans Blechnum spicant
f: Davalliaceae f: Blechnaceae f: Blechnaceae
This epiphytic fern from Malaysia Native to S.W. Europe and This evergreen fern has spreading
creeps over trees and other Algeria, this is a luxuriant fern of sterile fronds, and upright fertile
plants. Its rhizomes have furlike damp, shaded habitats. Its arching fronds with leaflets like narrow ribs.
scales and tips that resemble fronds sometimes produce bulbils It is native to the temperate
a squirrel’s foot. (small bulbs) at their tips. northern hemisphere.
 SOFT SHIELD FERN
Polystichum setiferum
f: Dryopteridaceae
Native to damp European
woodlands, soft shield fern has
feather-shaped fronds, with
leaflets turned at an angle,
instead of lying flat.

16 in
40 cm
OAK FERN
4 ft Gymnocarpium dryopteris
1.2 m f: Cystopteridaceae
Delicate bright green fronds, sprouting
separately, distinguish this northern fern. It
MALE FERN grows in shaded screes as well as in woodland.
Dryopteris filix-mas
f: Dryopteridaceae
One of the most common woodland
ferns in Europe, this species has a BRITTLE
crown of concentric fronds shaped like BLADDER-FERN
the tail of a shuttlecock. Cystopteris fragilis
f: Cystopteridaceae
Named for its swollen spore
capsules borne on the underside
4 ft of its fronds, this fern grows in
spore capsules 1.2 m temperate regions worldwide.
on underside
of fronds

23½ in
60 cm
16 in
40 cm

fronds form
cone-shaped
crown

winged
23½ in midrib 5 ft
60 cm toothed, pale 1.5 m
green leaflets

SENSITIVE FERN
Onoclea sensibilis
f: Onocleaceae
The fronds of this wetland fern OSTRICH FERN
from N. America and E. Asia seem Matteuccia struthiopteris
very sensitive to cold, dying soon f: Onocleaceae
after the first frosts. Native to the northern hemisphere,
this tall waterside fern has a
symmetrical crown of sterile fronds
in summer, followed by brown
6 in
15 cm fertile fronds in the winter. 12 in
WALL RUE 30 cm
Asplenium ruta-muraria
f: Aspleniaceae
Widespread across the
northern hemisphere, this
small tuft-forming fern
HART’S TONGUE FERN grows on limestone rocks
Asplenium scolopendrium and on walls containing
f: Aspleniaceae lime-rich mortar.
With its glossy, strap-shaped
fronds, this fern is often cultivated
as an ornamental plant. It grows
8 in wild in Europe, W. Asia,
20 cm and N. America. COMMON
STAGHORN FERN
Platycerium bifurcatum COMMON POLYPODY
MAIDENHAIR SPLEENWORT f: Polypodiaceae Polypodium vulgare
Asplenium trichomanes Found in Indonesia and f: Polypodiaceae
f: Aspleniaceae Australasia, this impressive Instead of forming clumps, this
Occurring from the tropics to subarctic epiphyte grows on tree trunks. fern sprouts separate fronds along
regions, this small clump-forming fern It has kidney-shaped sterile 35 in its creeping rhizome. It is common
grows in rocky places. Its fronds have fronds and spreading fertile 90 cm on rocks, trees, and leaf litter in
paired elliptical leaflets. fronds shaped like antlers. northern temperate regions.
116 C YC A D S , G I N KG O S ,
AND GNETOPHYTES
PLANTS • CYCADS, GINKGOS, AND GNETOPHYTES

Found mainly in warm parts of the world, the cycads, ginkgos, and gnetophytes are three
ancient divisions of nonflowering plants with strikingly varied shapes. They range from
climbers and low-growing shrubs to plants that are easily mistaken for palms.
DIVISION CYCADOPHYTA These three groups are traditionally classed, along trees and climbers from the tropics, densely
CLASS 1 with conifers, as gymnosperms—a word that branched shrubs from dry habitats, and the
ORDER 1 means “naked seed.” Unlike flowering plants, bizarre Welwitschia, which grows only in the
FAMILIES 2
SPECIES 330
gymnosperms form their seeds on exposed Namib Desert of southwest Africa.
surfaces, which are typically specialized scales.
DIVISION GINKGOPHYTA
Flowering plants produce their seeds in closed va ry i n g f o rt u n e s
CLASS 1 chambers, or ovaries. Cycads have a long history, dating back nearly
ORDER 1 Scientists have not yet resolved the exact 300 million years. Once an important part of
FAMILY 1 relationship between cycads, ginkgos, and the world’s vegetation, they have gradually lost
SPECIES 1
gnetophytes. Nor can they say where they stand ground in the face of competition from flowering
in relation to conifers and flowering plants. At plants. Today, nearly a quarter of cycad species
DIVISION GNETOPHYTA
CLASS 1
the cellular level, some features of gnetophytes are endangered, under threat from illegal plant
ORDERS 1 suggest they are more closely related to conifers collection and habitat change.
FAMILIES 3 than to cycads and ginkgos, but they also have Gnetophytes face fewer problems, and the
SPECIES 70 features in common with flowering plants. ginkgo has seen a dramatic improvement in its
Apart from the nature of their seeds, the status. For centuries it was preserved by Buddhist
three groups have little in common, and hardly monks, who grew it in temple gardens long
BUILT TO LAST
Like palms, cycads usually have a single ever grow in the same place. Cycads grow after it had vanished in the wild. Introduced to
crown of compound leaves, surrounding a mainly in the tropics and subtropics, while the Europe in the 1700s, it proved easy to grow and
central growing point, or apical meristem.
Their tough leaves are able to withstand single surviving species of ginkgo comes from highly tolerant of polluted air. It is now planted
strong sunshine and drying winds. China. Gnetophytes are more diverse, including worldwide in parks and city streets.
C YCA D S EASTERN CAPE
BLUE CYCAD
stiff, upswept leaves Encephalartos horridus 117
f: Zamiaceae
Unlike most cycads, this
low-growing semidesert

PLANTS • CYCADS, GINKGOS, AND GNETOPHYTES

species from South Africa has
gray-blue leaves, composed
of stiff leaflets armed with
sharp spines.

4 ½ ft/1.4 m 20 ft/6 m
trunk branches
10 ft/3 m when mature PRICKLY CYCAD
Encephalartos altensteinii
JAPANESE SAGO PALM f: Zamiaceae
Cycas revoluta Found near South Africa’s eastern
f: Cycadaceae coast, this tall, subtropical cycad
Unrelated to true sago, this palmlike spiny leaves with is named for the toothed edges
cycad from S. Japan has a thick stem recurved tips of its leaves. Its yellow cones
and glossy leaves. It is widely cultivated produce bright red seeds.
as an ornamental plant.
23 ft
7m
6 ft 4 ft
1.8 m 1.2 m

10 ft/3 m

6½ ft/2 m

MEXICAN FERN PALM MEXICAN HORNCONE COONTIE MACROZAMIA MOOREI BURRAWONG

Dioon edule Ceratozamia mexicana Zamia pumila f: Zamiaceae Macrozamia communis
f: Zamiaceae f: Zamiaceae f: Zamiaceae One of Australia’s tallest f: Zamiaceae
A cycad rather than a true palm, This robust cycad from E. Mexico Once used as a source of starch, this cycads, this palmlike plant Native to Australia’s southeast
this slow-growing plant from has a large crown of spreading leaves dwarf cycad from the Caribbean has giant seed cones up to coast, this cycad has large cones
E. Mexico has oval seed cones and gray-green cones with distinctive has a short, half-buried stem and 36 in (90 cm) long. It grows with red, fleshy seeds. It often
up to 12 in (30 cm) long. horns on their scales. upright reddish-brown cones. in dry woodland. grows in dense stands.

GNETOPHYTES GI N KG O
MELINJO
LONGLEAF 50 ft/15 m Gnetum gnemon MAIDENHAIR TREE
JOINT-FIR f: Gnetaceae Ginkgo biloba
Ephedra trifurca A nonflowering tree from f: Ginkgoaceae
f: Ephedraceae S.E. Asia and the Pacific,
100 ft/30 m
This distinctive tree is easily
Also known as Mormon melinjo has evergreen leaves recognized by its fan-shaped
tea, this mound-forming plant and nutlike seeds. Both are leaves, which turn bright
has stems ringed by used in cooking. yellow in the fall. Originally
scalelike leaves. It grows restricted to S. China,
in deserts in Mexico and it is now cultivated worldwide.
the southern US.
seeds with
fleshy coat
6½ ft/2 m
shredded leaf

edible
kernels

WELWITSCHIA
18 in/45 cm Welwitschia mirabilis
f:Welwitschiaceae
CHINESE EPHEDRA Endemic to the Namib
Ephedra sinica male cones Desert, this extremely
f: Ephedraceae attached to long-lived plant has a single
This straggling plant from E. Asia central stem pair of strap-shaped leaves.
contains powerful alkaloids Over the centuries they
and has a long history of use crack and split, creating a
as a medicinal herb. 3¼ ft/1 m tangled mound of foliage.
118 CONIFERS
Conifers evolved more than 300 million years ago, long before the world’s
PLANTS • CONIFERS

first broad-leaved trees. With their tough waxy leaves, conifers thrive in harsh
climates. They are less diverse than broad-leaved trees, but they dominate
forests on cold mountains, and in the far north.

DIVISION PINOPHYTA Although relatively few in species, conifers wither soon after shedding their pollen into the
CLASS 1 include the world’s tallest trees, as well as the air. The larger female cones contain one or more
ORDER 2 heaviest, the longest-lived, and some of the seeds. They can take several years to develop, and
FAMILIES 6
SPECIES About 600
most widespread. Conifers are traditionally usually become hard and woody when mature.
placed in a group known as gymnosperms, The cones of some species, including firs
along with cycads, ginkgos, and gnetophytes. and cedars, release their seeds by slowly
Unlike broad-leaved trees, they do not have disintegrating on the tree. Pine cones remain
flowers and produce pollen and seeds in cones. intact, and often remain on the tree long after
they are ripe. Most of them shed their seeds
l e av e s a n d c o n e s in dry weather, when their scales open wide,
Most conifers are evergreen, with highly but some hold onto their seeds until they are
resinous leaves that are good at withstanding scorched. This is a specialized adaptation that
cold winds and strong sunshine. Pines have allows pines to recolonize areas that have been
slender needles, arranged singly or in bundles, burned by forest fires.
while other conifers often have linear leaves A few conifers, the yews, junipers, and
or flat scales. A few conifers—including larches podocarps, have small, berrylike cones with
and some redwoods—are deciduous, with soft fleshy scales that do not fully enclose the seed.
leaves that are shed each year. These are scattered by birds as they feed.
Conifers have two types of cones, which
normally grow on the same tree. Male cones
Between areas of open tundra, conifers CONIFERS IN CONTROL
grow far to the north of the Arctic Circle, produce pollen. Small, soft, and often very Conifers often form extensive single-species stands. These pines
forming the largest forest on Earth. numerous, they typically appear in spring, and are growing in California’s Yosemite National Park in California.
 115 ft 245 ft/75 m 200 ft/60 m
35 m 119

PLANTS • CONIFERS
cone

four-sided 260 ft/80 m

rigid leaves
COLORADO BLUE SPRUCE GRAND FIR CALIFORNIA RED FIR EUROPEAN SILVER FIR
Picea pungens Abies grandis Abies magnifica Abies alba
f: Pinaceae f: Pinaceae f: Pinaceae f: Pinaceae
Bright blue-gray leaves, with The giant fir is a tall, fast-growing This drought-resistant fir is Named after the silver bands on the
prickly tips, make this a popular tree that is native to western found on dry mountain slopes underside of its leaves, this fir has
ornamental tree. Native to N. America. Its leaves have a in California. It has up-curved upright resinous cones. They disintegrate
western N. America, it typically citruslike scent when crushed. leaves and upright cones in order to scatter their seeds.
grows on mountains. up to 8 in (20 cm) long.

200 ft/60 m

SITKA SPRUCE CAUCASIAN FIR

Picea sitchensis Abies nordmanniana
f: Pinaceae f: Pinaceae
Thriving in cold, wet Originally from mountains in the
115 ft Black Sea region, this is a popular
conditions, the sitka spruce 35 m
is often grown as a forestry Christmas tree in Europe because
tree. It comes from the it retains its needles indoors.
coastal belt of western
N. America. long paired
needles
lower side of leaf 165 ft
has two blue- 50 m
white bands SCOTCH PINE
NORWAY SPRUCE Pinus sylvestris 65 ft/20 m
Picea abies f: Pinaceae
f: Pinaceae Ranging from the British Isles
245 ft/75 m An important lumber tree, to China, the Scots pine is the cone with
edible seeds
this fast-growing conifer has world’s most widespread conifer
spiky leaves and cylindrical after common juniper. Its upper
cones. Its natural range is branches have beautiful
N. and C. Europe. orange-red bark.

LODGEPOLE PINE STONE PINE

Pinus contorta Pinus pinea
100 ft f: Pinaceae f: Pinaceae
cylindrical cone 30 m Growing in coastal Prized for its edible seeds (pine
with toothed scales dunes and bogs, this nuts), this Mediterranean pine
N. American pine has has large, oval cones, and an
50 ft/15 m paired leaves and prickly elegant umbrella-shaped
cones. The cones release outline when mature.
SINGLE-LEAF PINYON their seeds after being
Pinus monophylla scorched by fire.
f: Pinaceae
150 ft
Unique among pines, this species has leaves 45 m
that grow singly, rather than in pairs or
clusters. It comes from rocky slopes in
Mexico and the American Southwest. male pollen-
producing cones

stiff, sharp
needles

130 ft
65ft/20 m 115 ft/35 m 40 m

AROLLA PINE MARITIME PINE AUSTRIAN PINE

Pinus cembra Pinus pinaster Pinus nigra 82 ft/25 m
f: Pinaceae f: Pinaceae f: Pinaceae MONTEREY PINE
Found on European mountains, Native to the W. Tall, rangy, and open-crowned, CHINESE PINE Pinus radiata
this slow-growing tree Mediterranean, the maritime this pine has long leaves Pinus tabuliformis f: Pinaceae
produces small cones that pine grows rapidly on poor arranged in pairs. Despite its f: Pinaceae Originally restricted to a small area
fall intact. Birds such as sandy soils. It has glossy name, it is widespread With age, this oriental pine develops a distinctive of California, this fast-growing pine
nutcrackers feed on the cones brown cones that are up throughout Europe, typically flat, spreading crown. It grows on mountains, is now widely planted for its lumber,
and disperse the seeds. to 8 in (20 cm) long. growing on limestone. and produces small, egg-shaped cones. particularly in the southern hemisphere.
 CONIFERS immature cone
120
male cone
withers after
shedding pollen

200 ft/60 m

DOUGLAS FIR
Pseudotsuga menziesii
f: Pinaceae
One of the world’s tallest
conifers, the Douglas
fir grows in western
N. America. Its cones have
projecting bracts formed
from modified scales.

mature cone
with three-
165 ft/50 m pronged bracts

130 ft
40 m

200 ft
60 m
WESTERN HEMLOCK GOLDEN LARCH
Tsuga heterophylla Pseudolarix amabilis
f: Pinaceae f: Pinaceae
From western N. America, Native to E. China, the
DEODAR this is the largest hemlock. golden larch turns brilliant
Cedrus deodara It thrives in cool, damp yellow in the fall before losing
f: Pinaceae conditions, and can live to its leaves. Its cones break up as
Native to the W. Himalayas, this be over 1,000 years old. they scatter their seeds.
fast-growing cedar has branches
that droop at their tips. Its cones
are purple-brown when ripe.

130 ft/40 m

130 ft 82 ft/25 m
40 m

fragile
ripe cone
98 ft/30 m 165 ft/50 m
JAPANESE
ATLAS CEDAR UMBRELLA PINE
Cedrus atlantica CEDAR OF LEBANON JAPANESE LARCH MONKEY PUZZLE Sciadopitys verticillata
f: Pinaceae Cedrus libani Larix kaempferi Araucaria araucana f: Sciadopityaceae
This N. African cedar f: Pinaceae f: Pinaceae f: Araucariaceae The only member of its
has short needles and A majestic conifer with A deciduous tree—like all Originally from the mountains of family, this ancient
upright barrel-shaped spreading branches, the cedar larches—this has soft leaves and Chile, this primeval-looking tree Japanese species is
cones. When ripe, the of Lebanon is now rare in the persistent woody cones with has spirally arranged, sharply distinguished from other
cones slowly break up to wild, but is widely planted as downcurved scales. It grows pointed leaves. Mature trees conifers by having 10–13
release their seeds. an ornamental tree. on mountains in N. Japan. develop an umbrellalike crown. needles clustered in whorls.
 121

PLANTS • CONIFERS
100 ft
30 m

65 ft
165 ft/50 m 230 ft/70 m 20 m

WESTERN RED-CEDAR LAWSON CYPRESS JAPANESE CEDAR WESTERN JUNIPER

Thuja plicata Chamaecyparis lawsoniana Cryptomeria japonica Juniperus occidentalis
f: Cupressaceae f: Cupressaceae f: Cupressaceae f: Cupressaceae
A large tree with flat sprays Like other cypresses, this has A cypress rather than a true This long-lived tree grows on
of small, scalelike leaves, this small cones and sprays of tiny, cedar, this tree has slender rocky mountain slopes in 82 ft/25 m
species grows in northwest scalelike leaves. Originally from leaves and small, rounded W. US. Like other junipers, it
N. America. Its wood is highly western N. America, it exists in cones. It grows on mountains produces seeds inside
rot-resistant, even when dead. many cultivated forms. in China and Japan. berrylike cones. CHINESE JUNIPER
Juniperus chinensis
f: Cupressaceae
Widespread in temperate
E. Asia, this shrub or small
tree has prickly leaves when
young, and scalelike leaves
312 ft/95 m 130 ft/40 m when mature.

linear leaves in
opposite ranks

aril turns red

when ripe

377 ft
115 m
ripening 82 ft/25 m
cone
GIANT SEQUOIA DAWN REDWOOD MONTEREY CYPRESS
Sequoiadendron giganteum Metasequoia glyptostroboides Cupressus macrocarpa
f: Cupressaceae f: Cupressaceae f: Cupressaceae
COAST This Californian redwood is Native to C. China, this Although widely cultivated,
REDWOOD the world’s most massive tree. deciduous redwood is the wild Monterey cypress
Sequoia sempervirens The largest living specimen is extremely rare in the wild. is restricted to a small region
f: Cupressaceae estimated to weigh 2,200 Until the 1940s it was thought of California’s coast. Mature
Native to the coasts of N. California and tons, with fireproof bark to be extinct, being known trees typically have an
Oregon, this is the world’s tallest tree. more than 2 ft (60 cm) thick. only from fossils. irregular spreading shape.
Mature trees have a soaring trunk and
relatively sparse branches, and can
be as much as 2,000 years old.
66 ft/20 m

245 ft/75 m 130 ft/40 m 98 ft/30 m 66 ft/20 m

TAIWANIA SWAMP CYPRESS CALIFORNIA CHINESE PLUM-YEW EUROPEAN YEW

Taiwania cryptomerioides Taxodium distichum NUTMEG Cephalotaxus fortunei Taxus baccata
f: Cupressaceae f: Cupressaceae Torreya californica f:Taxaceae f:Taxaceae
One of Asia’s largest conifers, Also known as the bald cypress, f:Taxaceae This small, densely branched This long-lived tree produces
this tropical species has a trunk this deciduous conifer grows This rare conifer is restricted to conifer has fleshy cones, which seeds in fleshy arils—modified
up to 10 ft (3 m) thick. It has in swamps in S.E. US. canyons and mountains in California. turn purple-brown when cone scales. Found wild in
spiny-tipped leaves, and small, Its trunk often has a It is unrelated to the true nutmeg, ripe. It grows in mountain Europe and S.W. Asia,
rounded cones. buttressed base. although it has nutlike seeds. forests in C. and E. China. it is commonly planted.
122 F L OW E R I N G
PLANTS
PLANTS • FLOWERING PLANTS

With more than 300,000 species, flowering plants—or angiosperms—

are by far the largest group of plants, as well as the most diverse.
They play a vital role in most land-based ecosystems, producing
food and shelter for animals and many other living things.

DIVISION ANGIOSPERMAE The first flowering plants evolved 140 million years
CLADES 10 ago, which makes them relative latecomers to life on
ORDERS 64 Earth. Since then, they have become the dominant
FAMILIES 416
SPECIES About 304,000
forms of plant life. The smallest flowering plants are
not much bigger than a pinhead, but this group also
includes all broadleaved trees, and a variety of other
species, from cacti and grasses to orchids and palms.
Flowering plants share several key features that
help to make them successful. Foremost among
these are flowers—actually a collection of highly
modified leaves. In most flowers, the outermost layers
are the sepals, then the petals. These surround the
male stamens, which make pollen, and the female
Wind-pollinated flowers such as hazel carpels, which collect pollen arriving from similar
catkins shed clouds of pollen grains into
the air. They are rarely brightly colored. flowers, so that their female ovules can develop into
seeds. Some flowers spread their pollen through the
air, but many more use animals to transfer pollen for
them. In these showy species, flowers lure animals
to sip the sugary nectar in the innermost part of the
flower, brushing past the pollen on the way. These
plants mostly attract insects, but birds and bats are
also important pollinators.
Animal-pollinated flowers are usually
conspicuous, and have sticky pollen grains. s t r at e g i e s f o r s p r e a d i n g
Hummingbirds transfer pollen as they feed. Flowering plants are not unique in making seeds, but
they are the only plants that produce them in fruits.
A fruit develops from a flower’s ovaries—the closed
chambers that house the seeds, close to the flower’s
central stem. Fruits have a double function: to protect
the seeds and to help them spread. Fleshy fruits do
this by attracting animals, which feed and scatter the
seeds in their excrement. Dry fruits work in a variety
Fleshy fruits evolved to attract animals. of ways. Some burst open when their seeds are ripe.
Wild cucumbers are eaten by antelopes, Others have hooks that latch onto clothing, skin, or
which scatter the seeds in their droppings.
fur. Still more drift in water or through the air. Many
flowering plants spread by the way they grow, often
producing new plants from creeping stems. This can
create vast, interconnected clones. The largest examples,
formed by North American aspen trees, stretch over
100 acres (40 ha), and may be 10,000 years old.

Dry fruits often burst open when ATTRACTING ATTENTION

their seeds are ripe. This willowherb is Hellebores use colorful sepals, rather than petals, to attract
scattering its fluffy seeds into the wind. pollinating insects—or multiple sepals, in this garden variety.
flow ering plants grou ps
DNA sequencing is revolutionizing
our understanding of relationships
between flowering plants. The
groupings below represent the
current thinking (see p.150 for
subdivisions within the Eudicots).
ba sa l a n g i o s pe r m s

124
m ag n o l i i d s

128
m on o c o t s

130
eu dicots

150
124 BA SA L ANGIOSPER M S
Of the more than 60 orders of flowering in widely scattered parts of the world. They DIVISION ANGIOSPERMAE

plants, or angiosperms, three, known include trees, shrubs, and climbers, as well as GROUPS BASAL ANGIOSPERMS
FLOWERING PLANTS • BASAL ANGIOSPER MS

ORDERS 3
as basal angiosperms, evolved early on water plants. The Amborellales order survives
FAMILIES 7
and still exist today. today as a single species of shrub on one South SPECIES 190
Pacific island. The Nymphaeales are aquatic
Modern DNA sequencing is a powerful way of plants with primitive but often showy flowers.
identifying groups of species that are closely related They include more than 70 species of water lilies, D E B AT E
to one another. However, the interrelationship found all over the globe. The Austrobaileyales MYSTERIOUS ORIGINS
between these groups is sometimes less clear, with order contains nearly 100 species of woody Angiosperms may have evolved from
different DNA markers suggesting different affinities. plants, mainly in the tropics. seed ferns, a group that died out over
In flowering plants, there seems to be a pattern of Two other orders feature on this page as 50 million years ago, or more probably
some flourishing groups that have diversified hugely, “orphans,” whose place in the phylogeny from gnetophytes (see p.117). DNA
leaving behind a few, remnant groups that have (evolutionary history) of angiosperm families and fossil evidence suggest that the
survived for millions of years but diversified little. is unclear. The most recent review suggests that Amborellales were the first angiosperms
“Basal angiosperm” is the term used for three the Chloranthales might be related to magnoliids, to appear, about 140 million years ago.
The Nymphaeales and Austrobaileyales
of these remnant orders. They are primitive and the Ceratophyllales to eudicots, but for now were the next two groups to evolve.
species, apparently with little in common, growing that remains uncertain and unproven.

CHLORANTHALES 6½ ft
2m
AMBORELLALES
The Chloranthaceae, the only family in the order This order of primitive evergreen shrub contains
Chloranthales, contains four genera and some 70 living a single family, which has a single genus with only
species, but its fossil record dates back over 100 million one species, Amborella trichopoda. It bears small
years. Its members are mainly flowers, males and females being produced on
tropical shrubs and trees separate plants, and red berries, each containing
with small, stalkless flowers. SARCANDRA GLABRA
f: Chloranthaceae a single seed.
With a range of medicinal uses,
5 ft this evergreen shrub inhabits
1.5 m damp ground, especially AMBORELLA TRICHOPODA
wooded banks of streams, in f: Amborellaceae
S.E. Asia, China, and Japan. Found only on the island of New Caledonia in the
South Pacific, this sprawling, white-flowered shrub
berry clusters is quite common in mountain forests.
in winter

AU S T RO BA I L E YA L E S
The order Austrobaileyales is made up of only three
families. Members of this order are trees, shrubs, and
climbers. The flowers of most species are single and stem, leaves,
have many petals. Perhaps the best known species is and flowers
the spice, star anise.

many-petaled
single flower

C E R ATOPH Y L L A L E S fruit
59 ft/18 m
A single family, the Ceratophyllaceae, makes up this STAR ANISE
order. Called hornworts, these plants are completely Illicium verum
unrelated to the nonflowering hornworts f: Schisandraceae
(see p.111). The four species are free- Widely used for flavoring,
star anise fruits are woody and
floating, rootless aquatics with whorls star-shaped. This woodland
of finely-divided leaves, tiny male and species is native to China
female flowers, and spiny fruits. and Vietnam.

3¼ ft RIGID HORNWORT
1m Ceratophyllum demersum AUSTROBAILEYA SCANDENS
f: Ceratophyllaceae f: Austrobaileyaceae
This is a submerged species with no The flowers of this rare primitive
roots, inhabiting ponds and ditches 50 ft/15 m climber, found only in rainforests in
in non-arctic Europe. It has tiny Queensland, Australia, smell of
flowers and whorls of leaves. rotting fish to attract pollinating flies.
NYMPHAEALES
This primitive order encompasses three families of star-shaped 125
semidouble flower
aquatic plants with floating, submerged, or, more
rarely, emergent leaves. The family Nymphaeaceae

FLOWERING PLANTS • BASAL ANGIOSPER MS

includes water lilies, which are grown in ornamental
ponds throughout the world for their showy flowers.

10 ft/3 m flower

AMAZON WATER
LILY
Victoria amazonica
f: Nymphaeaceae
Native to deep Amazonian
backwaters, this water lily
has enormous round leaves 6½ ft/2 m
with upright rims. The
rimmed leaves flowers open at night.

CAROLINA NYMPHAEA ‘SUNRISE’

FANWORT f: Nymphaeaceae
Cabomba caroliniana With large, showy, fragrant
f: Cabombaceae flowers and mid-green
From still, freshwater floating foliage, this hybrid,
habitats in US and S.E. thought to be of American
South America, this origin, is one of the largest
species has submerged flowering water lilies.
and floating leaves. It
can be invasive.
20 in
50 cm
WHITE WATER LILY
Nymphaea alba
f: Nymphaeaceae
This European species has
fragrant flowers and its fruits
ripen under water, releasing 5 ft/1.5 m
floating seeds. It inhabits lakes,
ponds, and slow-flowing rivers.
starlike, pure
white flower
PRICKLY FOXNUT
Euryale ferox
f: Nymphaeaceae
This inhabitant of deep,
slow-flowing water and still
water in parts of Asia has 5 ft/1.5 m
prickly leaves, flowers, and
many-seeded berries.

sharp prickles
protect stems,
flowers, and floating young stems, fruits,
leaves from animals and seeds are
all edible

bright purple
flower

grass-green to
olive-green shoots
 FLOATING FLOWERS
Water lily flowers are hermaphrodite—they have
126 both male and female organs. Pollen from the male
parts can fertilize ovules in the female reproductive
parts, so they can “self-pollinate.” However, female
parts mature slightly before the flower starts
releasing pollen. This increases the chances of
flowers being fertilized by pollen carried by insects.

LEAVES
The water lily has large leaves, up to 14 in
stamen (35 cm) in diameter. Unlike most leaves, their
stomata (breathing pores) are on the upper furled
surface, which has a water-repellent coating. young leaf
petal

FLOWER
The flowers have numerous
white petals and bright yellow aerenchyma
sepal stamens, and can grow to
8 in (20 cm) in diameter.

seeds
STEM SECTION
Stems have longitudinal air spaces
OVARY SECTION (aerenchyma), as well as structural
The female reproductive tissue, creating buoyancy and
organs (consisting of allowing oxygen to circulate.
ovules, ovary, and stigma)
are fused together. Cavities
contain ovules, which turn
into seeds when fertilized.

W H I T E WA T E R L I LY
Ny mphaea alba
One of about 75 wild species of water lily, this handsome plant lives in still
or slow-flowing water, casting a deep shade with its rounded, glossy leaves.
It grows in water up to 5 ft (1.5 m) deep, and flowers in mid- to late summer,
producing a succession of pure white blooms. Each flower lasts for three to
four days, opening in the morning and closing by late afternoon. They attract
pollinating beetles, which often spend the night inside the flowers, before
being released at dawn. White water lilies are useful to water animals: pond size Leaf diameter 4–14 in (10–35 cm)
snails glue their eggs to the underside of the leaves, and fishes hide beneath habitat Ponds, streams, lakes, backwaters
them, avoiding predatory birds. After the flowers are pollinated, they produce distribution Europe
leaf type Simple, orbicular with a basal notch
buoyant seeds. These float for several weeks, before sinking into the mud.
 flower bud enclosed
in protective sepals
127
flower supported
by long, thick stalk large floating leaf maximizes the
surface area, in order to capture
sunlight for photosynthesis

fibrous root

buoyant stem (or petiole)

enables the leaf to reach
the top of the pond

folded petal

stamen (male part

that produces and
releases pollen)

sepal

inner petal

stigma (female part

that collects pollen)

ovary (containing
ovules which,
when fertilized,
become seeds)

ROOT SYSTEM INSIDE THE BUD

The small, fibrous root system is This cross section shows the
embedded in the mud. Its primary reproductive organs of the water
purpose is to soak up water and lily. Each long, pointed bud has
absorb oxygen, while also helping four to five pale green sepals
to anchor the plant. which enclose the flower.
128 M AG NO L I I D S
The magnoliids are a large group of in color, size, and shape, known as tepals. The DIVISION ANGIOSPERMAE
primitive flowering plants that in fossil record shows that similar flowers existed
FLOWERING PLANTS • MAGNOLIIDS

CLADE MAGNOLIIDAE

botanical terms are between basal more than 100 million years ago. ORDERS 4
FAMILIES 18
angiosperms and monocots. SPECIES 10,000
c o m m o n c h a r ac t e r i s t i c s
Found in tropical and temperate regions, the Magnoliids are grouped together mainly by genetic
magnoliids form a major plant group that emerged evidence, but they also share features that can D E B AT E
early in the history of flowering plants. They be seen with a microscope, or with the naked PIONEERING PLANTS
take their name from the magnolia family, which eye. At a microscopic level, their pollen grains
is one of the largest in the group. Plants in this have a single pore. This small but significant Scientists have differing opinions
family nearly all have woody stems, and some feature links them with monocots, but about the growth habit, or shape and
lifestyles, of the first flowering plants.
kinds grow into large trees. However, the group distinguishes them from eudicots—by far
According to the “woody hypothesis,”
also includes herbaceous or nonwoody species, the largest group of flowering plants—which these plants were trees or shrubs,
both as free-standing plants and as climbers. have three-pored pollen grains. like many magnoliids are today.
Some herbaceous magnoliids have highly Most magnoliids have leaves with smooth However, a rival view—known as
specialized flowers. In birthworts and pipevines, margins, and a network of branching veins. the “paleoherb hypothesis”—
for example, the flower is shaped like a flaring Their fruits may be soft and fleshy, or hard and suggests that the first angiosperms
tube, lined with backward-pointing hairs. Flowers conelike, and can contain one or several seeds. were herbaceous, or nonwoody,
like these act as temporary traps for pollinating Species with fleshy fruits use animals to disperse plants with relatively fast life cycles.
flies, which are attracted by a powerful scent. But their seeds: many of them are swallowed whole This meant that they were good at
colonizing disturbed ground such as
these are an exception. Most magnoliid flowers and then scattered by birds. In prehistoric times,
riverbanks. Currently, neither view has
are structurally simple, with numerous spirally wild avocados may have been dispersed by giant proved decisive, although molecular
arranged parts, attached separately to a central ground sloths. Now that ground sloths are analysis does favor the idea that they
stem. Instead of sepals and petals (see p.122), extinct, avocados depend on human cultivation were woody plants.
they have a single layer of flaps that are similar to disperse their seeds and ensure survival.

CANELLALES PIPERALES M AGNOLI A L ES

There are two families in the order The order Piperales includes three families of Consisting almost exclusively of trees
Canellales: the Canellaceae and the herbaceous plants, trees, and shrubs, widely distributed and shrubs, the Magnoliales are a
Winteraceae. These are aromatic trees in tropical regions. The stems have scattered bundles primitive order, widely distributed
and shrubs with leathery entire leaves. of vascular tissue, a characteristic of monocots. in the fossil record. Although very
The flowers in most species have both Members of the family Piperaceae have tiny flowers variable, most have simple alternately
male and female parts and the fruit is which lack petals and are clustered in spikes. Many arranged leaves and flowers with both
a berry. The leaves and bark of some species are aromatic. male and female parts. There are
species can be used medicinally. The six families in the order, of
Winteraceae is a primitive family— which the Magnoliaceae are
its members have woody stems that the best known, being widely
contain no water-conducting vessels. 3¼ ft grown in gardens for their
1m spectactular flowers.
WINTER’S BARK 4 in
Drimys winteri 10 cm tepals
f: Winteraceae protect
Native to the coastal rainforest stamen
36 ft
11 m in Chile and Argentina, this tree
has aromatic bark and leaves BIRTHWORT
Aristolochia clematitis ASARABACCA
and fragrant flowers. Asarum europaeum 100 ft
f: Aristolochiaceae f: Aristolochiaceae 30 m
A foul-smelling and
poisonous perennial, This creeping species
birthwort was grown grows in European
for medicinal use. It is woodlands. Its glossy
native to Europe, evergreen leaves hide
growing in damp places. insignificant flowers.

BLACK PEPPER 100 ft CAMPBELL’S

Piper nigrum 30 m MAGNOLIA
13 ft
4m f: Piperaceae Magnolia campbellii
An evergreen TULIP TREE f: Magnoliaceae
climber from shady Liriodendron tulipifera Several of the 210 or so species
habitats in S. India f: Magnoliaceae of Magnolia are grown as
fruits, or and Sri Lanka, this A native of eastern N. America, garden ornamentals. This
peppercorns, species is widely the tulip tree grows in species is native to mountain
used dried cultivated for its woodland. The leaves turn yellow forests in China, India,
aromatic fruits. in the fall before dropping. and Nepal.
L AU R A L E S
The order Laurales consists of seven families
129
of trees, shrubs, and woody climbing plants.
A few genera grow in temperate parts

FLOWERING PLANTS • MAGNOLIIDS

of the world, but most are found in
tropical and subtropical regions. Their
classification is based on genetic analysis 8 ft/2.5 m
rather than morphological characteristics.
Many plants are aromatic and are used in
perfumery, cookery, and medicine. Others
provide lumber or are grown ornamentally. SASSAFRAS CAROLINA ALLSPICE
Sassafras albidum Calycanthus floridus
BAY LAUREL
f: Lauraceae f: Calycanthaceae leaf
This suckering deciduous Native to woods and cluster
Laurus nobilis
f: Lauraceae 80 ft
tree of woodland in eastern streamsides in the S.E. US,
Widely cultivated for its N. America has aromatic Carolina allspice has aromatic
25 m leaves which are brightly leaves and bark and large,
aromatic leaves used in
flavoring, this species colored in the fall. fragrant flowers.
grows in woods, scrub,
and rocky habitats around 60 ft
the Mediterranean. 18 m
edible
fruit

fruits 60 ft
18 m
60 ft
18 m leaves AVOCADO TREE
CINNAMON CALIFORNIA LAUREL Persea americana
unripe fruit Cinnamomum verum Umbellularia californica f: Lauraceae
f: Lauraceae f: Lauraceae Probably originating in S. Mexico,
The spice cinnamon comes Sometimes called “headache tree,” in well-drained parts of rainforests,
from the aromatic bark of this as scent from the crushed leaves the avocado tree is now widely
50 ft cinnamon species, found in lowland forests can cause headaches, this species cultivated for its edible,
15 m stick in Sri Lanka. from the W. US flowers in winter. pear-shaped fruits.

scented
flowers
white to dark pink
flowers open early
in spring before
the leaves
evergreen
leaves

26 ft
65 ft 8m
20 m
YLANG-YLANG
Cananga odorata SWEETSOP
f: Annonaceae Annona squamosa
Oil from the fragrant flowers of f: Annonaceae
ylang-ylang are used in perfumery. Believed to have Caribbean origins, sweetsop, or sugar
This evergreen tree comes from or custard apple, is widely cultivated. The fruits have
parts of Asia and Australia. edible flesh, which look and taste like custard.

glossy leaves NUTMEG

Myristica fragrans
f: Myristicaceae 26 ft
The spices nutmeg and 8m
mace are both
obtained from the
seeds of this evergreen
mace tree, a native of the
Molucca—or Spice—
Islands of Indonesia.
60 ft
seed, or nutmeg, 18 m
with mace covering

leaves

PAWPAW
Asimina triloba
edible
f: Annonaceae
fruits This deciduous tree is native
to damp woods of eastern
N. America. Its singly borne
flowers produce edible fruits.
130 MONOCOTS
Defined by their unique internal characteristic—revealed under a microscope— DIVISION ANGIOSPERMAE
anatomy, monocots include grasses involves the structure of their stems. Monocots
FLOWERING PLANTS • MONOCOTS

CLADE MONOCOTS

and palms, plus lilies, orchids, and have scattered bundles of vascular tissue—the ORDERS 11

many other ornamental plants. specialized cells that carry water and sap—unlike FAMILIES 77

eudicots, which have theirs in concentric rings. SPECIES 60,000

Early in the evolution of flowering plants, their This makes monocot stems more flexible than
family tree developed two major branches. A those of a typical eudicot, but it also makes it D E B AT E
smaller but still substantial branch became the harder for them to evolve into trees. Treelike DID MONOCOTS HAVE
monocots, while the larger branch became monocots, which principally include palms, A Q U AT I C O R I G I N S ?
the eudicots. Monocots get their name from have a very different way of growing to typical
their seeds, which have a single prepacked seed- broadleaved trees and conifers. Monocot trunks Living monocots include many
freshwater plants, and also some of
leaf, or cotyledon. Apart from this, there is no grow taller, but not thicker, and are usually topped the few flowering plants that live in
foolproof way of recognizing a monocot, although by a single rosette of leaves. the sea. According to a longstanding
there are strong clues. Most have long, narrow theory, monocots may have originally
leaves with parallel veins, and the flower parts survival strategies evolved in fresh water, before
(petals, stamens, and so on) are arranged in threes Smaller monocots vary greatly, from climbers diversifying and taking up life on land.
or multiples of three. The pollen grains have a to aquatic plants that survive harsh periods as This would account for the long, thin
single opening, unlike those of eudicots which underground storage organs, such as bulbs or tubers. leaves found in many species, and
have three. In many monocot flowers, such as Grasses thrive on the grazing effects of animals, also the internal structure of their
tulips, the sepals and petals are almost identical and are the only plant family that forms an entire stems (see left). Having spread to
land, some terrestrial monocots then
and are known as tepals, but many magnoliids habitat: grassland. In the tropics, many monocots came full circle and evolved into
share this feature. are epiphytic, growing high up in trees. This aquatic forms—duckweeds, for
Below ground, monocots usually have a cluster group includes bromeliads, and many orchids— example, originated in this way.
of fibrous roots, instead of a main taproot by far the largest monocot family, with more
with smaller roots leading off it. Another key than 25,000 species worldwide.

ACOR A L ES A L I SM ATA L E S
Only one genus and as few as This order includes many common aquatic plants as
two species make up the order well as the mainly land-based aroid family (Araceae).
Acorales. Called sweet flags, Aroids, which can be dramatic in appearance, have a male flower
these waterside and wetland distinctive reproductive anatomy consisting of a fleshy
plants have a fleshy flower spike of tiny flowers called a spadix and a leaflike
spike of small flowers and surround called a spathe. The other families in the
were once classified as order include many freshwater species, as well as
aroids (see right). several families of seagrasses.
Botanists now believe
they represent the
earliest branch of the WATER-PLANTAIN
Alisma plantago-aquatica
monocots family f: Alismataceae
tree, and may hold Found in waterside habitats, this
clues as to what the plant is common across Europe,
Asia, and N. Africa. It has white, 3¼ ft flower
first monocots pinkish, or pale purple flowers 1m stem
looked like. that last only a day.

oval arrow-
floating shaped leaf
leaf elliptical
3¼ ft
1m to oval
3¼ ft leaves
1m 3¼ ft
1m
branched whorl of three
flower cluster female flowers
SWEET FLAG ARROWHEAD
Acorus calamus Sagittaria sagittifolia
f: Acoraceae WATER HAWTHORN f: Alismataceae
Once cut to strew on Aponogeton distachyos This European wetland
floors because of its f: Aponogetonaceae plant grows arrowhead-
fresh citrus odor, Native to South Africa, this plant has shaped leaves above water,
this waterside plant become widely naturalized elsewhere. ribbonlike ones below
grows across the Its vanilla-scented flowers open just water, and sometimes
northern hemisphere. above the water surface. floating leaves too.
 20 in
50 cm
13 ft
4m
LORDS AND
LADIES
Arum maculatum
GIANT f: Araceae
TARO The spadix of this
Alocasia macrorrhizos spring-flowering
f: Araceae European native patterned leaves
Native from tropical Malaysia 6 in
heats up to attract 15 cm
to Queensland, Australia, this pollinating insects. glossy
huge-leaved aroid is cultivated leaf spathe
In the fall it
around the Pacific region produces poisonous
as an ornamental and minor red berries. FRIAR’S COWL
food crop. Arisarum vulgare
f: Araceae
The spathe of this widespread
23½ in 3¼ ft Mediterranean aroid curves over its spadix
60 cm 1m like a hood. Its leaves are arrow-shaped.
10 ft
3m
SWISS CHEESE DUMBCANE
PLANT Arum seguine
Monstera deliciosa f: Araceae
f: Araceae This aroid from tropical
The mature leaves of American forests causes
this tree-climbing severe swelling and pain if
aroid from C. America chewed. Houseplant varieties
become perforated and have showy, attractive foliage.
feathery divided. Swiss cheese
roots plant is widely grown
WATER as a houseplant. 20 ft/6 m
GOLDEN CALLA LILY LETTUCE 65 ft/20 m
Zantedeschia elliottiana Pistia stratiotes SWEETHEART PLANT
f: Araceae f: Araceae Philodendron scandens
This showy aroid with Of uncertain origin, this TARO f: Araceae
yellow spathes is found only lettuce-sized member of the Colocasia esculenta Very common in its
in cultivation. Its wild aroid family floats in warm
f: Araceae native C. America, this
relatives live in South Africa, fresh waters worldwide, This large-leaved fast-growing climbing
mainly in wetland habitats. often clogging waterways. aroid, native to aroid is also a popular
tropical Asia and the ornamental houseplant.
Pacific, has been
3⁄16 in cultivated extensively
5 mm since ancient times
giant spadix for its edible tubers.
with tiny flowers
at base
61⁄2 ft
giant spathe 2m

3¼ ft 3¼ ft
1m 1m
FAT DUCKWEED
Lemna gibba
f: Araceae DRAGON ARUM COBRA LILY
Despite appearances, Dracunculus vulgaris Arisaema consanguineum
duckweed belongs to the f: Araceae f: Araceae
aroid family. This widespread With its dark red spathe Native from the E. Himalayas to
floating plant grows in tiny and odor of rotting meat, China, this summer-flowering
ovals up to 5 mm across. this E. Mediterranean aroid aroid has a distinctive striped
is well adapted to attract spathe. The entire plant is
pollinating flies. very poisonous.
5ft
1.5 m
12 in
30 cm

1/32 in 20 ft
1 mm 6m
5 ft
1.5 m
TITAN ARUM
Amorphophallus
SPOTLESS WATERMEAL titanum WESTERN SKUNK BOG ARUM FLOWERING RUSH
Wolffia arhiza f: Araceae CABBAGE Calla palustris Butomus umbellatus
f: Araceae After its 10 ft (3 m) Lysichiton americanus f: Araceae f: Butomaceae
Found worldwide, this flower stalk has died f: Araceae This wetland and shallow- A showy waterside plant,
rootless duckweed is the down, this giant aroid Native to wet habitats in water aroid is a popular flowering rush is the only
world’s smallest flowering from Sumatra grows western N. America, this ornamental. It is native to member of its family native
plant, forming tiny green a single, long-lasting, aroid gives off a strong odor cool regions of the to Eurasia. It is invasive
ovals only 1 mm wide. treelike leaf. to attract insects. northern hemisphere. in N. America.
 A L I SM ATA L E S NEPTUNE GRASS
Posidonia oceanica
green
132 flowers in f: Posidoniaceae
short spikes This seagrass sometimes
forms dense patches
on sandy seabeds, at
FLOWERING PLANTS • MONOCOTS

61⁄2 ft depths of up to 115 ft

2m 3¼ ft (35 m) beneath the
1m Mediterranean Sea.
BROAD-LEAVED
SOUTH PONDWEED
AMERICAN Potamogeton natans
WATERWEED f: Potamogetonaceae tufts of
Egeria densa This freshwater plant has broad surface
ribbonlike
f: Hydrocharitaceae leaves and grasslike underwater ones. It is leaves brown
leaves
Native from Brazil common across the northern hemisphere. with age
to Uruguay, this
underwater plant, grown 3¼ ft
by aquarists, has escaped 1m
to clog lakes and
waterways across the
temperate world.

horizontal stem
buried in sand

COMMON EELGRASS
floating three- Zostera marina
leaf petaled f: Zosteraceae
TAPE GRASS flower Growing on sandy sea
Vallisneria americana 3¼ ft 8 in floors across the northern
f: Hydrocharitaceae 1m 20 cm hemisphere, this grasslike
This submerged, plant forms underwater
freshwater plant is found FROGBIT “meadows” that are an
from southern N. America Hydrocharis morsus-ranae important marine habitat.
to Colombia. Its male f: Hydrocharitaceae
flowers break loose and This free-floating freshwater plant from Eurasia
float, perhaps encountering spreads via horizontal underwater stems. Male 3¼ ft
1m
female ones to pollinate. and female flowers grow on separate plants.

A S PA R AG A L E S
funnel-shaped,
This diverse order of 14 families includes familiar light to deep
garden flowers such as daffodils and irises, blue flowers
desert-adapted plants such as agaves, and a few 3¼ ft/1 m
trees. Some of its members were previously
classified in the order Liliales, but modern genetic
techniques have changed our understanding of
these interrelationships. The order also includes
the highly specialized orchid family.
dense flowerhead
of small, bell-
shaped flowers

23½ in 8 in
ONION 60 cm 20 cm
Allium cepa
f: Amaryllidaceae
Evidence from
ancient Egypt shows
that the onion has
been cultivated for
at least 5,000 years.

edible bulb
32 in 20 in
80 cm 45 cm AFRICAN LILY
Agapanthus africanus SNOWDROP
f: Amaryllidaceae Galanthus nivalis
ROUND-HEADED LEEK RAMSONS Native to South Africa, f: Amaryllidaceae
Allium sphaerocephalon Allium ursinum this plant has a tall, The early spring
f: Amaryllidaceae f: Amaryllidaceae flowering stem and flowers of this
This European relative of the Related to and smelling of narrow, arching leaves. shade-tolerant
onion favors limestone soils. garlic, this plant often carpets It can survive fires in European native have
Gardeners grow it for its the floors of European its native habitat, three white sepals that
colorful, tightly packed woodlands in spring with its regrowing from a fleshy are much longer than
flowerheads. broad, green leaves. underground stem. the three inner petals.
 18 in 5 ft
45 cm 1.5 m 133
SWAMP LILY
Crinum x powellii
f: Amaryllidaceae
This bulb-forming plant is a
hybrid of two South African
species of Crinum. It is hardy
in cooler climates.

NATAL LILY
Clivia miniata
f: Amaryllidaceae
Horticulturalists have bred many varieties
of this South African woodland plant for its
showy flowers and straplike evergreen leaves. bract protects long-stalked,
developing trumpet-shaped
flowerhead flower

3¼ ft
1m

23½ in
60 cm

30 in large, showy
75 cm flower with
six tepals

AMARYLLIS FIRECRACKER FLOWER spreading rosette CENTURY

Hippeastrum sp. Dichelostemma ida-maia 26 ft PLANT
of gray-green, 8m
f:Amaryllidaceae f: Asparagaceae fleshy leaves Agave americana
This genus of showy, Native to Oregon and f: Asparagaceae
Native to Mexico and SPIDER PLANT
bulb-forming plants is native northern California, this mainly Chlorophytum comosum
to warmer parts of the woodland species has attractive southwestern US, this f: Asparagaceae
Americas. Many cultivated tubular flowers that make it succulent, or water-
varieties and hybrids exist. popular as a garden plant. retaining plant, flowers Forming new plantlets from its leaves
once after 10–30 years and arching stems, the variegated
and then dies. forms of this African species are
well known as houseplants.

straight, stiff
28 in evergreen
70 cm mature, leaves
feathery leaves

6½ ft
2m

COMMON
ASPARAGUS
Asparagus officinalis
f: Asparagaceae
Cultivated for its tender
young shoots (spears),
this European species
becomes a tall, feathery
plant if left to grow.

20 in 16 ft pendulous,
50 cm 5m 50 ft
bell-shaped 15 m
flowers

WILD DAFFODIL JOSHUA TREE

Narcissus pseudonarcissus ST. BERNARD’S LILY SPANISH DAGGER Yucca brevifolia
f: Amaryllidaceae Anthericum liliago Yucca gloriosa f: Asparagaceae
The ancestor of many f: Asparagaceae f: Asparagaceae Native to the Mojave
cultivated varieties of This European native with Like all yuccas, this coastal Desert, this woody,
daffodil, this native European grasslike leaves likes sunny species from S.E. US relies many-branched yucca is
species of mountains and hillsides and other open places. on a specialized yucca moth thought to live for
woods is becoming rarer. It prefers rich, moist soils. for pollination. hundreds of years.
 A SPA R AG A L E S
134
flower stem holds
up to 20 flowers 28 in 3¼ ft
FLOWERING PLANTS • MONOCOTS

70 cm 1m

10 in SOLOMON’S-SEAL
25 cm
Polygonatum multiflorum
f: Asparagaceae
A mainly woodland plant, this
Eurasian species has leafy, arched
stems and tube-shaped,
scentless flowers.

5 ft
BUTCHER’S BROOM
1.5 m Ruscus aculeatus
f: Asparagaceae
LILY-OF-THE- SEA SQUILL Flowers and fruits grow
VALLEY Drimia maritima straight out of the “leaves”
Convallaria majalis f: Asparagaceae (actually flattened stems)
f: Asparagaceae Native to Mediterranean of this shrubby
A sweetly scented plant coasts, this large- European plant.
from temperate Eurasia, bulbed plant grows
lily-of-the-valley has 23½ in a tall spike of white flowers
poisonous red berries. 60 cm in late summer after its
leaves have withered.
CAST-IRON PLANT
Aspidistra elatior
f: Asparagaceae
Native to Japan, this woodland plant is now
a popular houseplant. Its small, purple bell-shaped flower
flowers bloom close to the soil.

12 in 35 in
30 cm 90 cm

MOTHER-IN- 20 in
LAW’S TONGUE 50 cm
Dracaena trifasciata
STAR-OF-BETHLEHEM f: Asparagaceae PORTUGUESE SQUILL
Ornithogalum angustifolium This tropical Scilla peruviana
f: Asparagaceae W. African species f: Asparagaceae
This widespread European plant with stiff patterned From S.W. Europe, this showy,
closes its flowers in dull weather. leaves is a popular bulb-forming perennial has long,
Its narrow, grooved leaves have houseplant. It is also broad leaves growing from
a central white stripe. grown for its fibers. the plant’s base.

12 in APHYLLANTHES
30 cm Aphyllanthes monspeliensis
f: Asparagaceae
When not flowering, this
Mediterranean species looks
like a clump of rushes, with
many slender stems that are
almost leafless. 18 in
45 cm

spike of up to 50
fragrant flowers

12 in
30 cm

HYACINTH
Hyacinthus orientalis CAPE COWSLIP
“Blue jacket” Lachenalia aloides
f: Asparagaceae f: Asparagaceae
This is one of the many This bulb-forming species,
fragrant, colorful varieties native to the southwestern
derived from a S.W. Asian tip of Africa, has only one
species over many centuries. or two strap-shaped leaves.
 BLUE HOSTA
Hosta “Halycon”
f: Asparagaceae 135
This hybrid is one of many
garden varieties of Hosta, a genus 18 in
of shade-tolerant plants native 45 cm

FLOWERING PLANTS • MONOCOTS

to N.E. Asia.

3¼ ft 50 ft 23½ in
1m 15 m 60 cm

ENGLISH BLUEBELL GREAT CAMAS DRAGON TREE

18 in Hyacinthoides Camassia leichtlinii Dracaena draco
45 cm non-scripta f: Asparagaceae f: Asparagaceae
f:Asparagaceae The edible bulbs of several Rare in its native Canary
Native to W. Europe, this Camassia species were Islands and Madeira, this
bulb-forming plant sometimes cooked like potatoes many-branched tree survives
carpets entire woodland floors by Native American as a popular ornamental.
with blue flowers peoples as a staple food Its red sap was prized
in spring. called Quamash. as “dragon’s blood.”
TASSEL HYACINTH
32 in Muscari comosum
80 cm f: Asparagaceae
The flower heads of this
Mediterranean species
have fertile and sterile
flowers, the latter forming
the purplish “tassel”
at the top.

COMMON FRINGE LILY

Thysanotus tuberosus
f: Asparagaceae
Native to S.E. Australia, this
narrow-leaved plant has
fringe-petaled flowers that
each open for only one day.
65 ft 3¼ ft
20 m 1m
CABBAGE TREE FIELD
Cordyline australis GLADIOLUS
f: Asparagaceae Gladiolus italicus 3¼ ft 23½ in
One of New Zealand’s most common trees, f: Iridaceae 1m 60 cm
shown here as a young plant, the cabbage A Mediterranean
tree develops a sturdy trunk topped member of the iris
by dense tufts of narrow leaves and showy family, the field
spikes of white flowers. gladiolus flowers BEARDED IRIS MONTBRETIA
between March Iris germanica Crocosmia x
and June. f: Iridaceae crocosmiiflora
3¼ ft Botanists are unsure f: Iridaceae
1m whether this fragrant- A 19th-century hybrid
flowered garden iris was once created from two
PALE YELLOW- native to the E. Mediterranean South African species, this
EYED GRASS or originated as a garden popular garden plant is
Sisyrinchium striatum hybrid. invasive in some areas.
f: Iridaceae
Native to Chile KEW’S FREESIA
and Argentina, this Freesia x kewensis
representative of a large 12 in
30 cm
f: Iridaceae
New World genus has Derived from native
pale flowers marked South African
with fine purple stripes. ancestors, this freesia
is a sweet-smelling 23½ in
garden hybrid in the 60 cm
northern hemisphere.

18 in
45 cm
23½ in
60 cm
SAFFRON CROCUS
GRAND CHRISTMAS Crocus sativus
BELLS f: Iridaceae BARBARY NUT
Blandfordia grandiflora The dried stigmas of this Gynandriris sisyrinchium
f: Blandfordiaceae cultivated Mediterranean f: Iridaceae
Nectar-sipping birds seek out crocus have long been This wild Mediterranean iris
this large-flowered species of sold as saffron, used grows from a corm (a bulblike
coastal E. Australia. Its leaves for coloring and underground stem), which helps it
are narrow and grasslike. flavoring food. survive unfavorable conditions.
 A SPA R AG A L E S ONCIDIUM SP.
f: Orchidaceae
TRACY’S
CYMBIDIUM
136 Native to the tropical Cymbidium tracyanum
Americas, this genus of f: Orchidaceae
broad-lipped orchids This epiphytic orchid
includes about 330 from Burma, Thailand,
FLOWERING PLANTS • MONOCOTS

tree-living species of and S.W. China

8 in 3¼ ft varying sizes. 3¼ ft produces its strongly
20 cm 1m 1m fragrant flowers
in the fall.
FAIRY
SLIPPER
ORCHID 23½ in
Calypso bulbosa 60 cm
f: Orchidaceae PHRAGMIPEDIUM 32 in
X SEDENII 80 cm
Widespread across f: Orchidaceae
cooler parts of the northern This fragrant,
hemisphere, this fragrant, ground-living orchid
single-flowered orchid is a hybrid created HYACINTH ORCHID
prefers damp woodland from species of the Dipodium squamatum
and marshland. genus Phragmipedium, f: Orchidaceae
slipper orchids native A leafless, ground-living Australian orchid, this
to tropical regions of species depends on associating with underground
the Americas. fungi to survive. It lives in woodland habitats.
12 in
30 cm
23½ in 14 in 12 in
60 cm 35 cm 30 cm

LESSER BUTTERFLY
ORCHID
Platanthera bifolia
f: Orchidaceae
Found in various habitats
across temperate Eurasia,
this sweetly scented,
pale-flowered orchid is
visited and pollinated by
night-flying moths.
PYRAMIDAL ORCHID PINK LADY’S HAIRY SLIPPER
Anacamptis pyramidalis SLIPPER ORCHID
f: Orchidaceae Cypripedium acaule Paphiopedilum villosum
This chalk-loving orchid from f: Orchidaceae f: Orchidaceae
temperate Eurasia sticks its Widespread in eastern Native to S. China and
pollen sacs onto visiting N. America, this parts of S.E. Asia, this
butterflies and moths, as first two-leaved slipper slipper orchid has been
described by Charles Darwin. orchid favors acid soil used to create many
as in pine forests. ornamental hybrids.

¾ in
2 cm

WAGENER’S
MASDEVALLIA
Masdevallia wageneriana
f: Orchidaceae
The sepals of this small
epiphytic orchid from the
mountains of N. Venezuela
have narrow “tails” (spurs),
a feature typical
flowers have three of the genus.
outer sepals and
three inner petals
12 in 6 in 23½ in
30 cm 15 cm 60 cm

ORANGE GUARIANTHE central section RED HELLEBORINE

Guarianthe aurantiaca WINDOWSILL ORCHID has distinctive Cephalanthera rubra
f: Orchidaceae Pleione formosana frilly edge f: Orchidaceae
This tree-living species is native f: Orchidaceae This ground-living species with
to tropical C. America. Many Native to parts of China, this rose-pink or purplish flowers
ornamental varieties and hybrids small ground-living orchid dies grows in open woodlands from
have been bred by horticulturalists. back during the winter months. Europe eastward to W. Asia.
 VIOLET BIRD’S-NEST DENDROBIUM SP.
ORCHID f: Orchidaceae
Limodorum abortivum Including more than 137
f: Orchidaceae 1,000 species of diverse
This S. European orchid shapes, colors, and sizes,
lacks green leaves. For food, this genus of epiphytic

FLOWERING PLANTS • MONOCOTS

it depends entirely on Russula orchids is found from S.E.
fungi living in its roots. Asia to New Zealand. 3¼ ft
1m
32 in 6½ ft
80 cm 2m
LIPPEROSE MOTH ORCHID
Phalaenopsis ‘Lipperose’
35 in
f: Orchidaceae
90 cm This cultivated hybrid is one of
12 in many developed using the genus
30 cm Phalaenopsis from S.E. Asia with
TONGUE LIZARD its broad, flattened flowers.
ORCHID ORCHID
Serapias lingua Himantoglossum hircinum
f: Orchidaceae f: Orchidaceae
The lip of this unusual The long lips of this orchid
Mediterranean orchid’s from southern Europe have
flower hangs down like a fanciful resemblance to
a tongue, acting as a tiny lizards, hence its
landing platform for common name.
visiting insects.

three sepals
and two petals 23½ in 50 ft
on each flower 60 cm 15 m
23½ in
drooping 12 in
60 cm sepals 30 cm

MILITARY ORCHID COMMON DONKEY

Orchis militaris ORCHID
f: Orchidaceae Diuris corymbosa
This chalk-loving Eurasian f: Orchidaceae
orchid may have acquired Native to S.E. and S.W.
its name because the Australia, this orchid gets its
seed pod individual flowers resemble common name from the
helmeted human figures. shape of its side petals, which
leathery resemble donkeys’ ears.
evergreen leaves
ROTHCHILD’S VANDA FLAT-LEAVED VANILLA
Vanda ‘Rothschildiana’ ORCHID
f: Orchidaceae Vanilla planifolia
This cultivated hybrid was created f: Orchidaceae
Native to Mexico and BEARDED
by crossing species of the genus GREENHOOD
Vanda, which are tree-living orchids C. America, this climbing ORCHID
native to tropical Asia. orchid is the source of the Pterostylis sp.
flavoring vanilla. f: Orchidaceae
Scent from these
AUTUMN ground-living
LADY’S-TRESSES Australian orchids
lanceolate Spiranthes spiralis attracts small male
leaves on f: Orchidaceae flies. A feathery trap
each stalk This small, mainly grassland then spurs the fly
orchid from temperate to collect the
Eurasia is notable for its pollen sacs.
flowers being arranged
spirally up the flower stalk. flower resembles
a fat bumblebee
8 in
flower attacts 20 cm
pollinating sepal and petals form
insects protective hood over
inner flower

3¼ ft
1m flower trigger
snaps shut to
trap insect
16 in 20 in
40 cm 50 cm
DARK-RED lower sepals
HELLEBORINE NUN’S provide landing BEE ORCHID
23½ in Epipactis atrorubens ORCHID stage for insects Ophrys apifera
60 cm f: Orchidaceae Phaius tankervilleae f: Orchidaceae
The long roots of this f: Orchidaceae Although in a genus whose
fragrant Eurasian This fragrant ground-living flowers attract pollinating
orchid help it grow orchid is widely cultivated. male insects by mimicking
successfully even Its native range is tropical females, this Eurasian
within the cracks of and subtropical S.E. Asia orchid in practice often
limestone cliffs. and the South Pacific. pollinates itself.
 RAMBLING STEMS
The many-bulbed orchid grows all over its
138 chosen support structure, whether it be a
rock or another plant, and forms a dense
mat of stolons—the horizontal stems that
reach out from the base of the plant. broad, whitish petal
forms a lip, which acts
as a flag to attract
pollinating insects
outer three sepals
enclose the flower bud
M A N Y- B U L B E D O RC H I D 139
Dinema polybu lbon
This diminutive orchid is the only species in the genus Dinema. Rambling and
prolific, it is an epiphytic or lithophytic plant, meaning that it grows on trees or
rocks, using them for support. It gains its nutrients from the air, animal detritus,
or other plant remains, and absorbs water from rain or mist. Moisture collects on
the orchid’s leaves, which are thick and waxy, and have reduced stomata—all features
which help to reduce water loss. The leaves funnel water into upright, swollen
outgrowths from the stems, called pseudobulbs. These act as water-storage organs
to keep the orchid hydrated during tropical dry seasons. The creeping stems,
with the pseudobulbs, in turn connect to aerial and (sometimes) underground
roots. These absorb dissolved nutrients through a spongy,
outer layer of cells called velamen. In winter, each size Height 3 in (7.5 cm)
pseudobulb produces a single flower. Plants can survive habitat Humid, mixed forests
temperatures down to 45°F (7°C), and are a favorite distribution Mexico, C. America,
of orchid collectors. Jamaica, Cuba
leaf type Parallel veins

flattened leaf tip

has a notch

PSEUDOBULB
These small, oval,
bulblike outgrowths
from the creeping
stems store water for
dry periods. Each
produces one to
three glossy leaves.

FLOWER
The pseudobulb produces a small,
sweet-smelling flower with narrow,
yellow-brown sepals, purple petals,
and a spreading, white lip.
POLLEN SACS (POLLINIA)
Orchid pollen grains are contained
in two sacs. A sticky mass glues the
sacs onto pollinating insects, which
carry the pollen to another plant.
AERIAL ROOTS
The orchid’s aerial roots do not have
to connect to the soil. The older parts
of the root have a protective layer of
dead cells that act like blotting paper
to absorb and retain water.
 A SPA R AG A L E S 5 ft
1.5 m
LILIALES
140 Many plants once considered part
16 ft
tall spike of
16 ft of the lily family, including onions
5m 5m and hyacinths, have been moved into
tiny, white or
FLOWERING PLANTS • MONOCOTS

cream flowers the order Asparagales (see pp.132–39).

ORANGE DAY LILY
Hemerocallis fulva The ten families in the Liliales include
f: Asphodelaceae the true lilies and tulips, the
Each flower of this ornamental species sarsaparilla and colchicum families,
lasts just one day. Native to Asia, it can and the colorful alstroemeria
be an invasive weed in N. America.
family from the New
World. Beware that
several lily species
are highly toxic
to cats.

orange-red
flower spike
GRASS TREE NEW ZEALAND FLAX fades to yellow
Phormium tenax near base 18 in
Xanthorrhoea australis 45 cm
f: Asphodelaceae f: Asphodelaceae
Perhaps living for up to 450 years, This common New Zealand species
this bushfire-resistant Australian grows large leaves at ground level and WILD TULIP
plant develops a sturdy trunk and a a tall flower stalk bearing tubular Tulipa sylvestris
spherical crown of narrow leaves. reddish flowers. f: Liliaceae
Still widespread as a native
plant in southern Europe,
sap from leaves has this yellow-flowered relative
healing properties of the cultivated garden
tulip grows wild in
meadows, open
woodland, and
rocky places.

16 in 20 in
40 cm 50 cm
5 ft
1.5 m

COMMON ASPHODEL
Asphodelus aestivus
f: Asphodelaceae
This common narrow-leaved
Mediterranean plant produces many 3¼ ft 5 ft
pinkish white flowers on a tall stem. 1m 1.5 m
HERB PARIS SNAKESHEAD
MEDICINAL ALOE TORCH LILY Paris quadrifolia FRITILLARY
Aloe vera Kniphofia uvaria f: Liliaceae Fritillaria meleagris
f: Asphodelaceae f: Asphodelaceae Found in ancient f: Liliaceae
This spiny succulent Native to the S.W. woodland in temperate This bulb-forming
buds at top of adapted to dry areas has tip of Africa, this Eurasia, this European species has
spike open last been cultivated since multiflowered lily four-leaved plant has a distinctive checkered
ancient times for its with long, narrow single green flower that flowers. It grows wild in
reputed medicinal leaves is now a popular develops into a black damp meadows but is more
properties. ornamental specimen. inedible fruit. common in cultivation.

DIOSCOREALES
BLACK BRYONY
18 in This order contains just three families. It is dominated Tamus communis
45 cm
by the yams, a family of mainly tropical climbing f: Dioscoreaceae
plants. Several species of yam have been cultivated 13 ft/4 m This poisonous
European climbing
since ancient times for their edible tubers. plant of the yam
The small bog asphodel 16 ft
family is named for
family is mainly its black underground unripe
5m fruits are
north temperate. tuber. Its globular
fruits turn bright red green
when ripe.
edible
BOG ASPHODEL tuber
Narthecium ossifragum YAM
f: Nartheciaceae Dioscorea sp
This European plant f: Dioscoreaceae
lives in nutrient-poor Widely cultivated in the
upland habitats. After tropics, yams grow large,
fertilization, its seed starch-rich tubers. Plants heart-shaped
capsules develop a have stems that trail or climb leaf
fiery orange color. and heart-shaped leaves.
 MADONNA LILY GAGEA RETICULATA
Lilium candidum f: Liliaceae
f: Liliaceae 6 in This small, bulb-forming plant 141
Often encountered in Christian art as a symbol 15 cm grows in open habitats across
of purity, this widely cultivated lily is native much of temperate Asia, as well as
to the E. Mediterranean. S.E. Europe and N. Africa.

FLOWERING PLANTS • MONOCOTS

61/2 ft
2m

female stigma
receives the
pollen
61/2 ft
2m 61/2 ft/2 m

TURK’ S CAP LILY FLAME LILY

Lilium martagon Gloriosa superba
f: Liliaceae f: Colchicaceae
male stamen This widespread Eurasian This striking plant, found
releases the pollen lily has the reflexed from S. Africa to S.E. Asia,
(backward-bending) tepals is a climber, clambering
that form the flower shape upward with the aid
characteristic of the species. of tendrils.

flowerheads

6 in
15 cm
MEADOW
SAFFRON
Colchicum autumnale
f: Colchicaceae
The crocuslike flowers
10 ft of this European native
3m appear in the fall,
months before its
leaves. Despite being
GIANT HIMALAYAN LILY poisonous, it is
Cardiocrinum giganteum cultivated widely.
f: Liliaceae typical
leaf
Found from the Himalayas to
China, this giant lily grows for
several seasons before flowering,
after which the main plant dies.
flower with
six tepals
(lobes)

leaf twists from

stem so underside
faces upwards

up to 40
leaves arranged tubular flowers
spirally up stem
PERUVIAN LILY
TRAILING LILY Alstroemeria sp.
Bomarea multiflora f: Alstroemeriaceae
f: Alstroemeriaceae This lilylike S. American genus
This many-stemmed includes many popular
climbing plant, with clusters ornamentals. The leaves twist
of orange-red flowers, is 13 ft during growth to become 4 ft
native to S. America. 4m anatomically upside down. 1.2 m
 LILIALES 5 ft PA N DA N A L E S
142 1.5 m
Found mainly in the tropics, the five families in this order include over
1,300 species of trees, shrubs, climbers, and smaller plants. Many look
CHILEAN superficially like palms, except that they have simpler, strap-shaped leaves.
BELLFLOWER
FLOWERING PLANTS • MONOCOTS

Lapageria rosea About half of them belong to the single genus Pandanus (screw pines).
f: Philesiaceae THATCH SCREW PINE
Cultivated for its Pandanus tectorius
spectacular bell-shaped multi-seeded f: Pandanaceae
flowers, this twining
fruit
No relation of real pines, this
climber is native to tropical coastal tree has long
humid woodlands been a vital source of materials
of Chile. for Pacific Island cultures.
30 ft
10 m

50 ft
15 m

FALSE HELLEBORINE
berries SMILAX Veratrum sp.
Smilax aspera f: Melanthiaceae
f: Smilacaceae Native to the northern
flower Native to the Mediterranean hemisphere, this genus of 60 ft
cluster and S.W. Asia, this vigorous poisonous plants produces mature tree 18 m
climber bears male and female branched flower heads of
flowers on separate plants. greenish white flowers.

ARECALES
100 ft
Since 2016, a family of 16 endemic 30 m
Australian trees has been added
to the order Arecales, which is
dominated by more than 2,000
species in the palm family.
Typically growing from one 65 ft
20 m
central bud, palms vary from DATE PALM
towering trees to slender climbing SUGAR PALM Phoenix dactylifera
rattans. Their huge leaves are Arenga pinnata f: Arecaceae
f: Arecaceae Seen here as a young 100 ft
either feather-shaped or fanlike. This native of India and S.E. Asia 30 m
tree, this cultivated
Often associated with desert grows showy yellow flower heads. palm from the
islands, most species actually It yields many products, including Middle East grows
live in tropical rainforests. sugar from its sap and fibers. as separate male and
female trees.
80 ft
pinnate
25 m leaf

COCONUT PALM
Cocos nucifera
f: Arecaceae
Now widely
cultivated, this
species probably
originated in the
W. Pacific, colonizing
new islands via its
floating fruits.

mature single-seeded
tree fruit
60 ft
18 m
mature ripe DESERT FAN
tree fruit PALM
Washingtonia filifera PETTICOAT PALM
f: Arecaceae Copernicia macroglossa
BETEL NUT PALM Dead leaves hanging f: Arecaceae
Areca catechu below this palm’s Native to Cuba, this
f: Arecaceae crown provide shelter relatively small palm is
Cultivated for its seeds, which release a for birds and insects in named for the skirt of
psychoactive substance when chewed, this its native deserts of dead leaves that it retains 23 ft
palm originates from S.E. Asia. S.W. US. below its crown. 7m
 65 ft
20 m
pinnate leaf
143

FLOWERING PLANTS • MONOCOTS

20 ft
ripe fruit 6m 50 ft
15 m
AFRICAN OIL PALM BOTTLE PALM
Elaeis guineensis Hyophorbe lagenicaulis SAGO PALM
f: Arecaceae f: Arecaceae Metroxylon sagu
A plant of humid tropical lowlands, this palm This swollen-based palm comes from f: Arecaceae
is extensively cultivated beyond its native tiny Round Island near Mauritius. It is Shown here as a young plant,
Africa for the oils contained in its fruits. widely grown as an ornamental. this swamp-living palm
probably originated in New
Guinea but is now grown
throughout S.E. Asia.

100 ft
30 m
80 ft 30 ft
seed 25 m 10 m

palmate RAFFIA PALM

leaf Raphia farinifera
65 ft
f: Arecaceae
20 m At 65 ft (20 m) long,
the leaves of this
CHUSAN PALM African palm are
Trachycarpus fortunei the largest of
f: Arecaceae COCO DE MER ROYAL PALM any tree.
From C. China, this cold-tolerant species has Lodoicea maldivica Roystonea regia
separate male and female plants. The females f: Arecaceae f: Arecaceae
produce round, blue-black fruits. The seeds of this palm Often planted in
from the Seychelles are stately avenues
the biggest in the plant in the tropics, 65 ft
kingdom. They take six this handsome, 20 m
years to mature. smooth-trunked
species is native from
C. America to
Florida and Cuba.

fan-shaped
leaf up to 3¼ ft
(1 m) across

PALMYRA PALM
Borassus flabellifer
f: Arecaceae
This tall-trunked species from S. Asia favors
drier habitats. It is cultivated for its fruits
and sugar-yielding sap.

10 ft 80 ft
3m 25 m
BRAZILIAN EUROPEAN FAN PALM CHILEAN WINE PALM
WAX PALM Chamaerops humilis Jubaea chilensis
Copernicia prunifera f: Arecaceae f: Arecaceae
f: Arecaceae Often trunkless in the wild, Also known as the coquito
A waxy coating on the leaves of cultivated specimens grow palm, this massive-trunked,
this palm from N.E. Brazil helps it several short trunks from a cold-tolerant palm is native
50 ft resist drought. The harvested wax single base. It originates from only to C. Chile, where it is
15 m is used in polish and soap. Mediterranean countries. now a protected species.
 COMMELINALES 23½ in
60 cm
144 This order contains five families, two of which are
QUEEN'S SPIDERWORT
minor groups of about five species each, plus BLUE SPIDERWORT Dichorisandra reginae
the three larger families illustrated here. Most are Commelina coelestis f: Commelinaceae
f: Commelinaceae
FLOWERING PLANTS • MONOCOTS

low-growing plants, typically from warmer regions. Sometimes planted as

A popular garden flower
Many have attractive, three-petaled blue flowers grown in warmer regions, this
ground cover, this sprawling tropical woodland species is
(reduced to two petals in some species), making plant is native to Mexico from Peru. It has blue flowers
them popular as ornamentals. and C. America. with white centers.

TAHITIAN BRIDAL VEIL

Callisia procumbens 6 in 12 in
15cm 30 cm
f: Commelinaceae
Native to C. and S. America,
this species has weak
stems and three-petaled
white flowers.

4 in 2¼ ft
10 cm 70 cm
TURTLE VINE SILVER INCH PLANT
Callisia repens Tradescantia zebrina
f: Commelinaceae f: Commelinaceae
This succulent plant from With striped, succulent foliage,
forest edges in the American this ground-living species from
tropics spreads by rooting trailing the tropical Americas is also a
from creeping stems. stem popular houseplant.

POA LES SLENDER CLUB-RUSH

Isolepis cernua
The 14 families in the Poales include wind-pollinated 18 in f: Cyperaceae
45 cm With narrow, green stems and
families that dominate certain ecosystems: grasses in silvery flower heads, this
prairies and savannas, sedges and rushes in bogs and widespread temperate sedge is
heathlands, especially in the northern hemisphere, also dubbed “fiber-optics grass.”
and restios in heathlands and swamps in the southern
hemisphere. Bromeliads mainly live as epiphytes 12 in
30 cm
growing on tropical trees. SCARLET STAR
Guzmania lingulata
f: Bromeliaceae
PINK QUILL 20 in This tree-living bromeliad
Wallisia cyanea 50 cm has a wide native range
f: Bromeliaceae from C. America to
5 ft
Native to Ecuador and 1.5 m
Brazil. It is a popular
Peru, pink quill grows ornamental species.
on rainforest trees at up
to 2,800 ft (850 m)
above sea level. long, narrow
leaf

arching stalk of
flower cluster
12 in
30 cm
PINEAPPLE
Ananas comosus 12 in
f: Bromeliaceae 30 cm
red bracts Christopher Columbus introduced
surround Europeans to this cultivated bromeliad
flowers with its large, seedless fruits. Its wild
BIRD’S NEST origins are uncertain.
BROMELIAD
orange-red bracts protect Nidularium innocentii
tiny, white flowers f: Bromeliaceae spiny-leaved rosette
The small, white flowers
of this Brazilian bromeliad
nestle within the
chamber formed
14 in by the surrounding
vaselike rosette of leaves 35 cm
collects rainwater colored bracts.

LORENTZ’S
DYER'S TILLANDSIA BROMELIAD BLUSHING BROMELIAD
Racinaea dyeriana Deuterocohnia Neoregelia carolinae
f: Bromeliaceae lorentziana 10 in
f: Bromeliaceae
This epiphytic bromeliad is f: Bromeliaceae 25 cm At flowering time, the central
endangered in the wild due Native to the Andes of leaves of this Brazilian
to destruction of its native Argentina and Bolivia, this is bromeliad turn crimson, and it
mangrove forests in Ecuador. a ground-dwelling species. produces blue or violet flowers.
 yellow blotch on 3¼ ft/1 m
uppermost petal 145
of each flower
10 ft
3m

FLOWERING PLANTS • MONOCOTS

18 in
45 cm

TALL KANGAROO PAW

Anigozanthos flavidus
f: Haemodoraceae
From sandy areas of S.W. Australia,
this species gets its common name
from the appearance of its swollen leaf stalk
woolly-haired flower buds.

COMMON WATER PICKEREL WEED

HYACINTH Pontederia cordata
flowers in compact Pontederia crassipes f: Pontederiaceae
conical cluster f: Pontederiaceae Fast-growing and with
Native to Amazonia, this striking blue flower heads,
spike of male flowers floating plant is a major this water-edge species is
tropical pest, but can be common from eastern N.
used to absorb pollution. America south to Argentina.

PENDULOUS SEDGE 16 ft/5 m

Carex pendula
f: Cyperaceae 4½ ft 5 ft
Like other true sedges (genus Carex), this 1.4 m 1.5 m
European species has triangular stems and
separate male and female flower clusters.

spike of female
flowers

crimson CHINESE WATER

central leaves giant spike of CHESTNUT
more than Eleocharis dulcis PAPYRUS SEDGE
3,000 flowers f: Cyperaceae Cyperus papyrus
Native to Asia, this f: Cyperaceae
wetland sedge has This tall African perrenial is a
clumps of tubular wetland plant. The Ancient
stems. It is cultivated Egyptians wrote on a paperlike
evergreen for its edible material called papyrus made
variegated foliage underwater tubers. from its leaves.

spine-edged, yellow bracts

33 ft 3¼ ft striped leaf around small,
10 m 1m white flowers

stiff, spiny
leaves
ZEBRA PLANT HAHN'S CATOPSIS
Aechmea chantinii Catopsis paniculata
f: Bromeliaceae f: Bromeliaceae
This large, strap-leaved This epiphytic
epiphytic bromeliad comes bromeliad is native to
from the rainforests of the cloud forests of
S. America. It is pollinated S. Mexico and
by hummingbirds. C. America.
QUEEN OF THE ANDES
Puya raimondii 20 in
f: Bromeliaceae 50 cm
Native to the C. Andes, the gray-green
rosette of leaves with
world’s largest bromeliad waxy texture
strap-shaped leaves produces a single colossal
flower spike after many years’
growth and then dies.
 POA LES
146
SOFT RUSH
Juncus effusus
f: Juncaceae
FLOWERING PLANTS • MONOCOTS

This very widespread

rush thrives in damp, 23½ in
infertile soils. Its 60 cm
cylindrical stems are
filled with spongy
tissue, or pith. LARGE
QUAKING
GRASS
Briza maxima
f: Poaceae 6 ft 23½ in 30 in
This Mediterranean 1.8 m 60 cm 75 cm
annual grass gets its
name from the way
its fine-stalked
flowerheads shake at
the slightest breeze.
5 ft 4¼ ft
1.5 m 1.3 m FALSE OAT-GRASS SAND COUCH CRESTED DOG’S-TAIL
Arrhenatherum elatius Thinopyrum junceiforme Cynosurus cristatus
f: Poaceae f: Poaceae f: Poaceae
Common across its native This tough grass grows on the Low-growing apart from its
Europe, this tall-flowered seaward side of European sand flower stalks, this perennial
wild relative of the oat has dunes. Its roots, or rhizomes, help grass from Europe and W. Asia
now spread to many other to hold the windblown sand resists trampling and is often
parts of the world. together to form foredunes. used for lawns.
COCK’S-FOOT
Dactylis glomerata 8¼ ft
f: Poaceae 2.5 m
Often grown in hayfields young seeds
and pastures, this common
grass with distinctive tufty dried
seedhead
flowerheads is native JOB’S TEARS
to Eurasia and N. Africa. Coix lacryma-jobi
f: Poaceae
Native to C. Asia, this
grass is grown around
the tropics as a cereal
or fodder crop. The
hard seeds of wild
plants are used as beads.

evergreen
leaves
23½ in 20 ft
60 cm 6m
tall, broad
leaf

HARE’S TAIL GIANT REED

Lagurus ovatus Arundo donax
23 ft f: Poaceae f: Poaceae
7m The distinctive, soft-haired Originally from C. Asia, this
flowerheads of this mainly outsize marshland grass has
coastal Mediterranean grass been widely planted elsewhere.
make it popular in dried Its woody stems have many
flower arrangements. traditional uses.

BLACK BAMBOO
Phyllostachys nigra
f: Poaceae awns protect seeds
Like all the bamboos, this black-stemmed
species has hard, woody stems. It is native to THREE-AWNED
E. and S. China. Bamboos are the fastest GOAT GRASS
growing woody plant, growing up to Aegilops neglecta
35 in (90 cm) a day. f: Poaceae
A relative of wheat, this
low-growing, drought-resistant
annual grass is native to the
Mediterranean and Middle East.

14 in
finger-thick cane 35 cm
 147
awn

FLOWERING PLANTS • MONOCOTS

20 ft
6m
COMMON REED
Phragmites australis
f: Poaceae
Widespread in both temperate
and tropical regions, this
shallow-water grass can 3¼ ft 4 ft 6 ft 32 in 3¼ ft
colonize large areas via its 1m 1.2 m 1.8 m 80 cm 1m
creeping horizontal stems.

SWEET VERNAL GRASS MARRAM GRASS OAT BARLEY BREAD WHEAT

Anthoxanthum odoratum Calamagrostis arenaria Avena sativa Hordeum vulgare Triticum aestivum
f: Poaceae f: Poaceae f: Poaceae f: Poaceae f: Poaceae
Native to Eurasia, this This tough grass from This cultivated grass, Originating in the The world’s top-tonnage
early-flowering grass Europe flourishes on grown widely as a ancient Near East, this cereal, this species originated
contains a chemical called sand dunes, which cereal crop for cultivated cereal grass in the ancient Near East.
coumarin that makes it its long underground livestock and human is notable for the long, It is a hybrid between
smell pleasantly of stems and roots help consumption, thrives hairlike bristles (awns) the wild and earlier
new-mown hay. to stabilize. in cool, wet climates. on its flowerheads. cultivated varieties.

10 ft 10 ft
3m 3m

16 in
40 cm

CREEPING BENT-GRASS
Agrostis stolonifera
f: Poaceae 6 ft
1.8 m
This common perennial grass
with feathery flowerheads is
found worldwide. It spreads 20 ft
using creeping horizontal 6m
stems, or stolons.
CITRONELLA RICE CORN SUGARCANE
Cymbopogon nardus Oryza sativa Zea mays Saccharum officinarum
f: Poaceae f: Poaceae f: Poaceae f: Poaceae
A type of lemongrass, Native to E. Asia, this This major food crop was Possibly related to corn, this
citronella originates from major grain crop of first cultivated in ancient tropical cultivated grass may
tropical Asia. It yields an oil warmer regions is usually Mexico. It has male or have originated in New
used in perfumes and for cultivated in shallow water female flowerheads – edible Guinea. Sugar is extracted
repelling insects. or in flood-prone areas. corn is the female flower. from its thick stems.

PERENNIAL RYE-GRASS
Lolium perenne single stem
dense white
f: Poaceae flowerheads
Often used in pastures, lawns,
and sports fields, this common
Eurasian grass has now spread
around the world. PAMPAS GRASS
Cortaderia selloana
f: Poaceae
This tall grass from
southern S. America is
a popular ornamental
species. It has become
invasive in some areas.
10 ft 35 in 32 in
3m 90 cm 80 cm
10 ft
3m
VETIVER YORKSHIRE FOG
Chrysopogon zizanioides Holcus lanatus
f: Poaceae f: Poaceae
Native to India, this tropical A common European grass
grass is widely cultivated for found in damp meadows,
its fragrant oil and to bind Yorkshire fog has leaves
soils to prevent erosion. with soft, velvety hairs.
 POA LES ZINGIBERALES
148 Many species in this mainly tropical order of eight families
GIANT FEATHER
GRASS grow giant leaves at the end of stalks. Although there are
8 ft Stipa gigantea no true woody trees, some species such as the banana plant
2.5 m f: Poaceae
FLOWERING PLANTS • MONOCOTS

Gardeners cultivate this grow very large. Many Zingiberales have showy flowers and
tall grass for its showy foliage and have become ornamentals. The ginger family,
flowerheads, which Zingiberaceae, the largest in the order, includes several other
persist until winter. It important spice plants beside ginger itself.
is native to Spain,
Portugal, and
Morocco.

6½ ft
2m
INDIAN SHOT
Canna indica
f: Cannaceae
This tropical American
plant has unusual flowers in
which some of the petals
16 in 20 in are fused with pollen-
female flower 40 cm 50 cm producing stamens.
There are many
cultivated varieties.
NEVER-NEVER PLANT ETERNAL FLAME
Ctenanthe amabilis Goeppertia crocata
f: Marantaceae f: Marantaceae
Native to forest floors in the Brazilian Related to Ctenanthe and Maranta, this Brazilian
tropics, this warmth-loving plant requires species has more striking flowers than its
high humidity if grown in cultivation. relatives, although living in similar forest habitats.

male
flowerhead
23½ in 12 in
60 cm 30 cm

5 ft
1.5 m

BRANCHED BUR-REED
Sparganium erectum
f: Typhaceae
10 ft Separate ball-shaped clusters of
3m male and female flowers develop
on the same flower stalk in this SPIRAL FLAG SILVER-VEINED PRAYER
widespread wetland species of the Chamaecostus cuspidatus PLANT
northern hemisphere. f: Costaceae Maranta leuconeura
This orange-flowered plant f: Marantaceae
with dark green, glossy This Brazilian forest plant folds its
leaves is native to tropical leaves together at night to conserve
E. Brazil. It is also grown moisture. Cultivated varieties have
as an ornamental. strikingly patterned foliage.

40 ft/12 m

5 ft/1.5 m

30 ft each yellow bract

12 in 9m protects four or
30 cm five tiny flowers
CATTAIL
Typha latifolia YELLOW-EYED BANANA CHINESE YELLOW
f: Typhaceae GRASS ENSET Musa acuminata BANANA
This widespread Xyris sp. Ensete ventricosum f: Musaceae Musella lasiocarpa
wetland plant has f: Xyridaceae f: Musaceae The seedless hybrid of f: Musaceae
a distinctive, This genus of grasslike This African-relative of the Asian wild ancestors, Possibly extinct in the
cigar-shaped female plants, widespread in banana has long been cultivated cultivated bananas wild, this mountain species
flowerhead. The male warmer regions of the in Ethiopia for its nutritious grow sterile male from China produces a
flowers grow world, bears small, yellow rootstock and stem, not its flowers at the end of yellow flower spike that
above it in a tuft. flowers on slender stems. (inedible) fruits. their fruiting branches. can last for months.
violet-striped
white petal
white
flower
149

lanceolate

FLOWERING PLANTS • MONOCOTS

leaf
5 ft 12 in
1.5 m 30 cm

18 ft CARDAMOM LESSER AROMATIC

5.5 m Elettaria cardamomum GALANGAL GINGER
f: Zingiberaceae Alpinia officinarum Kaempferia galanga
This widely cultivated f: Zingiberaceae f: Zingiberaceae
seed tropical plant is native Native to E. Asia, this Native to tropical Asia,
pod to forests in S. India and relative of ginger this short-stemmed
Sri Lanka. Its dried, develops a swollen species develops small
unripe fruits provide underground stem and is flowers and is grown as
the spice cardamom. similarly used as a spice. an ornamental.

paddle-shaped rhizome
leaf, green at tip, and roots
yellow at leaf stem
3¼ ft/1 m

fleshy
rhizome
TURMERIC
symmetrical Curcuma longa
fan-shaped tree f: Zingiberaceae
Native to S.E. Asia, this large-leaved
plant develops a swollen underground
stem from which the yellow spice stem with
turmeric is obtained. flowerhead

3¼ ft
edible 1m
rhizome

GINGER
Zingiber officinale
f: Zingiberaceae
The spice ginger comes from
the underground stems of this
cultivated S.E. Asian plant,
which no longer grows wild.

purple flowers
develop in summer

10 in 12 ft
25 cm 4m
50 ft 6½ ft
15 m 2m
HUME’S ROSCOEA HELICONIA
TRAVELER’S TREE Roscoea humeana STRICTA
Ravenala madagascariensis BIRD-OF-PARADISE f: Zingiberaceae f: Heliconiaceae
f: Strelitziaceae Strelitzia reginae This relative of the In its natural habitat, this
Native to open forests in f: Strelitziaceae ginger plant grows large-leaved tropical plant
Madagascar, this relative Orange-and-blue bird-pollinated unusual orchidlike from northern areas of
of Strelitzia is pollinated flowers open one at a time from a flowers. Its native S. America is pollinated
by lemurs in its beaklike sheath in this South habitat is the mountains by nectar-sipping
natural habitat. African plant. of S.W. China. hummingbirds.
150 EUDICOTS
Almost three-quarters of the world’s stems as they grow. This “secondary growth” is DIVISION ANGIOSPERMAE
known flowering plants are classified absent in most monocots, which is why eudicots
FLOWERING PLANTS • EUDICOTS

CLADE EUDICOTS

in the eudicot grouping, which evolved make up most of the world’s flowering shrubs ORDERS 44

more than 125 million years ago. and trees. Below ground level, most eudicots have FAMILIES 312

a taproot, with smaller roots branching off it. SPECIES Over 210,000

Eudicots get their name from the cotyledons, Eudicot flowers usually have parts in fours or fives,
or seed-leaves, in the seed before it germinates. rather than the threes found in monocots, with sepals
Unlike monocots, which have a single seed-leaf, and petals that look different, both in color and in
eudicots have two. They include an enormous shape. Each species has its own distinctive kind of D E B AT E
variety of plants, from agricultural weeds to pollen, but the grains always have three pores; F O U R - W AY S P L I T
towering rainforest trees, and they are of huge monocot pollen grains only have a single opening. For many years, all flowering plants
economic importance, as well as being prized were classified in two groups—the
as garden flowers. Many eudicots are annuals, a v i ta l ro l e monocotyledons (monocots) and
with short lifespans measured in months or The fossil pollen record suggests that the eudicots the dicotyledons (dicots)—based
even weeks. Others are biennials and perennials, diverged from other flowering plants some 125 on the number of their seed-leaves.
which live from two years to a century or more. million years ago. Since then, they have colonized However, DNA analysis, combined
Despite their great diversity, eudicots have many every habitat on land, although they are less with palynology—the study of
physical features in common. Their leaves often common as waterplants. Their importance to pollen—has shown this classification
does not fully reflect plant evolution.
have a netlike tracery of veins, unlike the parallel animals is impossible to overestimate. Countless
As a result, flowering plants are now
veins of monocots, and their stems have a well- species—with the notable exception of grazing split into four overall groups: basal
developed vascular system, arranged in rings, for animals that feed on grasses (monocots)—depend angiosperms, magnoliids, monocots,
transporting water and sap. As well as getting on them for food and shelter, and in return, many and the eudicots, or true dicots.
taller, woody species thicken and strengthen their act as pollinators, or help to spread their seeds.

NEW PERSPECTIVES BUXALES

Computer analysis of DNA data has suggested new ways of Containing just a single family of about 120
subdividing the eudicots. These new hierarchical groupings, species, the order Buxales is found in temperate,
known as clades (see pp.30–31), give insights into the subtropical, and tropical regions. Most of the
relationships between species and how they evolved, but they members are trees or shrubs, with simple, 33 ft
are difficult to interpret in terms of physical characteristics of evergreen leaves and separate male and female 10 m
the resulting groups and do not fit neatly into the traditional flowers on the same plant. Many species are grown
taxonomic hierarchy. This book is organized around orders and ornamentally and box wood is used for carving.
families, which still provide the clearest way to understand the
diversity of flowering plants. However, the table below shows
how they “nest” together in the proposed new system of clades.

EUDICOTS (pp.150–209) 5 ft
1.5 m
(BUXALES—DIPSACALES)

CORE EUDICOTS (pp.155–209)

(GUNNERALES—DIPSACALES)

SUPERROSIDS (pp.156–181)
(SAXIFRAGALES—SAPINDALES)
ROSIDS (pp.158–181)
(VITALES—SAPINDALES)

As its name suggests, Japanese

rose belongs to the rosid group.

SUPERASTERIDS (pp.182–209)
(CARYOPHYLLALES—DIPSACALES)
ASTERIDS (pp.192–209)
(CORNALES—DIPSACALES)
SWEET BOX
Oxeye daisy is an asterid within Sarcococca hookeriana BOXWOOD
the larger superasterid group. f: Buxaceae Buxus sempervirens
This species bears clusters f: Buxaceae
of small, fragrant flowers in Native to rocky woods and scrub from Europe
winter, and is found in shady to N. Africa, this evergreen shrub or small
habitats in W. China. tree is often grown in gardens for topiary.
PROT E A L E S styles HONEYSUCKLE
GREVILLEA
Of the four Proteales families, the largest is the Grevillea juncifolia 151
23 ft/7 m f: Proteaceae
Proteaceae, evergreen trees and shrubs from Found in drier inland regions
the southern hemisphere. The Platanaceae are across Australia, this erect

FLOWERING PLANTS • EUDICOTS

northern hemisphere deciduous trees, and the shrub has narrow, gray-green
leaves and tight heads of
Nelumbonaceae are aquatic plants orange flowers with long,
from Asia, Australia, and seed 20 ft yellow styles.
heads
North America. 6m

10 ft
3 ft 3m
outer bract 1m
RED SILKY
OAK
lotus Grevillea banksii
leaf
f: Proteaceae
SACRED LOTUS Cultivated as an
Nelumbo nucifera ornamental species for its
f: Nelumbonaceae striking “bottle-brush”
Growing in shallow flowers, this native of
freshwater habitats in parts N.E. Australia grows in
of Asia and Australia, sacred woods and open habitats.
lotus carries large, fragrant
flowers on long stalks WARATAH
above the water. Telopea speciosissima
f: Proteaceae
From dry woodland in New South
seed pod Wales, Australia, common waratah
seen from 6½ ft/2 m is also known as Sydney waratah.
above
6½ ft
2m
collection of
flowers at center KING PROTEA
of flowerhead Protea cynaroides
f: Proteaceae
This species is native to hillsides and scrub in
South Africa. Clusters of small flowers are
enclosed by petallike bracts. MOUNTAIN DEVIL 5 ft
1.5 m
Lambertia formosa
f: Proteaceae
From coastal heathland to mountain forests PINCUSHION PROTEA
in New South Wales, Australia, this species Leucospermum cordifolium
has red-tinged bracts surrounding f: Proteaceae
simple, the flower clusters. This shrub of sandstone
evergreen leaf soils in South Africa’s
Cape Province is noted
for its brightly colored,
spherical flowerheads.

fruit takes
about six
months to ripen

6½ ft/2 m
50 ft
15 m NARROW-LEAF
DRUMSTICKS
SAW BANKSIA Isopogon anemonifolius
Banksia serrata f: Proteaceae
f: Proteaceae Spherical, yellow flowerheads
Saw Banksia grows in woodland above feathery leaves characterize
and scrub in E. Australia. Its bark this shrub, native to dry woodland
is fire-resistant, enabling it to and heathland in New South
survive bushfires. Wales, Australia.
thick,
stiff-textured,
leafy spike of red, or unripe broad leaf,
occasionally yellow nuts similar
or white flowers to maple
157 ft/48 m
30 ft
10 m MACADAMIA flower LONDON PLANETREE
NUT spike Platanus x hispanica
Macadamia integrifolia f: Platanaceae
CHILEAN FIRE BUSH f: Proteaceae Planted in London since the 17th
Embothrium coccineum Native to coastal century, this deciduous hybrid arose
f: Proteaceae rainforest in E. Australia, from the cross-fertilization of two
From forest and open habitats in S. Chile and Argentina, macadamias are species of Platanus in Spain. Tolerant
this species is cultivated for its flame-colored, 30-ft cultivated for their 50 ft of pollution, it is widely planted in
flowers, and thrives in sheltered gardens. edible nuts. 15 m urban parks and streets.
 RANUNCULALES oval, toothed
leaf
OREGON GRAPE
Berberis aquifolium
152 The order Ranunculales, consists of seven families. f: Berberidaceae
Flowering in springtime, Oregon
Four are relatively obscure, with few species, grape is a suckering evergreen shrub
but the buttercup family, Ranunculaceae—after from N.W. US. It grows in
which the order is named—contains about 3,200 shady habitats.
species. Together with members of the poppy and
barberry families, these include some of the most
common agricultural weeds worldwide and many 10 ft
familiar garden plants. 3m

SACRED BAMBOO
Nandina domestica
f: Berberidaceae oblong berry
Also called heavenly bamboo, this shrub from
mountain valleys in China and Japan is
unrelated to true bamboos.
BARBERRY 5 ft
Berberis vulgaris 1.5 m
f: Berberidaceae
A European species of hedges
and scrub, barberry has
distinctive triple spines, hanging
flower clusters, and red berries.
bright red
berries

33 ft 20 ft
BISHOP’S HAT 10 m 6m
Epimedium davidii
f: Berberidaceae
This evergreen species
evergreen from W. China grows
leaves in woods and scrub. The 12 in
6½ ft young leaves are coppery, 30 cm
2m later turning green.

clustered
flowers MOONSEED
twining stem Menispermum canadense
f: Menispermaceae
20 in Although the fruits of this climber
50 cm resemble black grapes, they are
extremely poisonous. The species
inhabits woodland and banks of
streams in Canada and the US.

CAROLINA
SNAILSEED
Cocculus carolinus
f: Menispermaceae
This climbing woodland
13 ft species from S.E. US has
4m tiny flowers; the males and
females are borne on
separate plants.
CHOCOLATE
VINE
fragrant flower
Akebia quinata 30 in
f: Lardizabalaceae 75 cm
Bearing sprays of scented flowers in
spring, chocolate vine inhabits forest
edges in China, Korea, and Japan.

16 in/40 cm 33 ft/10 m creamy-

colored
flower
JAPANESE pinnate
STAUNTON VINE leaf
LEONTICE MAY APPLE Stauntonia hexaphylla CLIMBING CORYDALIS
Leontice leontopetalum Podophyllum peltatum f: Lardizabalaceae Ceratocapnos claviculata
f: Berberidaceae f: Berberidaceae Native to woodland in f: Papaveraceae
Native to cultivated ground A native of N. America, Japan and South Korea, This scrambling annual
and dry hillsides in N. Africa May apple—also this vigorous evergreen species from W. Europe grows
and E. Mediterranean known as American climber has woody in woods and shady habitats
countries, leontice grows mandrake—grows in stems and fragrant on acid soils, supporting
from a tuber. open woodland. flowers. itself by leaf tendrils.
 YELLOW
HORNED POPPY
Glaucium flavum 153
f: Papaveraceae
Native to W. Europe and
the Mediterranean, this

FLOWERING PLANTS • EUDICOTS

biennial or perennial grows
on coastal shingle. Its long,
curved fruits are distinctive.

35 in 36 in
90 cm 90 cm
GREATER CELANDINE
Chelidonium majus
f: Papaveraceae 4 ft
Formerly grown by herbalists, 1.2 m
greater celandine is native to Europe
and N. Asia, growing in woodland,
12 in scrub, and rocky places.
30 cm
BLEEDING HEART
Lamprocapnos spectabilis
YELLOW CORYDALIS f: Papaveraceae
Pseudofumaria lutea Named for its heart-shaped flowers, this
f: Papaveraceae species grows in damp woodland edges
This European species grows in Siberia, N. China, and Korea.
on walls and in stony and
rocky places. It spreads
vigorously by seed.

spurred petal

CALIFORNIA POPPY MATILIJA POPPY

12 in Eschscholzia californica Romneya coulteri
30 cm f: Papaveraceae f: Papaveraceae
Native to open habitats in A species of scrub and grassland
western US and Mexico, this in California and Mexico, this
poppy is grown in gardens for poppy has fragrant flowers and 6½ ft
its brightly colored flowers. is often grown in gardens. 2m

20 in
50 cm

COMMON FUMITORY
Fumaria officinalis
f: Papaveraceae
Found in Europe and
pollen produced N. Africa on cultivated and
by a ring of dark waste ground, this species
cental anthers usually grows on light soils.

yellow or
orange flower

16 in 23½ in
40 cm 60 cm

23½ in
cut stem exudes 60 cm
poisonous latex OPIUM pinnate leaf
POPPY
Papaver somniferum
SICKLEFRUIT COMMON POPPY f: Papaveraceae WELSH POPPY
HYPECOUM Papaver rhoeas Grown as the source of Papaver cambrica
Hypecoum imberbe f: Papaveraceae opium, heroin, and f: Papaveraceae
f: Papaveraceae Arable and waste ground are home to poppy seed, this Inhabiting shady, rocky
This Mediterranean native grows this poppy, native to Europe, N. Africa, inhabits cultivated and places in hilly areas and often
in cultivated and waste ground and parts of Asia. It is used as a symbol disturbed ground 5 ft grown in gardens, Welsh
and on walls. to commemorate World War I. in Eurasia. 1.5 m poppy is native to W. Europe.
 RANUNCULALES seedhead
154
cup-shaped
flower
FLOWERING PLANTS • EUDICOTS

WINTER ACONITE
6 in Eranthis hyemalis
15 cm f: Ranunculaceae
This tuberous perennial
species is found in damp
woodland and shady places in
C. Europe. It flowers in late
winter (hence its common
name) and early spring.
climbing stems
TRAVELER’S JOY
Clematis vitalba
f: Ranunculaceae
Also known as “old man’s beard” for its
gray, feathery fruits, traveler’s joy is a
woody climber found in wood edges and
6½ ft hedgerows in Europe and N. Africa. 100 ft
2m
30 m

thin-stemmed
foliage
five petal-
like sepals
16 in 3¼ ft
40 cm 1m

PHEASANT’S EYE MEADOW BUTTERCUP

Adonis annua Ranunculus acris
f: Ranunculaceae f: Ranunculaceae
An increasingly uncommon Found in much of Europe
annual species, pheasant’s and temperate regions of 3¼ ft
eye is found in cultivated W. Asia, this perennial species 1m
and waste ground in is found in damp grassland.
W. Europe and N.W. Africa.

SCARLET LARKSPUR ROCKET LARKSPUR

Delphinium cardinale Delphinium ambiguum
f: Ranunculaceae f: Ranunculaceae
Also known as cardinal Growing on cultivated and
larkspur where it grows in disturbed ground on light soils,
California, and Baja rocket larkspur is an annual native
California, Mexico, this to the Mediterranean region.
short-lived perennial is found
on dry hillsides.
5 ft 5 ft
23½ in 20 in 1.5 m 1.5 m
60 cm 45 cm

CROWN ANEMONE
KINGCUP Anemone coronaria
Caltha palustris f: Ranunculaceae
f: Ranunculaceae Mediterranean countries are
Kingcup, or marsh marigold, home to crown anemone, a
inhabits marshes, ditches, tuberous perennial species
wet woods, and wet grassland which grows on stony
in much of Europe, Asia, hillsides, roadsides, and flower forms on
and N. America. cultivated ground. a long spike

12 in
30 cm
stem, leaves, and
roots are toxic
20 in
50 cm
MONKSHOOD
Aconitum napellus
f: Ranunculaceae
FIELD LOVE-IN-A-MIST Named for its
PASQUE FLOWER Nigella arvensis COMMON MEADOW RUE helmeted flowers,
Pulsatilla vulgaris f: Ranunculaceae Thalictrum flavum monkshood inhabits
f: Ranunculaceae This annual species is native to f: Ranunculaceae damp woods and
Chalk slopes with short turf are cultivated and disturbed ground This perennial species grows in streamsides in
the home of pasque flower, in C. and S. Europe. It sprouts wet meadows and fens beside Europe. It is an
a perennial species native in gardens elsewhere from fresh water in Europe and extremely poisonous
to Europe and W. Asia. spilled bird seed. temperate regions of Asia. perennial species.
 MOUSETAIL
Myosurus minimus
f: Ranunculaceae
The “tail” of mousetail is
the elongated fruit that
develops from its flower.
This species grows in
bare, damp ground across
Europe, Asia, and leafless stem
N. America. topped by a
single flower

28 in
70 cm
4 in fruit developing
10 cm from a flower

6 in
15 cm
green
flower GLOBEFLOWER HEPATICA
Trollius europaeus Hepatica nobilis
f: Ranunculaceae f: Ranunculaceae
Native to Europe and Native to much of Europe, this
W. Asia, globeflower is a perennial woodland species has
perennial species found in distinctive semievergreen,
damp mountain pasture. three-lobed leaves.

long, distinctive
spur
3¼ ft
1m
narrow,
23½ in threadlike leaf
60 cm

toothed leaf

leaves

HELLEBORE
Helleborus lividus COLUMBINE
f: Ranunculaceae Aquilegia vulgaris
The Balearic Islands, particularly f: Ranunculaceae
Mallorca, are home to this perennial A perennial species native to much of Europe,
species. Hellebore inhabits woods N. Africa, and temperate regions of Asia, columbine
and rocky hillsides. is found in shady and damp chalky habitats.

GUNNERALES DILLENIALES
The two families which make up the Gunnerales order were The order Dilleniales contains only the Dilleniaceae, a
previously classified in different orders, as they are visually mainly tropical family of around 330 species of trees,
very different. However, genetic analysis has recently shown shrubs, and climbers. Most species have alternately
the two families to be closely related. The family Gunneraceae arranged leaves and bisexual flowers (with male and
consists of a single genus of large herbaceous plants growing female parts), with five sepals, five petals, and numerous
in damp habitats, whereas Myrothamnaceae species are stamens. Some produce dry fruits that open to shed their 23 ft
inhabitants of African deserts. Gunnera species are often grown seeds, others have berries. A few are grown as ornamentals 7m
in gardens as ornamental plants. or for lumber used in construction and boat building.

8 ft
2.5 m 10 ft
3m
SIMPOH AIR
GUNNERA Dillenia suffruticosa
Gunnera manicata GOLDEN f: Dilleniaceae
f: Gunneraceae GUINEA VINE A large and vigorous
Massive leaves and tall Hibbertia scandens evergreen shrub endemic to
flower spikes characterize f: Dilleniaceae Malaysia, Sumatra, and
this perennial species. A vigorous climber or scrambler, this Borneo, simpoh air is found
It is native beside fresh evergreen shrub grows near the E. on swampy ground and
water in S. Brazil. coast of Australia and in New Guinea. forest edges.
 SA X I F R AGA L ES MEXICAN FIRECRACKER
Echeveria setosa
156 Of the 15 diverse families in the f: Crassulaceae
Named for its flame-colored
Saxifragales, five have just two flowers, this Mexican species has
members each and only three have 4 in rosettes of succulent, blue-
FLOWERING PLANTS • EUDICOTS

more than 500 species. Best known 10 cm green, variably downy leaves.
is the saxifrage family, after which
the order is named. The Latin saxifraga
literally means “rock breaker,” as these
plants often grow in cracks in rocks and
walls. The largest family is the stonecrops,
Crassulaceae, which includes many succulent or
water-retaining plants, adapted to dry conditions.
COMMON HOUSELEEK
Sempervivum tectorum
f: Crassulaceae
Originating from mountains in
C. Europe, houseleek is often
planted on roofs and walls. This
plant forms dense mats of
succulent rosettes.
20 in
50 cm SAUCER PLANT
Aeonium tabuliforme
f: Crassulaceae
This native of cliffs on the
N. coast of Tenerife, Canary
Islands, forms a flat,
succulent leaf rosette.
ORPINE
Hylotelephium telephium 23½ in
f: Crassulaceae 60 cm
Rocky terrain, woods, and NAVELWORT 20 in
hedgebanks are the habitats of Umbilicus rupestris 50 cm
orpine, a species native to Eurasia f: Crassulaceae
and introduced to N. America. 23½ in This species has round,
60 cm fleshy leaves, each with a
central dimple. Navelwort
inhabits rocks and walls in
S. Europe and N. Africa. 16 in
40 cm

16 in
40 cm

FLAMING KATY
Kalanchoe blossfeldiana
f: Crassulaceae
Arid areas of Madagascar are home
to this bushy species with glossy,
succulent leaves and brightly
colored flowers. ROSEROOT
Rhodiola rosea
f: Crassulaceae
BLACKCURRANT Inhabiting mountain rocks and coastal cliffs,
Ribes nigrum roseroot is a fleshy species found in the Arctic and
f: Grossulariaceae alpine regions in Europe, N. America, and Asia.
This species grows in damp
woods in much of Europe 6½ ft
2m
and C. Asia. Blackcurrant
is cultivated for its
flavorful berries.

3¼ ft/1 m

WESTERN WATER
BUFFALO CURRANT MILFOIL
Ribes aureum Myriophyllum hippuroides
f: Grossulariaceae f: Haloragaceae
Scented flowers and thornless This aquatic species has finely
stems characterize buffalo divided leaves. It is native to
currant, an inhabitant of rocky freshwater habitats in
6½ ft/2 m and sandy areas in C. US. western N. America.
 40 ft PARROTIOPSIS
12 m Parrotiopsis
jacquemontiana
f: Hamamelidaceae
Instead of petals, the
flowers of this forest
species from the
W. Himalayas have leaves changing
white bracts, or color
CHINESE SWEETGUM modified leaves. buds and
Liquidambar formosana flowers
f: Altingiaceae
A deciduous tree of damp woodland in 33 ft/10 m VIRGINIAN WITCH HAZEL
S.W. Asia, this species is noted for its Hamamelis virginiana
brightly colored fall foliage. f: Hamamelidaceae
20 ft/6 m The flowers of this witch hazel are fragrant
CHINESE ASTILBE and the leaves turn yellow in the fall. It
Astilbe rubra inhabits woodland in eastern N. America.
f: Saxifragaceae
3¼ ft/1 m With distinctive plumelike clusters of flowers, this
moisture-loving plant grows in damp woodland and
by streams from Myanmar to Siberia. 8 in
20 cm 50 ft/15 m

PERSIAN IRONWOOD
Parrotia persica
f: Hamamelidaceae
Native to forests in the Caucasus region and northern
MOTHER YELLOW Iran, Persian ironwood is a winter-flowering deciduous
OF THOUSANDS SAXIFRAGE tree with brilliantly colored fall leaves.
Saxifraga stolonifera Saxifraga aizoides
f: Saxifragaceae f: Saxifragaceae
Native to shady habitats Streamsides and wet
in China and Japan, this stony ground on
perennial spreads by mountains are home to 23½ in/60 cm
producing rooting this cushion-forming
plantlets at the end species from Europe,
of threadlike N. America, and
runners. W. Asia.
ALUMROOT
12 in Heuchera americana
30 cm f: Saxifragaceae
flowers form Inhabiting rocky woodland in N. America,
in a flattish-
topped cluster alumroot has glossy leaves which are
mottled when young.

ELEPHANT’S EARS
Bergenia stracheyi
f: Saxifragaceae
An inhabitant of damp woodland and
meadows in the W. Himalayas and 28 in
Afghanistan, elephant’s ears has fragrant 6 in/15 cm 70 cm
flowers and large, glossy leaves.

large,
glossy leaf
12 in
30 cm
OPPOSITE-LEAVED COMMON PEONY
GOLDEN-SAXIFRAGE Paeonia officinalis
Chrysosplenium oppositifolium f: Paeoniaceae
f: Saxifragaceae A herbaceous species
The procumbent stems of this found in woodland,
species form extensive patches meadows, and scrub
in damp, shady habitats in in parts of S. Europe,
W. and C. Europe. common peony is
known for its
PIGGYBACK showy flowers.
PLANT
Tolmiea menziesii
f: Saxifragaceae
Young plantlets growing on 28 in
leaf bases give piggyback 70 cm
plant its name. It is a hairy
perennial species found in
damp, shady places
in N. America.
 V I TA L E S
158 This order consists of a single family, the Vitaceae or
grape family. It contains 14 genera and 850 species, foliage turns red
in the fall
including the important grapevine and the ornamental
Virginia creeper. Members of the Vitaceae are mainly
GRAPEVINE native to the tropics or warm temperate regions. Mostly
Vitis vinifera vines or lianas, they usually have swollen nodes (where
f: Vitaceae leaves fork from the stem) and tendrils for climbing.
115 ft Humans have used grapes to make wine, food, Their flowers are usually held in
35 m and medicine since neolithic times. Very
widespread, the grapevine is native to the flat-topped clusters.
Mediterranean, Europe, and Asia.
VIRGINIA CREEPER
fruits smaller than Parthenocissus quinquefolia
domesticated varieties f: Vitaceae
This prolific climber from
N. and C. America clings onto 100 ft
CRIMSON GLORY smooth surfaces using the 30 m
VINE
Vitis coignetiae adhesive pads on its
f: Vitaceae small, forked tendrils.
Cultivated for its giant,
dimpled leaves
(12 in/30 cm across) and
fall color, this deciduous
climber is native to 50 ft
temperate Asia. 15 m

GERANIALES
Two families comprise the Geraniales order. The
Geraniaceae or geranium family contains about 800
species, including 400 cranesbills in the genus Geranium.
Many of the 200 species in the African genus Pelargonium
are of horticultural importance, including the garden 8 ft
2.5 m
plants called geraniums. The new family Francoaceae
encompasses four previously recognized families, mainly GIANT HONEYBUSH
Melianthus major
of trees and shrubs from Africa. f: Francoaceae
Nectar drips from the bronze
flower spikes of this South African
native. Touching the leaves causes
them to give off a strong odor.

12 in
30 cm 32 in
80 cm

20 in
50 cm
8 in
20 cm

APPLE PELARGONIUM HERB ROBERT

Pelargonium odoratissimum Geranium robertianum MEADOW CRANESBILL ROCK
f: Geraniaceae f: Geraniaceae Geranium pratense STORKSBILL
This perennial originates Widespread in the f: Geraniaceae Erodium foetidum
from South Africa and has northern hemisphere, Preferring grasslands on chalky f: Geraniaceae
spreading flower stalks. It is this sprawling species soil, this perennial is native to Storksbills take their name
cultivated for geranium oil, has red stems and long Europe and Asia. Its flowers are from the shape of their
which has a strong scent flower stalks. It has a pollinated by honey bees and seeds. This French native is
of apple and rose. strong, mousey smell. other wild bee species. cultivated in rock gardens.
M Y RTA L E S SWAMP
LOOSESTRIFE
PURPLE LOOSESTRIFE
Lythrum salicaria
The nine families in the Myrtales are most Decodon verticillatus f: Lythraceae 159
f: Lythraceae Numerous purple-red, square
common in warmer regions. The 5,800 species Native to N.E. America, stems grow from the woody,
in the Myrtaceae or myrtle family provide this shrub grows in creeping rootstock of this

FLOWERING PLANTS • EUDICOTS

essential oils, spices, and fruits such as guava. swamps. It has arching perennial. Native to Europe,
stems with leaves in Asia, S.E. Australia, and
They include more than 700 species of eucalypts whorls of three, and red N.W. Africa, it can
from Australia and New Guinea. The Lythraceae or purple flowers up to be invasive.
or loosestrife family consists mainly of tropical 1 in (2.5 cm) across.
trees and shrubs, producing fruits such as 8 ft
pomegranate and various dyes, and the 2.5 m triangular
Melastomataceae is a pan-tropical family of leaf
multi-seeded 23 ft
about 4,500 trees, shrubs, and climbers. fruit develops 7m
from flower

20 ft
6m
30 in 5 ft
75 cm 1.5 m

20 ft
6m
CRAPE MYRTLE POMEGRANATE
Lagerstroemia indica HENNA Punica granatum
f: Lythraceae Lawsonia inermis WATER CHESTNUT f: Lythraceae
Blooming for up to 120 days, this f: Lythraceae Trapa natans This spiny, shrubby tree, from
tree is native to China, Korea, Native to N. Africa and the Middle f: Lythraceae S.W. Asia, is widely cultivated
and Japan. Its bark is smooth, East, henna’s leaves make a This floating plant is from Europe and in the Mediterranean for its
mottled, pinkish-gray, and reddish-brown dye and its fragrant Asia. It has an edible, starchy seed in a pulpy fruits, containing
is shed each year. flowers yield essential oils. nut with four hornlike barbed spines. numerous seeds.

35 in
90 cm
100 ft
30 m

59 ft
18 m

CIGAR FLOWER
Cuphea ignea
f: Lythraceae
A densely branched, perennial INDIAN ALMOND
shrub, cigar flower is a garden Terminalia catappa
and house ornamental, native f: Combretaceae
to Mexico. Its fruits are RANGOON CREEPER Found on coasts of the
paperlike capsules. Combretum indicum Indo-Pacific oceans, this tree
f: Combretaceae has horizontal branches.
This climber from tropical Asia has clusters Its corky fruit, dispersed by
of tubular red flowers. Its ellipsoidal fruits, water, contains an edible,
each with five wings, taste of almonds. almond-tasting nut.

flower has five

or six petals
ROSE GRAPE
Medinilla magnifica
f: Melastomataceae
This ornamental from the
Philippines is an epiphyte,
growing on trees. Its
10 ft leaves have the longitudinal
3m veins characteristic of
the melastomes.
16 ft
5m
BRAZILIAN SPIDER
FLOWER
Tibouchina urvilleana VIRGINIA MEADOW BEAUTY
f: Melastomataceae 23½ in Rhexia virginica
In warm regions this Brazilian 60 cm f: Melastomataceae
ornamental blooms for most of This hairy, perennial herb from E. US
the year. Its velvety leaves have and Canada grows on wetlands. It has
red edges and three to five square stems and stalkless leaves
obvious longitudinal veins. with toothed margins.
 M Y RTA L E S TASMANIAN
SNOW GUM
120 ft
36 m
160 woody seed Eucalyptus coccifera
capsule f: Myrtaceae CIDER GUM
With gray-white bark, peeling Eucalyptus gunnii
in long strips to reveal a f: Myrtaceae
stamens form
FLOWERING PLANTS • EUDICOTS

creamy-white trunk, this This hardy Tasmanian tree

bottlebrush-shaped species of Tasmanian uplands
flowers forms juvenile leaves that
has unstalked, opposite leaves. 82 ft are round and silvery,
25 m maturing to scythe-shaped
and blue-gray. It is used in
10 ft 130 ft
3m 40 m paper production.

TONGHI RED GUM

BOTTLEBRUSH Eucalyptus camaldulensis
Melaleuca subulata f: Myrtaceae
f: Myrtaceae Widespread in Australia,
A spreading shrub with small, with smooth, pale bark and elliptical
woody fruits containing blue-green leaves, this leaf smells of
hundreds of seeds. It is found riverside tree has durable peppermint
mainly in New South Wales lumber and its nectar
and Victoria, Australia. makes good honey.
10 ft
3m
40 ft
12 m
82 ft 16 ft
GREEN 25 m 5m
BOTTLEBRUSH
Melaleuca virens 65 ft
f: Myrtaceae 20 m
Resisting snow, frost,
and drought, this
sprawling subalpine
shrub from Tasmania
has leaves with sharp,
pointed ends. It attracts
birds and butterflies.

CAJUPUT COMMON MYRTLE ALLSPICE URN-FRUITED

Melaleuca cajuputi Myrtus communis Pimenta dioica GUM
f: Myrtaceae f: Myrtaceae f: Myrtaceae Eucalyptus urnigera
Native from S.E. Asia to N. Australia, Native from the Mediterranean From the Caribbean, S. Mexico, and f: Myrtaceae
this tree has aromatic leaves that to Pakistan, myrtle produces C. America, this unisexual tree has This tree from S.E. Tasmania has
contain a pale yellow medicinal oil. fragrant flowers and blue-black small, white flowers that form brown, urn-shaped fruits, blue-gray juvenile
berries. Its aromatic leaves berrylike stoned fruits. Unripe fruits leaves, and white flowers in clusters
yield essential oils. are dried and ground to make “allspice.” of three with numerous stamens.

50 ft 10 ft
15 m 3m evergreen
leaves
SCARLET KUNZEA
Kunzea baxteri
f: Myrtaceae
Kunzea species are
unusual in having
colorful stamens much
longer than their petals,
which adds to the showy 40 ft
12 m
appearance that attracts
pollinating birds.

65 ft CHILEAN MYRTLE
20 m Luma apiculata
f: Myrtaceae edible fruit
This slow-growing tree has a contorted
trunk and smooth gray-orange peeling
bark. Its fruit is a black berry.
65 ft GUAVA
20 m Psidium guajava
f: Myrtaceae
POHUTAKAWA CLOVE This tree from tropical and
Metrosideros excelsa Syzygium aromaticum subtropical America has flaky,
f: Myrtaceae f: Myrtaceae coppery bark. The fruits have
Native to North Island, New Zealand, this tree The dried flower buds of this native of Indonesian islands a sweet, musky smell when
produces red flowers in December. Old trees are used as a spice. Its cream-colored flowers have red ripe, and contain numerous
have beardlike masses of hanging aerial roots. stamens and ripen to purple, single-seeded berries. hard, yellow seeds.
 large, purple-
pink flower

FLOWERING PLANTS • EUDICOTS

3¼ ft 10 ft
1m 3m

5 ft
1.5 m

10 ft
3m ROSEBAY
WILLOWHERB
GODETIA FUCHSIA FULGENS HARDY FUCHSIA Epilobium angustifolium
Clarkia amoena f: Onagraceae Fuchsia magellanica f: Onagraceae
f: Onagraceae Typically found clinging f: Onagraceae Widespread in the northern
This annual is native to coastal to rocks or tree branches This widely planted hemisphere, this herbaceous
hills of western N.America. by its long, tuberous deciduous, ornamental perennial proliferates by
Cultivated in gardens, its roots, this deciduous perennial originates from spreading its roots and
flower has four broad petals shrub grows near water Chile and Argentina, where wind-dispersed seeds. Some
and forms a dry capsule, in the mountains it usually grows with its taxonomists put it in the
containing numerous seeds. of Mexico. roots in water. genus Chamerion or
Chamaenerion.

20 in 5 ft
50 cm 1.5 m

SHOWY EVENING EVENING PRIMROSE

PRIMROSE Oenothera biennis stigma has
Oenothera speciosa f: Onagraceae four lobes
f: Onagraceae This biennial from eastern 6½ ft
2m
This herbaceous, smooth- N. America has bluish green
stemmed perennial is from leaves in a basal rosette and
S.E. US and Mexico. Its white erect stems bearing flowers that
flowers become pink with age open in the evening. The seed
and close in full sun. oil has medicinal properties.
GREAT
WILLOWHERB
28 in ENCHANTER’S NIGHTSHADE characteristic Epilobium hirsutum
70 cm Circaea lutetiana four petals f: Onagraceae
f: Onagraceae This very hairy perennial
This woodland plant is found in Europe, forms extensive patches from
N. Africa, and the Middle East. Flowers of its underground stems. It is
two sepals and two notched petals form native to most of Europe,
small, round, hairy fruits. N. Africa, and parts of Asia.

SEEDBOX
Ludwigia alternifolia
f: Onagraceae
Native to eastern N. America, this is a
plant of damp places. It has box-shaped 3¼ ft
seed pods borne on angled stems. 1m

CELASTRALES
The two families in this order are
20 ft
characterized by a nectar-secreting disk SPINDLE TREE 6m
in the flower. The Lepidobotryaceae Euonymus europaeus GRASS OF 12 in
f: Celastraceae PARNASSUS 30 cm
comprises just two tropical tree species, Native to Europe, the wood Parnassia palustris
but the Celastraceae or spindle tree of this tree was used to f: Celastraceae
family has 1,200 species, mainly of vines, make spindles—spikes used This rosette plant grows
to spin thread. Its fruits nectar in bogs in the northern
shrubs, and trees. Genetic analysis is yet ripen to red and split into disk temperate zone. Each
to determine how the Celastrales order four to reveal orange- elliptical leaf flower is borne on a stem,
relates to other rosids. coated, poisonous seeds. clasped by a stalkless leaf.
 C UC U R BI TA L E S WHITE BRYONY
Bryonia cretica ssp. dioica
162 Eight mainly tropical families of trees, shrubs, 13 ft
f: Cucurbitaceae
4m Common in hedgerows and scrub on
herbs, and climbers make up this order. Six calcium-rich soils in S. Europe and
have few members, but the Begoniaceae or N. Africa, this poisonous climber has
FLOWERING PLANTS • EUDICOTS

begonia family has 1,400 species, 130 of which a swollen, tuberous root.
are horticultural plants. The 850 species in the
Cucurbitaceae or gourd family include notable large, spherical fruit stem
food plants such as squash and pumpkins. Both fruit narrows
of these families have male and female flowers above middle
on the same plant (monoecious).
6½ ft
2m
large leaf edible CUCUMBER hairy
flower leaves
Cucumis sativus
f: Cucurbitaceae
Originally from tropical
Asia, cucumber has been
cultivated for 3,000 years immature
and there are many fruit
varieties, including the
nutritious gherkin. The plant has
fruit tubular yellow flowers.
15 ft/4.5 m

3¼ ft
1m

fruit leaf sprig

SQUASH SMOOTH LOOFAH
Cucurbita pepo Luffa cylindrica
f: Cucurbitaceae f: Cucurbitaceae CALABASH 16 ft
With five-sided prickly stems and This climbing, annual Lagenaria siceraria 5m
large, yellow-orange flowers, vine, originally from Asia, f: Cucurbitaceae
pumpkins, marrows, and zucchini produces cylindrical fruits. This vine produces whitish, crinkly flowers.
are all varieties of Cucurbita pepo Their fibrous interior can Its edible, hard-shelled fruit floats in the sea
from C. America. be eaten young, or dried for months and is used as a container.
and used as a sponge.

3¼ ft/1 m
SQUIRTING
CUCUMBER
3¼ ft Ecballium elaterium
1m f: Cucurbitaceae
The ripe fruit of this
Mediterranean plant
becomes so swollen with
juices that it explodes when
16 in/40 cm
touched, squirting its seeds
up to 20 ft (6 m). WATERMELON BEGONIA LISTADA
Citrullus lanatus f: Begoniaceae
f: Cucurbitaceae Native to Paraguay, this sprawling
With yellow-green flowers, this annual, prostrate species has thick, velvety leaves with a
vine originated in southern Africa and its fruits red underside and white-pinkish flowers.
are cultivated throughout warmer climates. It was first described in 1981.

BALSAM PEAR
poisonous, Momordica charantia
exploding fruit f: Cucurbitaceae
Also called bitter gourd,
this tropical vine produces
bitter-tasting fruits. Their
tips split into three to reveal
yellow pulp and seeds encased
in scarlet coats (arils).
10 ft 150 ft
3m 45 m

TETRAMELES NUDIFLORA
f: Datiscaceae
Props at the trunk base support
this tall tree from Asia and
leaf is up to N. Australia. Male flowers grow on
4 in (10 cm) long different trees than the female flowers,
which form round seed capsules.
FA BA L E S
Found all over the world, except for Antarctica, the
163
Fabales have compound leaves with tiny outgrowths
(stipules) at the base and seed pods that open when

FLOWERING PLANTS • EUDICOTS

mature. Called legumes, these plants have nodules—
swellings that contain bacteria—on their roots that
help them take nitrogen from the air into soil. The
Fabaceae, the largest of the four families, includes
plants like the pea with flowers that have a large 20 in
upper petal with adjacent smaller petals. 50 cm 20 in/50 cm SENSITIVE PLANT
pinnate leaf Mimosa pudica
PEANUT f: Fabaceae
Arachis hypogaea This prickly S. American plant’s leaves close when
f: Fabaceae touched. Its flower heads (⅜ in/1 cm) usually
Native to Brazil, this legume has have pink flowers and long, pink-purple stamens.
yellow, reddish-veined, pealike seed pod herbaceous plant
flowers. Its seed pod matures in the
ground, producing peanuts.
GOLDEN SHOWER flower
TREE raceme
Cassia fistula
f: Fabaceae 65 ft
This slender tree from 20 m
32 in S.E. Asia has pendulous,
80 cm pealike flowers and pinnate deciduous
leaves with three to eight leaflet
pairs of leaflets. Its seeds
are poisonous.

glossy
leaves

pod has
65 ft edible pulp
20 m

SILVER WATTLE SAINFOIN

Acacia dealbata Onobrychis viciifolia
f: Fabaceae f: Fabaceae
This tree has fragrant flowers This fodder crop from 130 ft
and silvery bark, aging to S. Europe has green, oval, 40 m
black. Native to S.E. Australia, pinnate leaves with 6–14 pairs
silver wattle is often called of leaflets. Densely packed pink
mimosa elsewhere. flowers appear on a main stem. WEST INDIAN
LOCUST TREE
Hymenaea courbaril
f: Fabaceae
This hardwood tree has a
straight, thick trunk and stems of
leaf white-purplish flowers with large
sprig petals and long stamens. Orange
gum oozes from its trunk, which
4 ft is used in incense and perfumes.
1.2 m
flowers are clusters
of stamens with
23½ in reduced petals
60 cm 65 ft
20 m
KIDNEY VETCH TAMARIND
Anthyllis vulneraria Tamarindus indica
f: Fabaceae f: Fabaceae
A plant of dry, grassy places This evergreen tree with seed
pods
throughout Europe, this drooping branches from East
perennial has silky-haired Africa and Asia has yellow-
stems and clusters of yellow orange flowers on stems and
to red flowers, with a ruff edible pulp inside the seed pod.
of downy bracts beneath.

FLAT WATTLE
Acacia glaucoptera 40 ft/12 m SILK TREE
f: Fabaceae Albizia julibrissin
The globular flowers of f: Fabaceae
this spreading shrub leaf with This deciduous tree is native
from S.W. Australia 6–12 pairs of to S.W. Asia. It has dark
grow from a modified pinnate leaflets green bark, striped vertically
stem looking like a with age, and is fast-growing
twisted leaf. but short lived.
 FA BA L E S evergreen JUDAS TREE
Cercis siliquastrum
leaves
164 f: Fabaceae
This deciduous, ornamental and lumber
tree from the E. Mediterranean flowers
35 in profusely in the spring, then later
90 cm
develops flat 4 in (10 cm) seed pods.

5 ft glossy
1.5 m pinnate 33 ft
leaf 10 m
WILD LICORICE
Astragalus glycyphyllos
f: Fabaceae
With similar leaves to true
licorice, this European
grassland herb has curved
pods. It can be used pulpy
50 ft pods
as a tea. 15 m seeds
hard
seed
BLUE FALSE INDIGO CAROB TREE
Baptisia australis Ceratonia siliqua
f: Fabaceae f: Fabaceae
The sap of this herbaceous 130 ft/40 m This Mediterranean tree
perennial, native to woods and has a thick trunk, dense
streamsides in E. US, turns BLACK BEAN TREE foliage, and small,
purple when exposed to air, and is Castanospermum australe dull-green flowers. Pulp
used as an indigo dye substitute. f: Fabaceae from the seed pods is a
The orange-and-red flowers of this chocolate substitute.
Australian lumber tree develop into
woody pods containing three to young seed pod
20 in
five beanlike seeds. 50 cm
5 ft/1.5 m CHICK PEA
Cicer arietinum
f: Fabaceae
5 ft
1.5 m One of the earliest cultivated
vegetables from the Middle
East, its seed pod contains one
to three seeds, which can be
cooked to make hummus.
GOAT’S-RUE
Galega officinalis 10 ft
f: Fabaceae 3m
Naturalized in temperate
areas, this perennial, with
long, cylindrical, red-brown
seed pods, is believed by some
to improve lactation and
reduce fevers and diabetes.
MOUNT ETNA BROOM
Genista aetnensis
f: Fabaceae
Native to Sicily, around the slopes of
purplish- Mount Etna, and Sardinia, this small
pink tree has few leaves; flattened green
flower shoots assist photosynthesis.

DORYCNIUM
Lotus hirsutus
f: Fabaceae
10 ft/3 m This Mediterranean perennial 20 in
forms gray-green shrubby 50 cm
mounds. Its red-brown
cylindrical pods can be
SMALL-FLOWERED GORSE mistaken for small berries.
Ulex parviflorus
f: Fabaceae pod
contains
This dense, spiny, perennial shrub 10–15 seeds
from the W. Mediterranean has
yellow leaves and short, brown-black
pods. Fire and brush clearance aid COMMON BIRD’S-
seed germination. FOOT TREFOIL
Lotus corniculatus 5 ft/1.5 m
f: Fabaceae
BROAD-LEAVED EVERLASTING PEA Found in grassland in COMMON VETCH
Lathyrus latifolius Europe, Asia, and Africa, Vicia sativa
f: Fabaceae it has yellow flowers and 12 in
f: Fabaceae
Distributed through S. Europe and N. Africa, five lobed leaves, three 30 cm This scrambling, widespread annual is
this vigorous, climbing perennial has of which sit above the native to Europe and the Mediterranean. An
winged stems and clusters of 5–15 pinkish others. Its seed pods animal-feed crop, its flowers are normally
flowers. Its seeds form in long pods. resemble a bird’s foot. paired and its tendrils can be branched.
 ALFALFA
Medicago sativa
pealike
flowers f: Fabaceae 165
This deeply rooting perennial
grows on chalk grassland
throughout S.W. Asia, Europe, and

FLOWERING PLANTS • EUDICOTS

the US. It is a useful animal-feed
crop and has medicinal properties.
12 in
30 cm 32 in 5 ft
twisted seed 80 cm 1.5 m
31/4 ft pods
1m HORSESHOE VETCH densely packed
Hippocrepis comosa flowers
f: Fabaceae
CROWN VETCH This prostrate perennial, from S. Europe
Securigera varia and the Mediterranean, is a major food
f: Fabaceae plant for blue butterfly caterpillars.
This rapidly spreading Its seed pods are twisted into
perennial of S. Europe and horseshoelike segments.
W. Asia has thick leaves and
deep roots, good for binding 31/4 ft 31/4 ft
soil and controlling erosion. 1m 1m

6½ ft
2m

BROOM RED CLOVER

Cytisus scoparius Trifolium pratense
f: Fabaceae f: Fabaceae
Found in heaths across Europe, A food crop for grazing
this shrub has strongly scented animals, this perennial
flowers and many slender, ridged, has long-stalked basal
and angled branches traditionally leaves, and an oblong
used for brooms. fruit pod hidden within
compound its flowerhead.
leaves blue-violet
flowers
strong stem

root LICORICE
Glycyrrhiza glabra
f: Fabaceae
This feathery perennial has smooth, small,
oblong pods and deep roots. Extract
from the roots is 50 times sweeter than
sugar and is medicinal.

12 ft
narrow leaflet
3.7 m
COMMON
SCARLET LABURNUM
RUNNER BEAN Laburnum anagyroides
Phaseolus coccineus f: Fabaceae
f: Fabaceae This deciduous tree is
Native to C. American native to C. and
mountains, the shoot of this S. Europe. Clusters
perennial twists clockwise of brown hairy pods
around its support. Its seeds (3 in/7.5 cm) contain
(beans) are variously colored. toxic black seeds.

40 ft
12 m

82 ft/25 m

GARDEN LUPIN
16 in Lupinus polyphyllus
40 cm f: Fabaceae
Originating from
NICEAN MILKWORT western N. America
BLACK LOCUST Polygala nicaeensis and naturalized
Robinia pseudoacacia f: Polygalaceae throughout Europe,
f: Fabaceae This perennial from France this popular
This vigorous deciduous tree, and Italy has flowers with two ornamental has hairy,
originally from S.E. USA, spreads sepals and three joined petals, black pods and
by suckers. It has poisonous bark one of which is fringed, and a undersides of leaves. Its
and leaves, and flat brown seed pods. small capsule as a fruit. flowers are fragrant.
 FAGA L E S 80 ft 80 ft
25 m
EUROPEAN HORNBEAM
Carpinus betulus
166 25 m
f: Betulaceae
The seven families in this order include Frequenting hedgerows, this
some of the world’s best-known trees, which European native has small, green,
dominate the woodlands in which they grow: male and female catkin flowers that
FLOWERING PLANTS • EUDICOTS

beech trees in the Fagaceae, birches in the develop into nuts surrounded by
three-lobed bracts.
Betulaceae, southern beeches in the Nothofagaceae,
walnut trees in the Juglandaceae, and Australian
she-oaks in the Casuarinaceae. Most have simple bract
JAPANESE HOP
leaves and small, wind-pollinated flowers. HORNBEAM male catkin
Ostrya japonica
65 ft fruit and f: Betulaceae
20 m seed develop This species from the Far East
from catkins has gray-brown scaly bark. Its
nutlet seeds are enclosed in
FOREST OAK husks, like hops, and hang
Allocasuarina torulosa in long clusters.
f: Casuarinaceae
The lumber of this W. Australian
tree is prized by wood-turners.
Its pendulous branches carry
needlelike, green twigs with tiny 100 ft
leaves and warty cones. 30 m

female 43 ft
flower 13 m

EUROPEAN HAZELNUT
Corylus avellana
f: Betulaceae RED
This European shrubby tree ALDER
is harvested for its edible nuts. Alnus rubra leaf bud
Its male catkins and female f: Betulaceae
flowers appear in early spring. A native of western N. America, this tree
prefers damp slopes and margins of
watercourses. It has pale gray bark that
becomes red when bruised or scraped.
catkin is a
cylindrical cluster
of male flowers

80 ft
25 m
pinnate
leaves

RAULI PLATYCARYA
Nothofagus alpina STROBILACEA
f: Nothofagaceae f: Juglandaceae
A lumber tree, rauli is native to Native to E. Asia, this
Argentina and Chile. The young tree has clusters of erect 50 ft
leaves are bronze. Greenish female male catkins and a fruit, 15 m
flowers grow in clusters and form like a conifer cone, with
bristly husks enclosing small nuts. ripe 130 ft winged seeds.
pecan 40 m
nut clusters form PECAN BUTTERNUT
in leaf axil Carya illinoinensis Juglans cinerea ERMAN’S BIRCH
f: Juglandaceae f: Juglandaceae Betula ermanii
6½ ft The pecan is native to This deciduous tree, native
f: Betulaceae
100 ft
2m N. America and cultivated for to N. America, has 30 m Native to mountain areas from
its edible nuts. The nuts are yellow-green catkins. Small Siberia to Japan, this tree is widely
enclosed in a husk that splits bunches of egg-shaped nuts planted in parks and gardens. It
four ways when ripe. contain sweet seeds. has peeling, whitish bark.

100 ft
30 m 100 ft
SWEET GALE 30 m ripe
Myrica gale walnut
f: Myricaceae ENGLISH WALNUT
This sweet-smelling shrub, Juglans regia
traditionally used as an insect f: Juglandaceae
repellent, grows in peat bogs A species from the mountains of S.W. Asia, the
in northern temperate zones. walnut tree is valued for its fruits and lumber.
It has either male or female It has smooth gray bark and male catkins up leaf sprig with
reddish catkins. to 6 in (15 cm) long. unripe fruits
 130 ft ENGLISH OAK SCARLET OAK 80 ft
Quercus robur Quercus coccinea
40 m
f: Fagaceae f: Fagaceae
25 m
167
Long-lived, with valuable lumber, The leaves of this N. American tree
deciduous scalycup
acorn this tree is common in W. Europe, turn a characteristic deep red in the
leaf especially Britain. It has male fall. This oak bears long catkins
flowers in long, hanging catkins, of yellowish-green male flowers and its
and long-stalked acorns. acorns have a glossy cup.

catkin fall spring color

sprig leaf

toothed leaf
30 ft
10 m
100 ft
30 m

spiny burr
KERMES OAK encases nuts
100 ft Quercus coccifera
30 m f: Fagaceae
An evergreen, shrubby tree from
the Mediterranean area, this
50 ft species has hollylike leaves that
15 m are bronze when young, and
yellow-brown male catkins.

JAPANESE STONE OAK AMERICAN BEECH EUROPEAN CHESTNUT

Lithocarpus edulis Fagus grandifolia Castanea sativa
f: Fagaceae f: Fagaceae f: Fagaceae
This Japanese evergreen tree has This broadly spreading tree from Cultivated for 3,000 years for its nuts,
upright creamy-white catkins, which eastern N. America has gray-brown this tree is from S.E. Europe and
have female flowers at the base and bark. It has glossy deciduous W. Asia. It bears catkins with male
males above. The edible acorns ripen leaves and two fruits (beechnuts) flowers in the upper part and
over two years. in each husk. female flowers beneath.

MALPIGHIALES 32 in
One of the largest and most diverse orders, 80 cm
Malpighiales comprises 36 mainly tropical
families with more than 16,000 species,
grouped together by their DNA but widely
different in form. Half the families, including one
only recognized in 2016, have fewer than 100 poisonous
fruits
members and many are unfamiliar outside their
native regions, but they also include the huge
spurge family (Euphorbiaceae), with 6,300 species. TUTSAN
Hypericum androsaemum
f: Hypericaceae
This small shrub from W. Europe has
coarsely toothed, reddish, two-ridged stems. Its
ovate leaf aromatic leaves have medicinal uses,
but its berries are poisonous.
100 ft
30 m
paired
leaves

32 in
80 cm

PERFORATE ST
JOHN’S-WORT
Hypericum perforatum
f: Hypericaceae
This Eurasian perennial is a
CEYLON IRONWOOD common plant of banks,
Mesua ferrea fields, and roadsides. It has
f: Calophyllaceae round stems with ridges on
This Asian tree yields heavy lumber, hence either side, leaves with
its common name. It has large flowers with four translucent dots, and
white petals and bright red young leaves. many-seeded capsules.
 MALPIGHIALES 6 ft
1.8 m
168
trilobed
leaf 8 in
50 ft 16 ft 20 cm
FLOWERING PLANTS • EUDICOTS

15 m 5m

4 ft
nut containing 1.2 m
hard seed

CANDLE-NUT TREE CASSAVA

Aleurites moluccanus Manihot esculenta CROWN OF THORNS BASEBALL PLANT
f: Euphorbiaceae f: Euphorbiaceae Euphorbia milii Euphorbia obesa
Oil from the nuts of this The tuberous roots of this f: Euphorbiaceae f: Euphorbiaceae
tropical tree was burned in S. American plant are boiled and This semi-succulent, climbing Rare in the wild, this ball-shaped
lamps. It has stems of small, ground as food. It has small shrub from Madagascar has succulent originates from the Great
creamy flowers and its variable flower clusters that grow on very spiny stems. Its leaves are Karoo, South Africa. Its tiny flowers
leaves are gray-green secondary branches. mainly on the new shoots. grow from “eyes” at its top.
when young.
20 ft
13 ft 6m
palmate 4m
leaf CROTON
Croton tiglium
f: Euphorbiaceae
Used in Chinese medicine,
this tree from S.E. Asia has
malodorous leaves and produces
sticky oil-yielding fruit capsules containing three
flower head leaves seeds
poisonous seeds.
CASTOR OIL PLANT JATROPHA LARGE MEDITERRANEAN
Ricinus communis GOSSYPIIFOLIA SPURGE
f: Euphorbiaceae f: Euphorbiaceae Euphorbia characias
Castor oil is extracted from the This invasive, toxic plant, f: Euphorbiaceae
This ornamental, Mediterranean 12 in
poisionous seeds of this species banned in parts of Australia, is 30 cm
from tropical Africa. With native to tropical America. It perennial has upright, purplish,
upright flower heads, the upper has sticky leaves, watery sap, woolly stems, bare and smooth at the
female flowers have red stigmas and flowers with purple bracts. base. The fruit, or seed capsule, is
and lower males yellow anthers. 13 ft hairy and berrylike.
4m

130 ft
40 m

16 in
40 cm
PETTY SPURGE SAUSAGE SPURGE
Euphorbia peplus Euphorbia guentheri
RUBBER TREE f: Euphorbiaceae f: Euphorbiaceae
Hevea brasiliensis This poisonous, weedy annual, This evergreen succulent from
f: Euphorbiaceae native to Europe, N. Africa, and Kenya has white flower bracts
16 in W. Asia, has dividing stems and with purple markings. It has
A native of Brazil, this tree is famous 40 cm
for the milky sap (latex) under its flower stalks with three branches. fleshy, sickle-shaped leaves
bark used for rubber. It has leaves mainly on young growth.
with three leaflets and pungent DOG’S MERCURY
yellow flowers. Mercurialis perennis
f: Euphorbiaceae five-lobed
This downy perennial has a leaf and
single upright stem. Plants are flower
either male or female and
produce tiny green flowers
on long, thin stalks.

berry with
numerous seeds

leaves have twining

tendril at base
23¹⁄2 in 10 ft 65 ft
PERENNIAL FLAX 3m 20 m
60 cm Linum perenne BLUE PASSION FLOWER
f: Linaceae COCA Passiflora caerulea
This slender, often Erythroxylum coca f: Passifloraceae
spreading perennial is f: Erythroxylaceae This ornamental S. American vine has
native from C. Europe to The leaves of this evergreen shrub scented flowers associated with
China. Its flowers form from northwestern S. America yield Christian symbolism. Flowers have ten
buds at intervals at the cocaine. It has oval leaves and clusters very similar sepals and petals, five
top of the stem. of small, yellowish white flowers. stamens, and three purple stigmas.
 23¹⁄2 in
60 cm

RAFFLESIA
33 ft 80 ft Rafflesia arnoldii
10 m 25 m f: Rafflesiaceae
This leafless parasite on S.E Asian 80 ft
AYAHUASCA RED MANGROVE 25 m
Banisteriopsis caapi Rhizophora mangle rainforest vines has the world’s largest
f: Malpighiaceae f: Rhizophoraceae flowers, up to 3¼ ft (1 m) in diameter.
This woody vine, native to the Found throughout the tropics, Their foul smell attracts flies.
Amazon, is traditionally used to especially in swampy salt marshes,
make a sacred, medicinal drink. the red mangrove has prop roots 40 ft
It produces stalks of pink and its seeds germinate before 12 m
flowers and winged seed pods. leaving their parent tree.
PUSSY WILLOW
Salix caprea
70 ft f: Salicaceae
21 m This shrubby tree, native to Europe and
Asia, has downy, oval, toothed, alternate
leaves. Female catkins form capsules,
releasing cottony seeds.

IDESIA
Idesia polycarpa
f: Salicaceae WHITE WILLOW
A mountain tree from E. Asia, this has 80 ft
Salix alba
flower is up to f: Salicaceae
5 in (12 cm) across smooth, gray bark. Small yellow-green, 25 m
GIANT fragrant flowers produce deep red berries. This waterside tree is
GRANADILLA native to Europe and
Passiflora quadrangularis Asia. Trees have either
f: Passifloraceae male or female catkins,
This passion flower is never both. The bark
ASPEN is a source of salicin,
a perennial native of Populus tremula
S. America. It bears f: Salicaceae the active ingredient
oblong fruits on of aspirin.
Native to Europe and Asia,
four-sided stems. the gray bark of young trees
has diamond-shaped scars.
fragrant flower, may be Flattened stalks cause the 100 ft
white, red, or purple leaves of aspen to quiver. 30 m

WHITE POPLAR
Populus alba
f: Salicaceae
This deciduous tree, native to
C. Europe and C. Asia, tolerates salt
water. It is unisexual, and most trees
are female. Catkins form capsules that
release fluffy seeds.

50 ft
15 m
33 ft
10 m
AZARA
MICROPHYLLA
:
f Salicaceae
This evergreen tree
from Argentina and Chile
has small, vanilla-scented
flowers with yellow
stamens. Each leaf has a
circular growth (stipule)
at the base.

12 in/30 cm 4 ft/1.2 m

WILD PANSY SHRUB VIOLET

Viola tricolor Pigea floribundus
f: Violaceae f: Violaceae
This sprawling, short-lived This Australian woody
perennial is native to Europe perennial accumulates
and W. Asia, growing in grassy nickel. Shrub violet has
habitats with neutral to acid bluish petals with a yellow
soils. Used in herbal remedies, patch and small, dark
it is also known as heart’s-ease. green, lance-shaped leaves.
 OX A L I DA L E S ALBANY PITCHER
PLANT ROSA L E S
170 This order contains about 2,000 species,
Cephalotus follicularis
This order of plants contains nine families, among
f: Cephalotaceae
grouped within seven families. Of these, This carnivorous plant is native them the Rosaceae (rose family), Cannabaceae
the Cephalotaceae contains only one to the coast of S.W. Australia. (hemp family), Moraceae (mulberry family),
FLOWERING PLANTS • EUDICOTS

species—the carnivorous Albany pitcher It has basal, oval leaves, and a Rhamnaceae (buckthorn family), Ulmaceae (elm
liquid-filled pitcher that traps
plant. The Cunoniaceae are woody prey but it is not a member of family), and Urticaceae (nettle family). Members
plants that produce woody fruit capsules the pitcher family (see p.194). of the Rosales are often grown for
containing small seeds. The wood sorrel their fruits or other products.
family (Oxalidaceae) is the largest, with 8 in Plants in this order tend to
20 cm
800 species within five genera. They have five sepals and numerous
are characterized by divided leaves that stamens. Mostly insect-
exhibit or open out by day and then pollinated, they are
close up at night. often thorny
or hairy.

flower has
five petals
13 ft
4m

CHRISTMAS BUSH
Ceratopetalum gummiferum
f: Cunoniaceae
From coastal E. Australia,
this shrub produces
insignificant white flowers in
14 in spring. The pink and red
35 cm sepals enlarge in winter,
enclosing the fruit.

PINK SORREL
Oxalis articulata 40 ft
f: Oxalidaceae 12 m
Growing from a swollen
rhizome, or root stalk,
this plant from S. America
forms leafy mounds flower
topped with clusters of cluster
flowers. These produce
seed capsules that explode.
BLACK
WATTLE
Callicoma
serratifolia
f: Cunoniaceae
The wood of this shrubby tree from the
east coast of Australia was used by early
settlers to make wattle-and-daub shelters.
Its young foliage is bronze.

oil-rich berry

SEA BUCKTHORN
ripe flower Hippophae rhamnoides
fruit sprig f: Elaeagnaceae
leaflet STAR FRUIT Widespread through Asia
cluster Averrhoa carambola and Europe, this shrub
f: Oxalidaceae bears tiny flowers before
This bushy tree, native to S.E. Asia, is the leaves. The small,
widely cultivated for its edible star-shaped 33 ft bright-orange berries are
fruits. It can flower four times a year. 50 ft/15 m 10 m rich in vitamin C.
 20 ft/6 m 6½ ft 23 ft/7 m
2m
171

female plant
and flowers

male plant
and flowers
OLEASTER HEMP HOP
Elaeagnus angustifolia Cannabis sativa Humulus lupulus
f: Elaeagnaceae f: Cannabaceae f: Cannabaceae
Native to Eurasia, this spreading, Originating from C. and W. Asia, this This climbing perennial from N. Europe and
deciduous tree has spiny shoots, annual is a source of cannabis from its N. Asia is widely naturalized elsewhere.
covered in silver scales. Its egg-shaped, leaves, fiber for making rope, and The conelike female flower or hop is used
yellow-red fruits are edible. oil is extracted from its seeds. to flavor and preserve beer.

65 ft BLACK MULBERRY
20 m young fruit Morus nigra
43 ft f: Moraceae
13 m This deciduous, spreading tree,
widely cultivated for its rich-
flavored fruits, is native to the
Middle East. It has bumpy,
fissured orange bark.

JACKFRUIT
Artocarpus heterophyllus
f: Moraceae
From lowland S.E. Asia, this tree
yields the largest tree-borne fruit,
weighing up to 110 lb (50 kg) and
36 in (90 cm) long when ripe.

33 ft
10 m 50 ft/15 m

100 ft conical sprig of female

glossy fleshy 30 m fruits flowers
leaf fruit
COMMON FIG SACRED FIG PAPER MULBERRY
Ficus carica Ficus religiosa Broussonetia papyrifera
f: Moraceae f: Moraceae f: Moraceae
As with most figs, flowers of this Under this sacred tree, Buddha is said Fine paper is made from the inner
Mediterranean and Asian species develop to have gained enlightenment. Native to bark of this tree from Japan and S.E.
inside a flask-shaped receptacle and are S.E. Asia, its flowers, then its fruits, are China. Male catkins produce large
pollinated by a single species of wasp. enclosed within purple, flecked figs. quantities of wind-dispersed pollen.

65 ft
20 m

26 ft/8 m

30 in
leaf sprig 75 cm
33 ft/10 m
OSAGE ORANGE JUJUBE COMMON BUCKTHORN NEW JERSEY TEA
Maclura pomifera inedible Ziziphus jujuba Rhamnus cathartica Ceanothus americanus
f: Moraceae fruit f: Rhamnaceae f: Rhamnaceae f: Rhamnaceae
Native to the S.E. US, this tree This thorny, shrubby tree is widely cultivated A shrubby, invasive tree that has This bush from eastern N. America has
is used for hedging and its roots for its fruits in China and Korea. The spine-tipped shoots, tiny yellow-green purple, three-lobed capsules, containing
and wood were valued by immature, smooth, oval, green stoned flowers, and black berries. It is native seeds. Its red roots and hairy leaves
native Americans. fruit tastes like apple. to Europe, Asia, and N.W. Africa. have been used to make tea.
 ROSA L E S 80 ft
172 25 m

40 ft/12 m
32 in
80 cm
FLOWERING PLANTS • EUDICOTS

leaves and
SILVERWEED edible fruits
Potentilla anserina spring
f: Rosaceae flowers
This silky-haired, SNOWY MESPILUS WILD CHERRY
creeping perennial grows Amelanchier lamarckii Prunus avium
in wasteland, pastures, f: Rosaceae f: Rosaceae
and dunes in Europe, Frothy clusters of star-shaped flowers The wild ancestor of orchard cherries
CHINESE PHOTINIA Asia, and N. America, in spring are followed by dark red grows in woods and hedges in Europe,
Photinia serratifolia and has been widely berries in summer in this showy tree Asia, and N. Africa and is naturalized
f: Rosaceae introduced elsewhere. from eastern N. America. in N. America.
A plant of Chinese forests, this tree
is commonly cultivated for
ornamental purposes. Its dense
26 ft/8 m wood is used in furniture making. 40 ft
12 in 12 m
30 cm ORCHARD
PLUM
Prunus domestica
summer edible f: Rosaceae
flower fruits These trees originated
35 ft/11 m as a cross between the
STRAWBERRY cherry plum of China
Fragaria vesca and the European
f: Rosaceae blackthorn, but
The wild strawberry is a perennial without the sharp
in European and N. American thorns of the latter.
woodlands. Tiny edible fruits form double flower
from the swollen flower receptacle. garden variety

PRAIRIE CRAB APPLE three-lobed

Malus ioensis leaf
f: Rosaceae
One of several crab apples native
to N. America, this species is cultivated
for its rather tart fruits.

32 in
6 1/2 ft/2 m 80 cm

JAPANESE ROSE
Rosa rugosa
f: Rosaceae
Tolerant of salt spray, this rose from
E. Asia is often planted as a hedge near
the sea. It has prickly stems and pink
flowers with wrinkled petals.

white or
pink flowers

10 ft 61⁄2 ft
3m 2m

fragrant, deep
pink flower
DOG ROSE SWEET BRIAR APOTHECARY’S ROSE
Rosa canina Rosa rubiginosa Rosa gallica var. officinalis
f: Rosaceae f: Rosaceae f: Rosaceae
The thorny, arching branches of this One of the most deeply colored wild A popular garden plant, this European
rose are familiar in hedgerows in roses, sweet briar enlivens hedgerows and scrub parent of many floribunda and hybrid tea
Europe, Asia, and N. Africa. This plant around Europe, Asia, and Africa. Glands on its roses has a long pedigree. “Attar of roses”
is also naturalized in N. America. leaves produce an apple scent when crushed. is a fragrant oil distilled from its petals.
 COTONEASTER
Cotoneaster horizontalis
20 in f: Rosaceae 10 ft/3 m 173
50 cm This creeping Chinese
shrub is commonly
grown in gardens for its
flowers and fruits, and
occasionally escapes. The white or pink
oaklike eight-petaled flowers
leaf flower semievergreen leaves
form flattened clumps.
MOUNTAIN AVENS
Dryas octopetala
f: Rosaceae 3¼ ft edible
The flowers of this Arctic and 1m
FIRETHORN fruits
mountain undershrub turn to follow Pyracantha rogersiana
the sun to warm their centers and f: Rosaceae 8 ft
attract pollinating insects. many small This thorny evergreen shrub from
flowers 2.5 m
E. China belongs to a genus often BLACKBERRY
planted for their attractive but Rubus fruticosus
80 ft inedible, orange, berrylike fruits. f: Rosaceae
25 m Familiar in Europe as scrambling hedgerow
bushes bearing edible fruits in the fall,
blackberries form a group of very closely
related “microspecies.”

40 ft
12 m 20 ft/6 m
WILD SERVICE TREE AMERICAN MOUNTAIN ASH
Sorbus torminalis Sorbus americana
f: Rosaceae f: Rosaceae
This rare tree grows in ancient woods This deciduous woodland tree is native
in Europe, Asia Minor, and N. Africa. to eastern N. America. Its orange berries five-petaled
Its name is derived from the brew, last into winter, providing food for white flower
cerevisia, made from its fruits. thrushes and jays. MEDLAR
Mespilus germanica
five-petalled
f: Rosaceae
white flower Originally from C. and S. Europe,
medlar produces a hard, yellowish-
brown fruit. This turns soft and edible
in the fall, then decays.

SALAD BURNET
Sanguisorba minor fruits with remains
f: Rosaceae of sepals
Native from lobed leaf
Europe to Iran, and
introduced in
N. America, this
perennial of
lime-rich grassland
has edible leaves,
hence its name.
231⁄2 in
60 cm
WILLOW-LEAVED toothed flower heads
PEAR leaflets
Pyrus salicifolia
f: Rosaceae red fruits with
Cultivated for its pendulous, single stem
silvery foliage, not its 40 ft showy
inedible fruits, this Middle 12 m five-petaled
Eastern tree is endangered flower
in the wild in Turkey.

231⁄2 in
60 cm

red fruits,
231⁄2 in or haws
60 cm
nodding COMMON HAWTHORN fragrant blossom
summer burrlike Crataegus monogyna
flowers fruit LADY’S MANTLE f: Rosaceae
WATER AVENS Alchemilla vulgaris Occurring in woods and hedges
Geum rivale f: Rosaceae in the wild from Europe and N.
f: Rosaceae This name encompasses several Africa to Afghanistan, this densely
This downy perennial grows in damp closely related grassland species branched small tree produces
places in Europe, Asia Minor, and N. from Europe, Asia, and eastern massed white blossom in spring,
America. Hooked hairs on the fruits N. America. Their leaves are said followed by deep red fruits. It 52 ft
latch onto animals for dispersal. to resemble a trailing gown. is also cultivated in gardens. 16 m
174 SWEETBRIAR
R osa r u biginosa
Sweetbriar, or eglantine rose, is one of the
FLOWERING PLANTS • EUDICOTS

most attractive of a range of wild roses that

brighten hedgerows and scrub across Europe
into Asia and Africa. They have five-petaled
flowers that are usually white or pale pink. To
nonbotanists, they seem rather similar, and are
often called “dog roses.” This derogatory name
is said to be because they are less showy and size Erect stems to 61⁄2 ft (2 m) tall
prickly, spreading
strongly scented than the garden roses bred by habitat Open scrub, hedgerows, parks,
stem
and roadsides
horticulturalists over centuries, although some distribution Europe to Asia,
would argue they have a more subtle beauty and introduced elsewhere
delicate scent than their fancy relatives. Many
are prickly shrubs with scrambling, arched
stems, for which “briar” is a descriptive term.
Sweetbriar has deep pink flowers and leaves
that emit the scent of apples when crushed. leaflet teeth fringed
It has been introduced to the Americas and with stalked glands
Australia, where it has become invasive.

DEVELOPING HIPS
The flower base swells to form a
succulent hip, technically a “false fruit” ROSEBUD
because it is not formed from the ovary. Sepals around the
This contains numerous, small, one- flower bud protect
seeded “pips” which are the true fruits. the developing petals
sepals remain as and fertile parts. The
a leafy tuft stubble of glandular
hairs may insulate the
bud and repel pests.

UNDERSIDE
OF LEAF
The underside of
the leaf is densely
hairy, mixed with
many brownish,
stalked glands.
These release sweet-
smelling terpene
chemicals that may
deter grazers.
STAMENS
Numerous stamens
cluster in whorls
around the centrer
of the sweetbriar
flower. They have
pale stalks and
bright yellow
anthers, which
produce copious
amounts of pollen. leaflet with
saw-toothed
edge

THORNS RIPE HIPS

Hooked thorns, Hips shed their sepal tuft before
interspersed with ripening. Ripe, red hips are rich in
strong bristles, vitamin C and are eaten by thrushes
discourage large animals and pigeons, which spread the seeds
from grazing the leaves. in their droppings.
 showy flower up to 5- or 7-lobed leaf,
1½ in (4 cm) in paler beneath
diameter 175

FLOWERING PLANTS • EUDICOTS

five overlapping
petals, slightly
notched at tip
flower
base

SIDE VIEW
Five sepals spread beneath the
petals. After pollination by bees
or butterflies, the flower base
swells to form a hip.

pollen-producing
stamen

THE SWEET SMELL OF SUCCESS

Scent plays different roles in plants, to attract
pollinators or repel pests. We may enjoy the smell
of sweetbriar leaves, but old herbals say leaf extracts cluster of sticky,
were “harsh on the throat” and would “purge the pollen-receiving
chest,” so the apple-scented terpenes in the leaves stigmas
may well be distasteful to grazing animals.
 ROSA L E S JAPANESE ZELKOVA
Zelkova serrata
176 f: Ulmaceae
In the US elms killed by Dutch
elm disease are sometimes
replaced by this valuable Asian
FLOWERING PLANTS • EUDICOTS

lumber tree. The Japanese grow 12 in

it in bonsai form. 30 cm
100 ft
30 m
red berry FRIENDSHIP PLANT
toothed leaf Pilea involucrata
130 ft
f: Urticaceae
40 m Several Pilea species make valuable
houseplants, including this species
from C. and S. America with
strongly veined leaves.

HACKBERRY EUROPEAN FIELD ELM STINGING NETTLE 6½ ft/2 m

Celtis occidentalis Ulmus minor Urtica dioica
f: Cannabaceae f: Ulmaceae f: Urticaceae
Native to N. America, hackberry This tree used to be a major Stinging hairs discourage animals
has bright green, elmlike leaves, feature in European from eating nettle leaves. This species
and red fruits eaten by many 120 ft landscapes, but was decimated grows in disturbed ground in Europe,
birds and mammals. 36 m by Dutch elm disease. Asia, N. Africa, and N. America.

BRASSICALES
WILD CABBAGE
The order Brassicales has 17 families. Many members have bitter or Brassica oleracea
fragrant oils in their leaves, stems, or swollen roots. Although these oils f: Brassicaceae
3¼ ft Humans have grown this
evolved to deter grazers, they make many species pleasantly edible for 1m W. European plant for
humans, and have culinary, perfumery, or herbal uses. The cabbage millennia. Cauliflower,
family (Brassicaceae), is the largest group, with 3,300 species. broccoli, and brussels
sprouts are cultivated forms
of the same species. stalkless blue-
green leaf

flower head
male
flower

30 ft
10 m

30 ft/10 m
PAPAYA
Carica papaya horseradish flower
f: Caricaceae tree leaf stem
This C. and S. American HORSERADISH TREE
treelike plant produces yellow Moringa oleifera
flowers. The female flowers f: Moringaceae
develop into large orange- This tropical Asian tree has corky gray
fleshed fruits bark and fernlike leaves. A condiment 231⁄2 in
of the same name. can be made from its crushed roots. 60 cm

5 ft
1.5 m

NASTURTIUM
Tropaeolum majus
f: Tropaeolaceae
A colorful annual from
C. and S. America, this is a
popular garden plant. The
10 ft/3 m flowers and leaves can be
eaten in salads. 20 in/50 cm

COMMON WALLFLOWER
CAPER MIGNONETTE Erysimum x cheiri
Capparis spinosa Reseda odorata f: Brassicaceae
f: Capparaceae f: Resedaceae Probably first developed
A perennial spiny bush, caper is Originally from N. Africa, common as a hybrid in Greece, this
native to the Mediterranean. Its mignonette is grown in gardens species has been cultivated
flower buds are used salted and across S. Europe. An oil from its throughout Europe since
pickled for culinary use. fragrant flowers is used in perfumery. medieval times.
 HONESTY
Lunaria annua
f: Brassicaceae 177
Native in the wild in S.W.
Europe, honesty is often grown
in gardens. Its seed pods are used 4 ft
in dried flower arrangements. 1.2 m

HORSERADISH
5 ft Armoracia rusticana
1.5 m f: Brassicaceae
The pungent taste of the
roots, which are used for
28 in 12 in horseradish sauce, evolved to
70 cm 30 cm deter herbivores from eating
honesty this Caucasian perennial.
seed pod

CORALROOT BITTERCRESS AUBRIETA

Cardamine bulbifera Aubrieta deltoidea
f: Brassicaceae f: Brassicaceae
A plant of C. European beechwoods, Originating from the Aegean
this creeping perennial occurs from region, aubrieta grows readily
the British Isles eastward to the on walls throughout the warmer
Caucasus and Asia Minor. parts of Europe. honesty
flower stem

16 in
flower 40 cm
stem

edible
leaf
3¼ ft 12 in
1m 30 cm
WINTERCRESS SWEET ALISON
Barbarea vulgaris Lobularia maritima
f: Brassicaceae f: Brassicaceae
Native throughout Europe, This Mediterranean annual
wintercress was once is widely cultivated for its
cultivated for winter salads, sweet-scented flowers.
and introduced for this “Alison” from alyssum in Greek
purpose to N. America, refers to its reputation for
Australia, and New Zealand. curing madness.
lobed leaf
at base of stem
scented purple,
red, or white
flower head flowers

SHEPHERD’S PURSE
four-petaled Capsella bursa-pastoris
flower f: Brassicaceae 32 in/80 cm
Purse-shaped pods rounded
that split to release massed leaf is gray
seeds, have helped this beneath
species spread around the
world from its native
231⁄2 in Eurasian range.
60 cm

231⁄2 in
60 cm

distinctive wild radish leaf

leaf flower stem stem
SEA KALE
Crambe maritima WILD RADISH WATERCRESS
f: Brassicaceae Raphanus raphanistrum Nasturtium officinale HOARY STOCK
This cabbagelike perennial f: Brassicaceae f: Brassicaceae Matthiola incana
grows on shingle beaches and Also naturalized in This wild Eurasian perennial grows 231⁄2 in f: Brassicaceae
seacliffs around Eurasia. Its N. America, this Eurasian near water and is also cultivated 60 cm Native to coastal rocks in
spherical fruits, which float native may be the ancestor of in tanks. Its young shoots and S.W. Europe, stock is much
for many days, are dispersed garden radishes but its root peppery leaves, rich in vitamin C, cultivated. “Ten Weeks Stock”
by the sea. is not round. are used in salads. is a short-lived garden variety.
 M A LVA L E S
178 The ten families in the Malvales include many 33 ft/10 m
shrubs and trees, found mainly in tropical and
warm temperate regions, but extending into
FLOWERING PLANTS • EUDICOTS

cooler parts. The main members are the rockrose

family (Cistaceae), mostly shrubs from the spiny fruit
3¼ ft/1 m
northern hemisphere; the more widespread mallow ANNATTO
family (Malvaceae) of herbs, shrubs, and massive Bixa orellana
trees; and the pantropical Dipterocarpaceae, which f: Bixaceae
includes some of the most important tropical The food dye annatto comes from the
lumber trees. spiny fruits of this pink-flowered shrub
or small tree, native to tropical America.
COMMON ROCKROSE
Helianthemum nummularium
f: Cistaceae
Found on sunny banks
throughout most of Europe,
this low shrub prefers lime-rich
soils. Different subspecies grow
in Europe’s mountains.
13 ft/4 m
20 in
50 cm HAIRY ROCKROSE CALIFORNIAN FLANNELBUSH
Cistus incanus Fremontodendron californicum
f: Cistaceae f: Malvaceae five-petaled
Several scientific names The flowers of this spreading shrub produce rose-red flower
have been given to this showy masses in early summer. It lives high
widespread Mediterranean up in the granite mountains of California.
shrub, because its leaves
vary greatly in hairiness CHINESE HIBISCUS
and size. Hibiscus rosa-sinensis
oblong f: Malvaceae
leathery leaves
Possibly native to India,
despite its name, this
tropical shrub is one of
flowering several Hibiscus species
sprig cultivated for their
showy blossoms.

ribbed
pod 15 ft/4.5 m

40 ft
12 m
COCOA
Theobroma cacao
6½ ft/2 m f: Malvaceae
fruiting Originally from Brazilian
rainforests, this tree is cultivated toothed leaf
stem
around the tropics. Cocoa comes
strongly from the seeds, known as beans,
scented inside its fruit pods.
flowers

caffeine- nutritious
containing baobab fruit
seed
32 in
80 cm mature tree
82 ft/25 m

glossy oval 82 ft
leaves 25 m
MEZEREON MUSK MALLOW COLA NUT TREE BAOBAB TREE
Daphne mezereum Malva moschata Cola nitida Adansonia digitata
f: Thymelaeaceae f: Malvaceae f: Malvaceae f: Malvaceae
Damp woods and shady gorges are Native to N. Africa and S. Europe, The caffeine-rich seeds (called This massive African tree is called
the typical habitat of this deciduous and a garden plant farther north, cola nuts) from the pods of this an upside-down tree because,
shrub, which is found across this tall perennial grows in W. African tree are chewed for when leafless, its branches look like
most of Europe. grassy and bushy places. their stimulant effects. roots. It can live for 1,500 years.
 pollen-collecting TRAILING ABUTILON
stigma Callianthe megapotamicum
f: Malvaceae 6 ft
179
Native to dry mountain 1.8 m
pollen is attached to filaments valleys of Brazil, this
on central stamen tube spreading shrub is also

FLOWERING PLANTS • EUDICOTS

popular as a colorful
ornamental in warm,
sunny gardens. glossy
leaves

spiny
fruit

130 ft
40 m
DURIAN
Durio zibethinus
f: Malvaceae
The spiny fruits of this Asian
rainforest tree smell of sweaty
socks. This attracts animals to eat 120 ft 6 ft
the fruit then disperse the 36 m 1.8 m
seeds in their droppings.
AMERICAN LIME
Tilia americana
flower f: Malvaceae
sprig This medium to large
deciduous tree adds to fall
colors in woodlands across
eastern N. America.
It often grows alongside
sugar maple.

WILD HOLLYHOCK
Alcea pallida
palmate f: Malvaceae
leaves This tall evergreen
perennial, a close relative
of garden hollyhocks,
230 ft comes from the
70 m E. Mediterranean. It
5 ft grows in rocky places
open 1.5 m
boll and scrubland.

KAPOK UPLAND COTTON

Ceiba pentandra Gossypium hirsutum
down-
filled pod
f: Malvaceae f: Malvaceae
Massive kapok trees grow wild in This C. American shrub is the most
W. Africa, and C. and S. America. common cultivated cotton species.
Fiber from their fruits, or pods, is The cotton fibers protect seeds
used as a stuffing for toys. inside the fruit, or boll.

SA PI NDA LES evergreen

leaves
The Sapindales is an important order of nine
families, mostly of trees, shrubs, and woody vines, 130 ft/40 m
often with divided leaves. It includes many dominant
woodland species and commercially important ripe
species such as citrus fruits. More than half its fruit
members belong to two families: the maple family 16 ft/5 m
(Sapindaceae), with about 1,900 species; and the rue CASHEW
Anacardium occidentale
family (Rutaceae), with 1,700 species, which mostly f: Anacardiaceae
originate in Australia and South Africa. Originally from S. America, this
shrubby tree was taken to Asia
33 ft/10 m and Africa in the 1500s, and
40 ft/12 m cultivated for its nuts.
STAG’S HORN
SUMACH SMOKE BUSH MANGO
Rhus typhina Cotinus coggygria Mangifera indica
f: Anacardiaceae f: Anacardiaceae f: Anacardiaceae
This deciduous shrub or Finely branched clusters A native of Asia, mango
small tree grows on forest of pale green flowers is one of the most widely
edges and wasteland in give this bush a smoky cultivated fruits in the
eastern N. America. It has appearance. It is found in tropical world, and is
spiky clusters of red berries. S. Europe and Asia. rich in vitamin A.
 SA PI NDA LES 6½ ft
2m
180 130 ft
40 m
FLOWERING PLANTS • EUDICOTS

80 ft
25 m
ripe pinnate
fruits leaves
CRABWOOD 180 ft NEEM TREE CHINESE MAHOGANY SCENTED BORONIA
Carapa guianensis 55 m Azadirachta indica Toona sinensis Boronia megastigma
f: Meliaceae f: Meliaceae f: Meliaceae f: Rutaceae
The dark lumber of this tropical Valued for its lumber, medicinal oils, Chinese people eat the leaves of this This erect shrub of wet sandy sites in W. Australia
S. American tree is sometimes sold and edible shoots, this tree is grown E. Asian tree as a vegetable. Furniture is has bell-like flowers, brownish on the outside
as Brazilian mahogany. Soap is throughout the Old World tropics. made from its hard, reddish timber. and golden-green inside.
made from its seeds. Indian farmers produce insecticides
from the oil and leaves. 23½ in
60 cm

23½ in
60 cm

FRINGED RUE 5 ft
Ruta chalepensis 1.5 m
f: Rutaceae
A native of rocky PEPPER AND SALT SYDNEY
habitats in S. Europe Philoteca spicata ROCKROSE
and S.W. Asia, this is f: Rutaceae Boronia serrulata
thought to be the Widespread in sandy and gravelly places in f: Rutaceae
rue mentioned in S.W. Australia, this low shrub has narrow This small shrub, in a genus
the Bible. leaves and pink, white, or bluish flowers. confined to Australia, grows in
coastal heaths near Sydney. It has
bright pink, cup-shaped flowers.

prominent
6½ ft midrib
2m

20 ft
6m 3¼ ft/1 m
HOP TREE SALMON CORREA
Ptelea trifoliata Correa pulchella
f: Rutaceae f: Rutaceae
MEXICAN ORANGE This small tree, native to eastern A native of South Australia, this
Choisya ternata N. America, is grown for ornament. small shrub is grown in gardens
f: Rutaceae Its fruits were once used as a for its delicate, pendulous,
Originating from Mexico but common substitute for hops in brewing. tubular flowers.
in gardens elsewhere, this irregular,
bushy, evergreen shrub has branched
clusters of sweet-scented, white flowers.
stem 20 ft
6m

33 ft evergreen
10 m leaves

golf ripening unripe fruit

ball-sized fruit
fruit
6½ ft/2 m 30 ft
26 ft/8 m 9m
SKIMMIA
COMMON PRICKLY ASH Skimmia japonica JAPANESE BITTER ORANGE LEMON SEVILLE ORANGE
Zanthoxylum americanum f: Rutaceae Citrus trifoliata Citrus x limon Citrus x aurantium
f: Rutaceae Much planted in gardens, f: Rutaceae f: Rutaceae f: Rutaceae
This spiny N. American tree grows as parks, and amenity areas, this The small, inedible yellow fruits of Thought to have originated by Unlike the sweet orange we eat raw
far north as Quebec, Canada. Native evergreen, aromatic shrub this spiny shrub resemble oranges hybridization in Assam or China, (C. x sinensis), this species’ bitter fruits
Americans chewed its bark to help from E. Asia produces red with a downy skin. They have this evergreen tree is now widely are only good for cooking. Both
soothe toothaches. berries in late summer. several medicinal uses. cultivated for its fruits. originated as Asian hybrids.
 52 ft JAPANESE MAPLE SYCAMORE
16 m Acer palmatum Acer pseudoplatanus
f: Sapindaceae f: Sapindaceae 181
Centuries of breeding Native to mountain
have produced many woods in Europe and
cultivated varieties of Asia, sycamore is widely
this native Japanese planted elsewhere. Its
tree, with varying leaf winged seeds spread
shapes and spectacular flowering in the wind.
fall colors. stem
SUGAR MAPLE winged seeds
Acer saccharum
f: Sapindaceae GOLDEN-RAIN TREE
This tree is native to N.E. US Koelreuteria paniculata 100 ft
and S.E. Canada. Maple syrup
40 ft
f: Sapindaceae 30 m
is made by boiling the sap This showy tree from E. Asia
collected from the tree 12 m
is much planted in temperate
115 ft in spring. regions for its cascading
35 m conker in
yellow blossoms and spiky shell
distinctive bladderlike
seed pods.
young
flowers have
yellow “eye”

100 ft 130 ft
30 m 40 m

white
spring
flowers
LYCHEE HORSE CHESTNUT
Litchi chinensis Aesculus hippocastanum
f: Sapindaceae f: Sapindaceae
Probably originating in S. China, Native to S.E. Europe, this tree is
this tree is cultivated for its often planted for shade along city
fruits. Their sweet flesh is streets. Its fruits resemble chestnuts
encased within a tough shell. but are hard and inedible.

130 ft
40 m

bright
green edible
leaves nuts
JAVA ALMOND
Canarium indicum
f: Burseraceae
Native to rainforests in
the Pacific islands, this
is one of the region’s
most useful trees,
providing lumber, oil,
toothed leaflet and edible nuts.
26 ft
8m

65 ft
20 m

redder eye of
older flower

pinnate
leaf
flower
spike 26 ft
26 ft 8m
8m
YELLOWHORN TREE OF HEAVEN BITTERWOOD FRANKINCENSE
Xanthoceras sorbifolium Ailanthus altissima Quassia amara Boswellia sacra
f: Sapindaceae f: Simaroubaceae f: Simaroubaceae f: Burseraceae
This small tree grows wild in Hailing from China, this slightly Boiled extracts from the bark Frankincense, a gum resin used in
China. Its scientific name records rank-smelling tree is planted along and leaves of this tree are used incense and to fix perfumes, oozes
its leaves, which resemble those city streets because it endures to make tonics against malaria as milky juice from cuts in
of rowan (Sorbus). pollution and most soil types. in tropical America. trunks of these Arabian trees.
 C A RYO P H Y L L A L E S
182 The Caryophyllales is a diverse order of 38 families
12 in
30 cm
of trees, shrubs, climbers, succulents, and
herbaceous plants, ranging from carnations to
cacti. Many grow in difficult environments, but
have evolved special adaptations for survival.
Some have fleshy leaves to store water in
drought conditions. Another extreme example 2 in 1 in
5 cm 2.5 cm
is the ability of carnivorous species to trap and
digest insects for extra nutrients. CONOPHYTUM MINUTUM WARTY TIGER JAWS PURPLE DEWPLANT
f: Aizoaceae Faucaria tuberculosa Disphyma crassifolium
many-petaled yellow A minute, clump-forming species, f: Aizoaceae f: Aizoaceae
or light pink flower with fleshy pebblelike leaves, Native to semidesert areas of South This procumbent species—with
this succulent perennial inhabits Africa, this species has warty leaves that stems that trail along ground—has
semidesert areas in South Africa. resemble an open mouth, hence the name. fleshy leaves and daisylike flowers.
Native to South Africa, Australia, and
New Zealand, it lives on saline soils.
4 in 16 in
10 cm 40 cm

1¼ in
3 cm

SCHWANTESIA LAMPRANTHUS AURANTIACUS TITANOPSIS CALCAREA

RUEDEBUSCHII f: Aizoaceae f: Aizoaceae
f: Aizoaceae This succulent plant grows in sandflats The encrusted leaves of this South
A clump-forming species with along the west coast of South Africa, African succulent disguise
unequal pairs of keeled, fleshy leaves, where it is known locally as “vygie.” it among desert limestone
fleshy leaf this succulent grows on hillsides in Its daisylike flowers are about rocks. It flowers in late
Namibia and South Africa. 2 in (5 cm) across. summer and fall.

1¼ in
HOTTENTOT FIG LIVING STONES 3 cm
Carpobrotus edulis Lithops aucampiae
f: Aizoaceae f: Aizoaceae
This sprawling, fleshy South Growing among pebbles
African species with showy in semidesert regions of
flowers and edible figlike fruits South Africa, this dwarf
grows in open dry habitats, clump-forming species
where it can be invasive. has bulbous leaves.

12 in
30 cm
carmine-red flower

paired leaves

4 in bulbous
10 cm leaf

3¼ in
8 cm

12 in
30 cm FAIRY GIBBAEUM
ICE PLANT ELEPHANT’S FEET VELUTINUM
Mesembryanthemum crystallinum Frithia pulchra f: Aizoaceae
f: Aizoaceae f: Aizoaceae Pairs of unequal-sized fleshy
Popularly named for the This small succulent grows in leaves, joined near their
tiny glistening swellings covering montane grassland in South bases, characterize this
the entire plant, this species inhabits Africa. In drought its leaves mat-forming species. It is
saline areas in parts of Africa, shrink, pulling the plant into the found in semidesert areas
Europe, and W. Asia. soil to protect it from desiccation. in South Africa.
 ANNUAL SEABLITE
Suaeda maritima
f: Amaranthaceae
12 in
Mainly coastal, inhabiting salt 30 cm
marshes in Europe, but growing
inland in parts of Asia and
61⁄2 ft 6 ft N. America, annual seablite has
2m 1.8 m green leaves which turn red.

8 ft/2.5 m
panicle of
LOVE-LIES-BLEEDING ACHYRANTHES tiny flowers
Amaranthus caudatus BIDENTATA
f: Amaranthaceae f: Amaranthaceae 5 ft heart-
Believed to originate in Forest edges, streamsides, and 1.5 m shaped
leaf
S. America, this annual moist shady places in China,
plant has long been Japan, India, and Nepal are
cultivated for its edible home to this species.
leaves and seeds. tall,
leafy
POLPALA stem
Aerva lanata
f: Amaranthaceae
keel, or ridge With catkinlike flower clusters, this
perennial from tropical regions of
Asia and Africa grows on open, SEA BEET
disturbed ground. Beta vulgaris
f: Amaranthaceae
The wild ancestor of the
beet, sea beet is a fleshy
3¼ ft 12 in inhabitant of bare
1m 30 cm ground by the sea in
parts of Europe,
N. Africa, and Asia.

SEA PURSLANE
Atriplex portulacoides
f: Amaranthaceae
Salt marshes, especially edges
of tidal channels and pools, are
home to sea purslane, a sprawling, COMMON GLASSWORT
silvery species from S. Europe Salicornia europaea
and the Mediterranean. f: Amaranthaceae
Common glasswort is a bushy
species of muddy salt marshes
in W. Europe. Its succulent
spike of reddish stems are sometimes cooked
flowers as a vegetable.

MEXICAN TEA
Dysphania ambrosioides
edible
salad
f: Amaranthaceae
leaf A short-lived inhabitant of
cultivated and waste ground
in tropical America, this
aromatic species is used as
white, pink, or a seasoning and in tea.
lilac flower 3¼ ft
1m

GARDEN ORACHE 4 ft
Atriplex hortensis 1.2 m
f: Amaranthaceae
Probably originally from
seashores in S.W. Asia, this
spinachlike species has
long been cultivated for its 6½ ft
edible leaves. 2m

PLUMED COCKSCOMB
Celosia argentea
paired fleshy leaf f: Amaranthaceae
This showy plant is native to dry
slopes and stony ground in tropical
Africa, but is widely planted
elsewhere for its colorful foliage.
 C A RYO PH Y L L A L E S spines protect funnel-shaped
184 slow-growing
plant
flower

PARODIA ERIOSYCE
HASELBERGII SUBGIBBOSA
FLOWERING PLANTS • EUDICOTS

f: Cactaceae f: Cactaceae
This cactus has a This spherical species
spherical stem and grows in dry, stony
funnel-shaped flowers. places, often on the
It inhabits mountainous coast in its native Chile.
parts of Brazil.

6 in 35 in
15 cm 90 cm

23 ft
7m PERUVIAN BARREL CACTUS
OLD MAN Echinocactus sp.
CACTUS f: Cactaceae
Espostoa lanata The barrel-shaped
f: Cactaceae members of this genus
Identified by long, white are confined to the
hairs on the columnar deserts of S.W. US and
stem, this is a Mexico.
slow-growing species flesh in leaf
from the hilly areas of 16 in 4 in/10 cm
40 cm stores water
Peru and S. Ecuador.
REBUTIA HELIOSA
40 ft f: Cactaceae
23½ in/60 cm 12 m
A clump-forming species with
CLEISTOCACTUS OLD MAN CACTUS brightly colored flowers,
23½ in BROOKEAE Cephalocereus senilis this native of Bolivia grows
60 cm f: Cactaceae f: Cactaceae in partially shaded,
With semi-erect or Long white hairs on mountainous habitats.
spreading fleshy the stem give this
single stems, this cactus its common GOLDEN COLUMN
species inhabits name. It is native to Weberbauerocereus
mountainous rocky areas in Mexico. johnsonii
areas of Bolivia. f: Cactaceae
This is a tall species
from Peru, where it
inhabits sandy soils.

LEUCHTENBERGIA
PRINCIPIS
f: Cactaceae
The stem of this species is either 20 ft
13 ft/4 m 6m
spherical or short and cylindrical, and
bears fragrant flowers. It inhabits
semidesert regions of N. Mexico.
52 ft
16 m
arm, or branch,
supports extra
flowers

SAGUARO CACTUS
Carnegiea gigantea MISTLETOE CACTUS
f: Cactaceae Rhipsalis baccifera
Living for up to 150 f: Cactaceae
years in desert areas of The only cactus found naturally outside the
Mexico, Arizona, and Americas, this epiphyte was perhaps carried
California, saguaro to tropical Africa, Madagascar, and
cactus is an extremely Sri Lanka by migrating birds.
tall species.

73⁄4 ft 16 ft
2.4 m 5m

62 ft
19 m QUEEN OF THE LOPHOCEREUS
NIGHT SCHOTTII
Harrisia jusbertii f: Cactaceae
GIANT CARDÓN f: Cactaceae A tall, slow-
Pachycereus pringlei Possibly of natural growing species
f: Cactaceae hybrid origin in the from Mexico
The tallest living cactus hills of Argentina and S. Arizona,
species, giant cardón and Paraguay, this this species
grows in the desert of Baja columnar cactus opens has unpleasant-
California, Mexico. Its its flowers at night, smelling flowers
flowers open at night. hence its name. that open at night.
 12 in
30 cm
28 in/70 cm flower bud 185

FLOWERING PLANTS • EUDICOTS

RAT’S-TAIL
CACTUS
Aporocactus
flagelliformis
5 ft f: Cactaceae
1.5 m Living on trees or
rocks in wooded areas
MATUCANA INTERTEXTA MONK’S HOOD CACTUS CLARET CUP HEDGEHOG of Mexico, rat’s-tail
f: Cactaceae Astrophytum ornatum Echinocereus triglochidiatus cactus has trailing
A mountain valley region f: Cactaceae f: Cactaceae fleshy stems and
of Peru is the only home of this A spherical or columnar stem with Growing in deserts, scrub, and on colorful flowers.
clump-forming species with a long, brownish yellow protective rocky slopes in S. US and N. Mexico,
5 ft
spherical or short cylindrical stem. spines characterizes this species. this variable cactus is pollinated 1.5 m
It grows in arid areas of Mexico. by hummingbirds.

8 in 8 in 4 in
20 cm 20 cm 10 cm long, thin,
trailing stem

BRAIN CACTUS
Stenocactus multicostatus GYMNOCALYCIUM
f: Cactaceae HORSTII
This spherical species f: Cactaceae
inhabits dry grasslands This spherical,
in N.E. Mexico. Funnel- clump-forming cactus
shaped buds open into pink probably grows wild only
striped flowers. in rocky grassland near
Rio Grande do Sul
in Brazil.

TURK’S CAP CACTUS OLD LADY CACTUS

Melocactus salvadorensis Mammillaria hahniana
f: Cactaceae f: Cactaceae
With a characteristic flower- Growing in semidesert regions, 15 cm
bearing structure on top of a old lady cactus is a Mexican 6 in
spherical stem when mature, turk’s species with grayish hairs on
cap cactus inhabits open, rocky the spherical stem.
ground in N.E. Brazil.
newly formed 16 ft/5 m
prickly pear fruit

12 in
30 cm
CRAB CACTUS
Schlumbergera truncata
f: Cactaceae
An epiphytic winter-flowering species of
tropical rainforest in S.E. Brazil, crab cactus
is often grown as a houseplant.

bright yellow
flower
cluster of spines

paddlelike
green stem
8 in GLORY OF TEXAS PRICKLY PEAR
20 cm Thelocactus bicolor Opuntia ficus-indica
f: Cactaceae f: Cactaceae
Native to arid regions With flattened jointed stems and edible
of Texas and N.E. egg-shaped fruits, prickly pear is a native of
Mexico, this is a rocky hillsides and dry ground in Mexico,
spherical species. and is widely naturalized elsewhere.
 SPIKY HAIRY STAR
Viewed from above, this cactus
appears star shaped, with five to ten
(usually eight) outstretched ribs, and
crossbands of woolly, white scales
across the surface between them.

clusters of 5–11
spines grow from
a small hollow, or
areole, surrounded
by white hair
 MONK’S HOOD 187
CAC T U S
spine protects the
slow-growing plant
from being eaten
A s t rophy tum or natum

FLOWERING PLANTS • EUDICOTS

The genus name for this cactus, Astrophytum, means “star plant”
in ancient Greek. It was first collected in 1827 by Thomas
Coulter, an Irish doctor and botanist, who sent it to Professor
de Candolle at the Botanic Garden of Geneva. When de Candolle
unpacked the cactus he thought it was covered in fungus, but
discovered that the white spots were actually
tufts of hair or trichomes. These woolly
scales may help the cactus collect water
and protect it against the sun, but they
also provide camouflage. This species
is the most densely covered of all
the Astrophytum cacti, and also the
spiniest. It is now rare in the wild.

size 5 ft (1.5 m)
habitat Semidesert scrub
distribution N.E. Mexico
leaf type Spines

ROOT SYSTEM WOOLLY SCALES

Fibrous and shallow roots This cactus has white,
help the cactus to absorb scalelike tufts of
water from a large area, trichomes arranged
essential in an environment in bands between its
where after brief showers ribs. They are densely
only an inch or so of woolly in young
soil is wet. plants but become
sparse with age.
OUTER PETALS
The numerous, narrow outer
petals of the flower are pale
yellow and have brown tips.
pith, used for The flower itself grows to
water storage 4½ in (11 cm) in diameter.

fibrous roots

petal

stamen (male part

that produces and
releases pollen)
stigma

OVARY
Located below all the
other reproductive parts
FLOWER SECTION of the flower, the ovary
The yellow petals have a contains ovules that
broad, oblong shape with style (female part develop into seeds.
a slightly serrated apex. The that connects the
flower has a yellow carpel (the stigma to the ovary) STIGMA
female reproductive organs, The flower has a single stigma with 7–12 lobes. This
made up of the stigma, style, is the part of the flower used to catch pollen. The
and ovary) and yellow stamens. ovary stigma is about ½ in (1.5 cm) long.
 C A RYO PH Y L L A L E S
188
FLOWERING PLANTS • EUDICOTS

23½ in 3¼ ft 23½ in 10 in 12 in
60 cm 1m 60 cm 25 cm 30 cm

DEPTFORD PINK CORNCOCKLE COW BASIL SEA SANDWORT THYME-LEAVED

Dianthus armeria Agrostemma githago Vaccaria hispanica Honckenya peploides SANDWORT
f: Caryophyllaceae f: Caryophyllaceae f: Caryophyllaceae f: Caryophyllaceae Arenaria serpyllifolia
Inhabiting dry grassland, A native of E. Mediterranean An increasingly rare weed of grain A fleshy, prostrate species, sea f: Caryophyllaceae
especially on light, sandy soils countries, corncockle was fields in Europe and Asia, this sandwort is found on coastal Found in bare or disturbed
in much of Europe, Deptford formerly a widespread annual herb has pink flowers sand and shingle in Europe, ground in Europe, temperate
pink has starry flowers with cornfield weed, but is now and bluish green leaves. Asia, and N. America. Asia, and N. America, this
toothed petals. much rarer. species gets its common
name from its tiny leaves.
32 in
ALPINE CAMPION 80 cm
Viscaria alpina
f: Caryophyllaceae
This native of the European Alps,
pink, or occasionally Pyrenees, and subarctic areas of 12 in
white, flower Europe, W. Asia, and N. America 30 cm
grows on mineral-rich rocks.

flower stalk
lengthens as
flower matures 8 in
20 cm

five-
petaled
four or five small green flower
leaves per shoot lanceolate
leaves
RAGGED ROBIN FIELD MOUSE-EAR
Silene flos-cuculi Cerastium arvense
f: Caryophyllaceae f: Caryophyllaceae
4 in/10 cm Divided petals give ragged This is a native of dry grassland
robin its name. It is a European in Europe, N. Africa, N. America,
species found in marshy fields and W. temperate Asia.
and damp ground.

inflated
calyx

32 in/80 cm

MOSS CAMPION BLADDER CAMPION

Silene acaulis Silene vulgaris
f: Caryophyllaceae f: Caryophyllaceae
Cushion-forming and mosslike, Named for its inflated calyx,
moss campion grows on mountains bladder campion inhabits open
central cushion has in W., C., and N. Europe, as well as grassy places in Europe, N. Africa,
long tap root Asia, and N. America. and temperate regions of Asia.
 ROUND-LEAVED
sheath SUNDEW
around Drosera rotundifolia
flower f: Droseraceae
Inhabiting open, boggy
moorland and heathland

FLOWERING PLANTS • EUDICOTS

in Europe, N. Asia, and
20 in N. America, this
50 cm insectivorous species has
leaves with numerous
sticky hairs covered
in insect-dissolving
enzymes.
leaf hairs
trap insects

4 in
10 cm
20 in 3¼ ft
50 cm 1m

CHILDING PINK SOAPWORT COMMON CHICKWEED

Petrorhagia nanteuilii Saponaria officinalis Stellaria media
f: Caryophyllaceae f: Caryophyllaceae f: Caryophyllaceae VENUS FLY-TRAP
The flowers of childing pink open one at Once used to make soap, this species This sprawling species is found on cultivated
Dionaea muscipula
a time. This species inhabits dry, grassy is found along streams and damp and open ground worldwide and is
f: Droseraceae
places on sandy soils in W. Europe. ground in S. Europe and W. Asia. sometimes used as a salad vegetable. An insectivorous perennial
species from coastal bogs in
N. and S. Carolina, Venus
white flowers trap triggered by hairs fly-trap has hinged,
surrounded by bracts on leaf surface two-lobed leaves.

brightly colored
bract

when triggered,
26 ft/8 m leaf snaps shut, 6 in
trapping insect 15 cm

“teeth” prevent
insect from escaping
BOUGAINVILLEA
Bougainvillea glabra
f: Nyctaginaceae
What seem to be petals on nectar glands on
bougainvillea are actually lid attract insects
bracts. This slow-growing
evergreen climber from Brazil
is widely cultivated.

insects slither
12 in/30 cm off slippery rim 40 ft
into pitcher 12 m

SEA HEATH
Frankenia laevis
f: Frankeniaceae
A mat-forming species, sea heath
grows on bare sandy ground in
the drier parts of salt marshes in
W. Europe and N.W. Africa.

fluid in base of pitcher

digests insects

FOUR O’CLOCK FLOWER PITCHER PLANT

Mirabilis jalapa Nepenthes stenophylla
f: Nyctaginaceae f: Nepenthaceae pitcher
From dry, open habitats in In the forests of Borneo, grows from
tropical C. and S. America, this this scrambling vine gets tendril
species has fragrant flowers which extra nutrients by trapping at end
open during the late afternoon. 3¼ ft/1 m insects in its pitchers. of leaf
 C A RYO PH Y L L A L E S BLACK BINDWEED
Fallopia convolvulus
190 f: Polygonaceae
Growing on waste and
cultivated ground, black 3¼ ft/1 m
12 in/30 cm
bindweed is native to much
FLOWERING PLANTS • EUDICOTS

of Europe, N. Africa, and

temperate Asia.

61⁄2 ft
2m

wavy- 23½ in
edged seed
leaf stalk 60 cm
6½ ft/2 m

SULPHUR FLOWER ST. CATHERINE’S LACE

Eriogonum umbellatum Eriogonum giganteum
f: Polygonaceae f: Polygonaceae 23½ in
60 cm
Inhabiting well-drained montane With long-lasting flowerheads
forest and scrub in Canada and much visited by butterflies,
the N. and W. US, this this plant grows wild only
spreading, mat-forming species on the Channel Islands
has long-lasting flowers. of California.
COMMON BUCKWHEAT CURLED DOCK KNOTGRASS
small, green Fagopyrum esculentum Rumex crispus Polygonum aviculare
reddish flowers flowers f: Polygonaceae f: Polygonaceae f: Polygonaceae
pollinated Buckwheat originated in temperate Curled dock inhabits grassy places, This spreading species of open
by wind areas of Asia, and is cultivated for waste ground, and coastal shingle cultivated and bare ground in
its seeds, which are made into flour in Europe, Asia, and N. Africa. Europe and Asia grows on
seed and also fed to birds. It can be an invasive species. coasts and inland.
stalk

round
leaf
fleshy, kidney-
shaped leaf
12 in 35 in 40 ft 10 ft
30 cm 90 cm 12 m 3m

MOUNTAIN SORREL COMMON BISTORT CORAL VINE POKEWEED

Oxyria digyna Persicaria bistorta Antigonon leptopus Phytolacca americana
f: Polygonaceae f: Polygonaceae f: Polygonaceae f: Phytolaccaceae
Inhabiting damp ledges and streamsides on Inhabiting grassy areas in much This fast-growing species With poisonous fruits resembling
mountains in Arctic and temperate regions of Europe and C. Asia, common climbs by means of tendrils. blackberries, this unpleasant-smelling
of the northern hemisphere, mountain bistort has dense cylindrical It lives in tropical forests and perennial is native to open and shady
sorrel is often tinged red. flower spikes. scrub in Mexico. parts of eastern N. America and Mexico.

8 ft CHINESE 2 in
2.5 m RHUBARB 5 cm
Rheum palmatum
f: Polygonaceae
With enormous roots
and large, poisonous
leaves, Chinese 20 in
rhubarb inhabits 50 cm
streamsides and damp
ground in mountainous
areas in China.

large leaf
supported by
strong stem

SUMMER PURSLANE
Portulaca oleracea PHEMERANTHIS SEDIFORMIS
f: Portulacaceae f: Talinaceae
Native to the Mediterranean region and Africa, The flowers of this prostrate species open
but widely introduced elsewhere, purslane is an in the afternoon. Dry grassland and
edible, fleshy species of disturbed ground. scrub in western N. America are its home.
 FRINGED
REDMAIDS
Calandrinia ciliata 191
f: Montiaceae
12 in Native to grasslands from
30 cm S. Canada to Argentina,
this species has escaped to
the Falkland Islands and is
also grown in gardens.

11⁄2 in 12 in
4 cm 30 cm
SPRING BEAUTY
Claytonia perfoliata
f: Montiaceae
Also known as miner’s lettuce,
LEWISIA BRACHYCALYX this native of cultivated and
f: Montiaceae bare ground in western
Native to damp, stony meadows in mountainous N. America, Mexico, and
areas of S.W. US, this species has a basal Guatemala has two fused
rosette of fleshy leaves. leaves beneath the flowers.

CEYLON
LEADWORT
Plumbago zeylanica 5 ft
f: Plumbaginaceae 1.5 m
Growing mainly
in open, human-
created habitats,
this scrambling
shrub is found
throughout tropical
and subtropical
regions.

10 in 10 ft 6½ ft
25 cm 3m 2m
TAMARISK
STATICE SEA PINK Tamarix gallica JOJOBA
Limonium sinuatum Armeria maritima f: Tamaricaceae Simmondsia chinensis
f: Plumbaginaceae f: Plumbaginaceae Usually coastal, tamarisk f: Simmondsiaceae
Statice has winged, branching In much of W. Europe, thrift, or also grows in saline areas Deserts in Arizona,
stems and is found in rocky sea pink, inhabits coastal rocks inland. It originates from California, and Mexico
and sandy coastal habitats and and cliffs, salt marshes, and S. Europe, N. Africa, and the are home to jojoba, a
16 in in saline areas inland in mountains. It is a cushion- Canary Islands, and is species often
40 cm Mediterranean countries. forming species. planted elsewhere. cultivated for its oil.

SA N TA L A L E S
OSYRIS
Found mainly in the tropics and subtropics, the seven Osyris alba
families in the order Santalales include several important f: Santalaceae
lumber trees. They also feature many parasitic and berry can be This broomlike,
poisonous 30 ft
semiparasitic species
semiparasitic plants, notably the 900 species in the 9m
with fragrant flowers
showy mistletoe family (Loranthaceae) of the southern is native to S. Europe,
hemisphere. These live attached to other plants, from N. Africa, and S.W.
which they obtain all or most of the water and nutrients Asia, inhabiting
they need for growth. dry, rocky places.

SANDALWOOD
31/4 ft Santalum album
1m f: Santalaceae
Cultivated for its wood and
33 ft/10 m fragrant oil, this semiparasitic
tree grows in dry, rocky areas in
Asia and Australia.

FIRE TREE
Nuytsia floribunda
f: Loranthaceae MISTLETOE
This species is semiparasitic, Viscum album 4 ft
deriving moisture and nutrients f: Santalaceae 1.2 m
from the roots of surrounding Mistletoe forms spherical masses on tree branches.
plants in woodland in A semiparasitic species with white berries, it
S.W. Australia. inhabits much of Europe, N. Africa, and Asia.
 CORNALES FLOWERING
DOGWOOD ERICALES
192 Seven families constitute the revised order Cornales.
Cornus florida 40 ft Comprising 22 families, the Ericales is
f: Cornaceae 12 m
Five of these are small and relatively insignificant, The showy white a major order, both economically and as
one with just a single species of evergreen tree from “petals” of this one of the world’s main herbaceous groups.
Africa. The main family is the dogwood family N. American dogwood It is named after its largest member, the
tree are bracts
(Cornaceae), a loose taxonomic grouping of shrubs and (specialized leaves) heather family (Ericaceae), with more than
small trees found in temperate zones and on tropical around clusters of 4,000 species of flowering shrubs mainly
mountains. The hydrangea family (Hydrangeaceae) tiny flowers. in acid soils. It also includes 900 species
includes several popular garden plants. in the primrose family (Primulaceae),
largely confined to north temperate
HYDRANGEA mountain regions; herbs and shrubs in
Hydrangea macrophylla the Jacob’s ladder family (Polemoniaceae);
f: Hydrangeaceae and carnivorous plants in the pitcher plant
Native to Japan, this family (Sarraceniaceae).
showy shrub is cultivated DOVE TREE
for its rounded clusters of
Davidia involucrata
rose, lavender, blue, and
f: Nyssaceae
even white flowers. This small, flowering
tree from China is
impressive in bloom,
with flowerheads ¾ in (2
5 ft/1.5 m
cm) across, surrounded
by creamy bracts.
80 ft
25 m
MOCK ORANGE 3¼ ft
Philadelphus sp. 1m
f: Hydrangeaceae
About 45 species of this
shrubby genus, named for their
orange-blossom fragrance, are
native to Asia, western N.
America, and Mexico.

JACOB’S LADDER
Polemonium caeruleum
f: Polemoniaceae
This tall perennial has cup-shaped lavender
or white flowers. It grows in rocky and
grassy places in Europe and N. Asia.

15 ft
4.5 m 3 ¼ ft/1 m

PERENNIAL PHLOX
Phlox paniculata
f: Polemoniaceae
A pyramidal cluster of trumpet-shaped, pink or
lavender flowers top the stem of this perennial,
native to open woods in the S.E. US.
 193

FLOWERING PLANTS • EUDICOTS

3¼ ft 231⁄2 in
1m 60 cm
HEATHER DORSET HEATH
Calluna vulgaris Erica ciliaris 40 ft
f: Ericaceae f: Ericaceae 12 m
This evergreen shrub, with One of several similar spiky
spikes of pale purple flowers, ericas, this species is known
dominates vast areas of from S. England (including
moorland in N. Europe, Dorset), as well as W. Ireland,
eastward into Asia. France, Iberia, and Morocco.

scarlet,
bell-shaped
flower STRAWBERRY TREE
white, Arbutus unedo
bell-shaped
flowers
f: Ericaceae
100 ft/30 m Although related to heathers,
the warty, red fruits of this
acidic evergreen Mediterranean tree
TREE berries look remarkably like
RHODODENDRON strawberries but are tasteless.
Rhododendron arboreum 16 ft 12 in COWBERRY
f: Ericaceae 5m 30 cm Vaccinium vitis-idaea
There are about 1,000 species f: Ericaceae
of Rhododendron, most with This neat heather, with leathery leaves and
leathery, evergreen leaves and HIMALAYAN glossy red berries in the fall, is found across thick trunk covered
showy flowers. Many, like this PIERIS N. Europe, Asia, and N. America. in small leaves
one, come from the Himalayas. Pieris formosa
f: Ericaceae
Formosa means “beautiful,” 6 in
a reference to the hanging 15 cm
clusters of white,
urn-shaped flowers on this CHECKERBERRY
Asian bush or small tree. Gaultheria
10 ft
procumbens
3m
f: Ericaceae
This creeping, aromatic
shrub forms patches
beneath oaks and
conifers in eastern N.
deciduous America. Its red fruits
leaves
SWEET PEPPER last through the winter.
BUSH
Clethra alnifolia 65 ft
orange 20 m
f: Clethraceae fruits
This deciduous shrub PERSIMMON
grows in wet forests Diospyros virginiana
and bogs in eastern f: Ebenaceae 6 1/2 ft/2 m
N. America. Its 65 ft
leaves turn yellow This native N. America tree has yellowish white, 20 m
or orange in the fall. bell-shaped flowers and produces globular orange
fruits, about 11/2 in (4 cm) across.
INDIAN BALSAM
165 ft Impatiens glandulifera
fruits 50 m f: Balsaminaceae
33 ft Exploding fruit pods, which shoot
10 m round, their seeds outward, have helped this
woody fruit Himalayan species become established
by rivers in most of Europe.

black seeds
BOOJUM TREE
Fouquieria columnaris
f: Fouquieriaceae
This strange tree is
mainly confined to
green Baja, California.
flesh KIWI BRAZIL NUT Its stem and upright,
Actinidia chinensis Bertholletia excelsa spiny branches are
open f: Actinidiaceae f: Lecythidaceae green and can
fruit Also known as Chinese gooseberry, this Brazil “nuts” are actually seeds inside this photosynthesize.
multiple
woody climber was introduced from S. American tree’s hard, cannonball-like “nuts” inside
China to New Zealand, where the fruits. In the wild, agoutis (large
berries were sold as “kiwi fruits.” rodents) open the fallen fruits.
 194
FLOWERING PLANTS • EUDICOTS

FLOWER
HOODED PITCHER The flowers have five sepals, five
yellow petals (which redden as they
PLANT mature), and a whitish, umbrella-
like style. They hang upside down.
Sar racen ia minor petal
This insectivorous plant is highly adapted to living
in nutrient-poor, acidic swamps and bogs, digesting
insects to obtain phosphorus and nitrogen
from their bodies. Insects are attracted by ovary forms
seed capsules
nectar into the pitcher, the long, thin, tubular, when fertilized SEED CAPSULE
modified leaf from which the plant The broad, rough
gets its name. The interior of the capsules crack open
bowl-shaped style
from which insects and scatter many
pitcher is covered in a slippery, flower held well
above deadly collect pollen tiny knobbly seeds,
waxy coating and downward- pitchers which are about
pointing hairs that become ⅛ in (3 mm) long.
coarser nearer the bottom
of the tube. The insects
fall in and, unable to get
out, eventually die from
exhaustion. Digestive
glands in the tube walls
RHIZOME
exude enzymes which The plant’s domed
break down the insects, pitchers rise up from a
resulting in a liquid slurry horizontally branching,
which is then absorbed underground stem
known as a rhizome.
into the leaf, providing
the pitcher plant with
valuable nutrients.

INSECT TRAP
Insects caught in the
pitcher find it very
ROOTS difficult to escape.
The fibrous roots The leaf digests the
are 8–12 in soft parts, but the
(20–30 cm) long skeletons remain.
size Up to 12 in (30 cm) tall and grow along the
habitat Swampy areas in length of the “WINDOWS”
pinewoods and savannas rhizome. Insects fly toward the bright
distribution S.E. US translucent “windows” or areoles
leaf type Hooded, narrow, on the upper part of the pitcher,
funnel-shaped “pitcher” but then slide down into the tube.
 IN BLOOM
The yellow, odorless flowers
are produced from late March
to mid-May. Bees are the main
reddish purple colors pollinator, and will favor this
of the hood attract species over other nearby plants.
insects to the pitcher

HOODS
The concave hoods prevent
rainwater from filling up the
pitcher and also shade the
opening, making it harder for
trapped insects to find the exit.

CARNIVORE AT RISK
The hooded pitcher plant is now a rare and nectar glands
threatened species due to habitat loss. It is at the back
found in North Carolina, South Carolina, of the hood
Georgia, and Florida. There are two
varieties: S. minor grows to 12 in (30 cm),
while S. minor var. okefenokeensis, with
pitchers up to 4 ft (1.2 m) long, is found
only at Okefenokee Swamp in Georgia.
 ERICALES
196
FLOWERING PLANTS • EUDICOTS

15 ft 30 ft/10 m
4.5 m

GRANTHAM’S CAMELLIA
Camellia granthamiana
f:Theaceae
One of well over 100 Camellia species,
all from Asia, this endangered tree
from China was discovered in 1955.

paired oval leaf

with rounded tips
40 ft/12 m

EPAULETTE TREE
Pterostyrax hispidus 56 ft/17 m
f: Styracaceae
Native to E. Asia, this
large-leaved, deciduous tree is SILKY CAMELLIA
sometimes cultivated for its Stewartia malacodendron
hanging clusters of fragrant, f:Theaceae TEA
creamy-white flowers. Named after its silkily Camellia sinensis
haired young stems, f:Theaceae
this deciduous shrub or The dried, fermented leaves and buds of 231⁄2 in/60 cm
tree grows in woodland this evergreen Asian shrub are used to
in S.E. US. make tea. Their astringent tannins
100 ft/30 m deter grazing animals.

14 in
35 cm

6 ft
1.8 m

12 in
30 cm
CORALBERRY
Ardisia crenata
f: Primulaceae
A greenhouse favorite horizontal,
SPANISH CHERRY HOODED PARROT for its long-lasting, bright
Mimusops elengi PITCHER PLANT PITCHER PLANT creeping stem
berries, this Asian shrub
f: Sapotaceae Sarracenia minor Sarracenia psittacina has become an invasive
This evergreen Indian tree is f: Sarraceniaceae f: Sarraceniaceae pest in Hawaii, Florida,
planted in tropical countries Pitcher plants trap and Sarracenia species grow wild only and Texas.
for its fragrant flowers. Its digest insects to supplement in N.E. America. The pitchers
durable wood is used in their nutrient-poor soils. of this species, found from
shipbuilding and construction. This species is native Florida to Louisiana, form
to the S.E. US. horizontal traps.
SCARLET PIMPERNEL
Anagallis arvensis
f: Primulaceae
3¼ ft/1 m hood resembles Just as likely to have deep blue as
cobra ready to strike scarlet flowers, this sprawling
plant now grows in disturbed
COBRA PLANT ground virtually worldwide,
Darlingtonia californica except the tropics.
f: Sarraceniaceae
The only species of
Darlingtonia, this carnivorous
plant grows in coastal
bogs and beside mountain
streams in W. US.

12 in
30 cm
6 in CREEPING JENNY
15 cm Lysimachia nummularia
HAIRY ANDROSACE f: Primulaceae
Androsace villosa From Sweden to the Caucasus,
f: Primulaceae the spreading stems of this
In a genus of arctic-alpines, this perennial creep among
perennial of mountains from Europe vegetation in damp grassy
to the Himalayas is known for its dense places, shady hedgerows,
white flowerheads and silky leaves. and streamsides.
 BOR AGI NA L ES
Because of uncertainties over how the
197
forget-me-not family (Boraginaceae) relates TANSY PHACELIA
Phacelia tanacetifolia
to other families, a new order, Boraginales, f: Boraginaceae

FLOWERING PLANTS • EUDICOTS

12 in was proposed in 2016, just for this one This annual of dry places in
30 cm family of about 2,700 species. They range S.W. US flowers prolifically
from small annual herbs to large trees, often after rain. More than 160
MOSQUITO BILLS members of its genus are
Primula hendersonii with conspicuous hairs and swollen bases 3¼ ft
found in N. America.
f: Primulaceae on their stems or leaves. Some species are 1m
flower with One of a group of North American edible, and dyes are produced from others.
five petals primroses called “shooting stars”
from their rocket or dart shapes, this
species has magenta to white flowers.

6 in
WILD PRIMROSE 15 cm
Primula vulgaris
f: Primulaceae
Supposedly the prima rosa,
or “first flower” of spring,
this perennial grows in 12in 231⁄2 in 4 ft
30cm 60 cm 1.2 m
woodland clearings and
hedgebanks throughout
W. and S. Europe.
COMMON LUNGWORT BORAGE COMMON COMFREY
Pulmonaria officinalis Borago officinalis Symphytum officinale
f: Boraginaceae f: Boraginaceae f: Boraginaceae
With blotched leaves supposedly Stiff hairs cover the Symphytum means “growing
resembling unhealthy lungs, and once stem and leaves of this together,” named because this
thought to be efficacious against S. European wayside plant, perennial of damp ground across
NORTHERN PRIMROSE tuberculosis, this perennial grows in also cultivated for ornament Eurasia was traditionally used in
Primula scandinavica shady places throughout C. Europe. and its oil-rich seeds. the treatment of injuries.
f: Primulaceae
Growing on mountain slopes in Norway and
Sweden, this resembles the more common VIPER’S-BUGLOSS
European bird’s-eye primrose, Echium vulgare
but is much smaller. f: Boraginaceae
Widespread in grassy places
6 in 28 in in Europe and temperate
15cm 70 cm Asia, this rough-haired
231⁄2 in
60 cm biennial has pinkish flower
buds but deep blue flowers.

rough, hairy,
12 in lanceolate leaves
30 cm

WATER PURPLE
FORGET-ME-NOT GROMWELL
Myosotis scorpioides Lithospermum
COWSLIP f: Boraginaceae purpureocaeruleum
Primula veris This surface-breaking f: Boraginaceae
f: Primulaceae plant of streams and ponds Upright flower stems grow
A typical primrose, with a head of is widely distributed in from creeping ground stems
up to 30 nodding flowers, cowslip Eurasia, and introduced in this woodland perennial 3¼ ft
grows in lime-rich meadows in to the Americas and found on chalky soils in 1m
S. Europe and temperate Asia. New Zealand. S. Europe and S.W. Asia.

G A R RYA L E S SILK-TASSEL BUSH

Garrya elliptica
Any classification system f: Garryaceae
Growing in coastal scrub
4 in raises anomalies, and the order in California and Oregon,
10 cm Garryales is one of these. this small tree has showy,
Once placed in the Cornales, gray-green male catkins and
modern genetic investigation shorter, silvery
female catkins.
separated this order of
two families and about 16 ft/5 m
HARDY CYCLAMEN 20 species. The silky tassel 16 ft
Cyclamen hederifolium family (Garryaceae) includes
5m
f: Primulaceae SPOTTED LAUREL
Cyclamen leaves grow two genera: Garrya from Aucuba japonica
from underground stems, N. America and Aucuba from f: Garryaceae
and their tubular fall E. Asia. The Eucommiaceae Some cultivars of this ornamental
flowers have back-folded covers just one species: Japanese shrub grow yellow-blotched
petals. This species grows leaves. Plants are either male, with
in bushy, shady places Eucommia ulmoides, a tree of upright flower spikes, or female with
in S. Europe. Chinese mountain forests. small, clustered flowerheads.
 GENTIANALES
198 The order name celebrates the gentians of mountains and gardens, 40 ft
12 m
but the Gentianaceae family has only about 1,600 members. The
madder family (Rubiaceae) is the largest in the order, with more
FLOWERING PLANTS • EUDICOTS

than 13,000 species, including tropical shrubs such as coffee. The oval
oleander family (Apocyanaceae), strychnine family (Loganiaceae), leaf
3¼ ft
and jessamine family (Gelsemiaceae) complete the order. 1m five- COMMON
petaled FRANGIPANI
flower
Plumeria rubra
MADAGASCAR
f: Apocynaceae
PERIWINKLE Native from Mexico to Venezuela, the flowers of
Catharanthus roseus this ornamental tree are fragrant at night and
f: Apocynaceae attract sphinx moths for pollination.
The leaves of this garden ornamental,
endangered in its native Madagascar,
contain tiny amounts of alkaloids 6 in/15 cm
5 ft used to treat childhood leukaemias.
1.5 m This use has saved it
from extinction.

13 ft/4 m
BEAD PLANT
GREATER OLEANDER
Nertera granadensis
PERIWINKLE Nerium oleander f: Rubiaceae
Vinca major f: Apocynaceae Named for its small, beadlike fruits,
f: Apocynaceae All parts of this evergreen 30 in
this perennial with tiny, green flowers
Long, rooting shrub are poisonous. It is native to Australia, New Zealand,
75 cm Pacific islands, and S. America.
stems of this evergreen grows by streams from the
perennial creep across Mediterranean to China,
woodland floors in producing clusters of
S. and C. Europe fragrant pink flowers. incurved,
and N. Africa. rudimentary leaf
BUTTERFLY WEED
Asclepias tuberosa
f: Apocynaceae
10 ft succulent
3m Native Americans chewed the roots of this showy
CLEAVERS perennial as a cure for pleurisy. It grows in fields stem
Galium aparine and on roadsides in N. America.
f: Rubiaceae glossy,
Prickles on the stem evergreen
and leaf margins leaf CROSSWORT
attach to animal fur, Cruciata laevipes
dispersing parts f: Rubiaceae
of this agricultural With stem leaves in
weed of Europe crosslike whorls and
and Asia. clustered, honey-
scented flowers, this
perennial grows in
grassy places across 231⁄2 in
Europe and Asia. 60 cm

50 ft
15 m
QUININE TREE
Cinchona calisaya
f: Rubiaceae
The anti-malaria drug quinine comes
from the bark of this S. American tree.
Seeds were smuggled to Asia in the
mid-19th century for cultivation there.

6½ ft
33 ft/10 m 2m
COFFEE SHRUB
Coffea arabica CAPE JASMINE
f: Rubiaceae Gardenia jasminoides
Native to Ethiopia but transported f: Rubiaceae
elsewhere for cultivation, this Originating in Asia, this evergreen
evergreen shrub produces crimson shrub has fragrant, waxy, tubular
drupes (stone fruits) within which white flowers that turn yellow with
fruits red when ripe are two seeds (coffee beans). age. These ripen to berrylike fruits.
 oval 12 in
leaves 30 cm
20 in
50 cm
5 in
12 cm

FLOWERING PLANTS • EUDICOTS

82 ft
25 m
strychnine
STRYCHNINE seeds PERSIAN VIOLET
Strychnos nux-vomica Exacum affine
f: Loganiaceae f: Gentianaceae
The poison strychnine comes Grown as an indoor ornamental but native to
from seeds of this evergreen Yemen and nearby Socotra Island, this evergreen
tree from S.E. Asia. biennial has purple flowers with yellow centers.

SPRING GENTIAN COMMON

Gentiana verna CENTAURY
f: Gentianaceae Centaurium erythraea
With ground-level leaf f: Gentianaceae
rosettes and deep blue, tubular Grassy places and
flowers, this perennial grows dunes from Europe to
in Arctic and mountain S.W. Asia are home
regions of Europe and W. Asia. to this annual with
funnel-shaped,
five-lobed pink flowers.

12 in
30 cm

basal rosette
of leaves
flower’s center emits
stench of rotting meat 20 ft
6m
MADAGASCAR
JASMINE
CARRION FLOWER Stephanotis floribunda
Stapelia gettlifei f: Apocynaceae
f: Apocynaceae Originally from Madagascar,
Carrion flower is a succulent, this woody, twining vine is
spiny plant from southern Africa. a popular greenhouse plant
hairy edge The stench of its barred or with leathery leaves and
to petals blotched flowers attracts carrion clusters of fragrant, waxy,
flies as pollinators. white flowers.

LAMIALES 5 ft
stout
flower
crinkly
lobed
1.5 m
spike leaf
Modern taxonomy has expanded the
scope of this order to include 25 families,
typically with tubular flowers and
unequal petal lobes. The largest are
the mint family (Lamiaceae, often
known by its older name Labiatae) and 3 1/4 ft
the figwort family (Scrophulariaceae), 1m
each with 5,000–6,000 species. Others
include the olive family (Oleaceae)
and plantain family (Plantaginaceae). BEAR’S
BREECHES
Acanthus mollis
f: Acanthaceae
dark green leaf This robust perennial of rocky places in
31⁄4 ft with creamy veins the W. Mediterranean has spikes of
1m purple-veined white flowers with a
three-lobed lower lip.

50 ft
SHRIMP BUSH 15 m
Justicia brandeegeana
f: Acanthaceae ZEBRA PLANT BLACK
This popular ornamental Aphelandra squarrosa MANGROVE
from Mexico has spikes of f: Acanthaceae Avicennia germinans
white flowers surrounded Originating from coastal Brazilian f: Acanthaceae
by overlapping red- forests, this popular houseplant This tree forms thickets beside tidal
bronze bracts (specialized has pale-veined leaves and spikes estuaries along tropical Atlantic
leaves), looking like a of yellow flowers surrounded coasts. Pointed fruits, falling into
large shrimp. by yellow bracts. the mud, sprout new plants.
 LAMIALES COMMON
LAVENDER
200 Lavandula angustifolia
BUGLE f: Lamiaceae
12 in Ajuga reptans This is another evergreen shrub of
30 cm 6½ ft/2 m
f: Lamiaceae dry Mediterranean scrub habitats.
FLOWERING PLANTS • EUDICOTS

Growing around Europe, ROSEMARY The water-saving oils in its leaves

N. Africa, and S.W. Asia, Salvia rosmarinus are valued in perfumery.
this woodland and f: Lamiaceae
meadow perennial From dry Mediterranean
produces its flower stems habitats, the oils in the densely
from far-creeping, packed
leaves of this shrub reduce flowers
rooting runners. potential water loss from
transpiration. Rosemary is a
well-known culinary herb. 32 in
80 cm
tubular flower
with large, late
three-lobed lip summer
flowers

oval leaf,
often bronzy 32 in 32 in
underneath 231⁄2 in/60 cm 80 cm 80 cm

SAGE
Salvia officinalis edible leaves
f: Lamiaceae
BASIL
This grayish shrub Ocimum basilicum BETONY
from S.W. Europe and f: Lamiaceae Betonica officinalis
the Balkans is widely f: Lamiaceae
cultivated for its pungent Also known as sweet basil, this is an annual
leaves, used to from India and Iran. In cultivation, its A common hedgerow and grassland
flavor foods. pungent leaves are used as a culinary herb. herb in most of Europe, the Caucasus,
and N. Africa, betony has reddish
purple or white flowers.

branched cluster
of pinkish-
20 ft/6 m purple flowers

3¼ ft/1 m

lanceolate WILD
leaf MARJORAM
Origanum vulgare
5 ft
cluster of 4 ft f: Lamiaceae
1.5 m flowers 1.2 m A perennial aromatic herb
CHASTE TREE JERUSALEM SAGE WILD BERGAMOT of grassy and stony habitats
Vitex agnus-castus Phlomis fruticosa Monarda fistulosa in Europe and W. Asia,
f: Lamiaceae f: Lamiaceae f: Lamiaceae marjoram is cultivated for
Chaste tree grows in damp places from S. Europe Native to dry, rocky habitats in the A minty tea is made from the leaves many culinary uses.
to Pakistan. It was once thought to preserve chastity, E. Mediterranean, this evergreen of this showy American perennial of
and is used to regulate hormonal functions shrub with gray-felted leaves is a dry fields and thickets from New
in herbal medicine. widely grown garden ornamental. England to Texas. stalked,
oval leaf

3¼ ft
1m

10 ft/3 m 16 in/40 cm
COMMON THYME WATER MINT
Thymus vulgaris Mentha aquatica
ROUND-LEAVED MINT BUSH f: Lamiaceae f: Lamiaceae
Prostanthera rotundifolia Native to dry, rocky places in the Growing part-submerged in
f: Lamiaceae W. Mediterranean, this densely ponds and ditches in Europe,
This aromatic Australian shrub with circular leaves branched shrub is used as a Africa, and S.W. Asia, this
and pink to purple spring flowers grows in open culinary herb, and an ingredient herb is one parent of the
forests from New South Wales to Tasmania. in perfumes and soaps. hybrid peppermint.
 flower
sprig
201

FLOWERING PLANTS • EUDICOTS

egg-shaped
fruit
OLIVE
Olea europaea
f: Oleaceae
The fleshy fruits of this evergreen
Mediterranean tree contain 40 percent
unsaturated oils. Before eating, the
50 ft/15 m olives must be pickled in brine. heart-
shaped leaf

65 ft/20 m
10 ft/3 m

GOLDEN BELL MANNA ASH

Forsythia suspensa Fraxinus ornus
f: Oleaceae f: Oleaceae
Also called weeping forsythia, Unusually for an ash, this leaf up
this deciduous shrub has S. European tree has clusters to 5 in dense, pyramid-
hollow, pendulous stems and of showy, white flowers. (12 cm) long shaped cluster of
yellow flowers. It is native to Manna, a gum from its bark, scented flowers
23 ft/7 m
China and perhaps Japan. is used medicinally.
LILAC
Syringa vulgaris
12 in
30 cm
f: Oleaceae
Gardeners have developed
many hybrids and varieties
of this showy, deciduous trumpet-
shaped
tree, found wild on scrubby flower
hill slopes in S.E. Europe.
rosette
of sticky
leaves

40 ft COMMON JASMINE COMMON

12 m Jasminum officinale BUTTERWORT
f: Oleaceae Pinguicula vulgaris 7 in
Widely cultivated for its f: Lentibulariaceae 18 cm
showy, scented flowers, This bog plant from
this scrambling, deciduous N. Europe, Asia,
shrub, found from the and N. America
FALSE AFRICAN Caucasus to China, gains nutrients by
VIOLET twists counterclockwise trapping and digesting
Streptocarpus saxorum as it climbs. insects on its
f: Gesneriaceae sticky leaves.
40 ft
Originating from Kenya and 12 m
Tanzania, this evergreen
perennial has whorls of
small, hairy, almost succulent
leaves and five-lobed, 50 ft
trumpet-shaped flowers. 15 m

60 ft/18 m

beanlike
pods
231⁄2 in
6 in 60 cm
15 cm
HARDY GLOXINIA JACARANDA TRUMPET VINE
INDIAN BEAN TREE Incarvillea delavayi Jacaranda mimosifolia Campsis x tagliabuana
AFRICAN VIOLET Catalpa bignonioides f: Bignoniaceae f: Bignoniaceae f: Bignoniaceae
Streptocarpus ionanthus f: Bignoniaceae A hardy garden perennial Originally from Argentina A garden hybrid between
f: Gesneriaceae sprig with Despite its common name, with rosy-purple trumpet to Bolivia, this tropical tree a N. American and Asian
flower
Endangered in the rainforests of cluster this woodland tree grows in flowers, this herbaceous with hanging clusters of species, this vigorous
tropical E. Africa but popular as the S. US. It is planted farther plant grows wild in lavender-colored flowers is climbing shrub has clusters
houseplants, African violets were once north and in Europe for mountain grasslands widely planted for shade of orange-red, trumpet-
grouped in the genus Saintpaulia. its showy flowers. of China. and ornament. shaped flowers.
 LAMIALES 30 in dense
flower head
lance-
shaped leaf
broad,
oval leaf
greenish
flower spike
75 cm
202
GREATER
PLANTAIN
MARE’S-TAIL RIBWORT Plantago major
Hippuris vulgaris PLANTAIN f: Plantaginaceae
FLOWERING PLANTS • EUDICOTS

f: Plantaginaceae Plantago lanceolata This variable plantain

Wholly or partly f: Plantaginaceae grows in open habitats,
submerged, this aquatic A plant of grassy places, including trampled
perennial from Europe, with narrow, oblong, ground in gardens,
Asia, Africa, and N. and strongly veined leaves and throughout Europe,
S. America has whorls of a cylindrical flowerhead, N. Africa, and
narrow leaves up its stem. A this perennial grows in N. and C. Asia.
rapid grower, mare’s-tail is 31⁄4 ft most temperate regions. 20 in
a major garden pest. 1m 50 cm
231⁄2 in
60 cm

SHRUBBY GLOBULARIA
Globularia alypum
f: Plantaginaceae
Growing in dry, bushy places around the
Mediterranean, this poisonous, low,
evergreen shrub has globular heads
of sweet-scented, blue flowers.
16 in 61/2 ft 6 1/2 ft 32 in 32 in
40 cm 2m 2m 80 cm 80 cm
HEATH SPEEDWELL FOXGLOVE BEARDTONGUE SNAPDRAGON COMMON TOADFLAX
6 1/2 ft Veronica officinalis Digitalis purpurea Penstemon var “Red Garnet” Antirrhinum majus Linaria vulgaris
2m f: Plantaginaceae f: Plantaginaceae f: Plantaginaceae f: Plantaginaceae f: Plantaginaceae
A denizen of European and The heart drug digitalis is Members of this N. American Native to S.W. Europe This grayish perennial of
Asian heathlands, also introduced extracted from the dried genus, with colorful, but with many garden grassy banks around Europe
to N. America, this creeping leaves of this open-ground tubular flowers, have varieties, this perennial and W. Asia has tubular
perennial has small spikes biennial from C. and been cultivated into many has tubular, two-lipped yellow flowers with an orange
of lilac flowers. S. Europe and Morocco. garden varieties. flowers, with a yellow “throat” and long spur.
or white “throat.”
young flower
bud
231⁄2 in NEW ZEALAND LILAC 5 ft
older, open 60 cm Veronica hulkeana 1.5 m
flower f: Plantaginaceae
HEBE “RED EDGE” This evergreen shrub, also known
Veronica sp. as a hebe, is a popular garden
f: Plantaginaceae plant. It has showy stems of lilac
Often used in landscaping designs, flowers, growing wild on cliffs
this hardy hebe cultivar is thought on the eastern side of New
to be a cross between Veronica Zealand’s South Island.
albicans and Veronica pimeleoides.

85 ft/26 m
BUTTERFLY BUSH
Buddleja davidii tubular, early
f: Scrophulariaceae 10 ft/3 m spring flower
The tubular flowers of this resembles foxglove
semievergreen shrub,
originally from China, are
a rich source of nectar,
and frequently visited by
butterflies and moths.

20 ft
6m
30 in
75 cm

GREAT MULLEIN MONKEY FLOWER

Verbascum thapsus SPOTTED EMU BUSH FOXGLOVE TREE Erythranthe guttata
f: Scrophulariaceae Eremophila maculata Paulownia tomentosa f: Phrymaceae
Usually biennial, this f: Scrophulariaceae f: Paulowniaceae The tubular flowers of this
sturdy European and Widespread on seasonally flooded A deciduous Chinese tree, marshland and streamside
Asian plant of waste and land across Australia, this planted in parks for ornament, perennial from western
rough ground has a much-branched shrub has tubular Paulownia has strong-smelling, N. America have a
dense stem of tubular, yellow, orange, or red flowers, trumpet-shaped, two-lobed three-lobed lower and
five-lobed flowers. with spots inside. mauve flowers, paler inside. two-lobed upper lip.
 YELLOW RATTLE
Rhinanthus minor
f: Orobanchaceae
20 in A partial parasite whose roots tap grasses for
50 cm
nutrition, this variable, yellow-flowered annual
grows in grasslands in northern temperate regions.

3 in
8 cm

PURPLE
TOOTHWORT
Lathraea clandestina
f: Orobanchaceae
Lacking green leaves, this parasitic W. European
perennial taps the roots of willows and poplars
for food. Its flowerheads emerge directly from
underground shoots.

flowers attract
butterflies for
pollination

opposite, oval
shiny leaf

5 ft 3¼ ft 10 ft
1.5 m 1m 3m

LANTANA VERVAIN BODINIER’S

Lantana camara Verbena officinalis BEAUTYBERRY
f:Verbenaceae f:Verbenaceae Callicarpa bodinieri FLAMING GLORYBOWER
The tubular flowers of this Widely distributed f: Lamiaceae Clerodendrum splendens
prickly shrub open yellow in rough grassland in Related to American beautyberry, f: Lamiaceae
or orange, then turn red. temperate and tropical this Chinese ornamental shrub This African vine, with clusters of
Native to C. and S. America, regions, this stiffly hairy is grown in gardens for its tubular, red flowers, twines 12 ft
it is one of the most invasive perennial has slender spikes decorative purple berries, which around trees in its native forest 3.7 m
weeds in warm regions. of two-lipped, lilac flowers. are bitter but not poisonous. habitat, or trellises in gardens.

SOLANALES
The potato family (Solanaceae), an economically important family of
up to 4,000 species, dominates the order Solanales. Many species
contain poisonous alkaloids. The bindweed family (Convolvulaceae)
includes tropical climbers and low-growing herbs. The three other
families in this order are the Hydrolaceae, with one genus found
only in the Americas; five African trees in the Montiniaceae; and a
pan-tropical herb in the Sphenocleaceae.
three-
GREAT BINDWEED lobed leaf
Calystegia sylvaticus
f: Convolvulaceae
13 ft
A plant of hedgerows and 4m
waste ground, even in cities,
around the temperate
northern hemisphere, this
large-flowered perennial
spreads by far-creeping
rhizomes (underground
plant stems).
10 ft MORNING GLORY
3m Ipomoea tricolor COMMON DODDER 231⁄2 in
f: Convolvulaceae Cuscuta epithymum 60 cm
trumpet-shaped This twining herbaceous climber f: Convolvulaceae
flower with from Central America has lobed A Eurasian annual forming a dense network of
white or leaves and funnel-shaped flowers threadlike, twining stems, parasitic dodder taps
yellow center that open only in the morning. into gorse, heather, and other species for food.
 SOLANALES 5 ft/1.5 m
204
FLOWERING PLANTS • EUDICOTS

leafy
flowerhead
30 ft/10 m
glossy black
berry
DEADLY NIGHTSHADE RED ANGELS’ TRUMPETS
Atropa belladonna Brugmansia sanguinea
f: Solanaceae f: Solanaceae
Packed with powerful narcotics and truly This evergreen tree from western
deadly, this tall, perennial herb grows in woods S. America, with poisonous foliage, is grown
and scrub in Europe, N. Africa, and W. Asia. in warm, sunny gardens for its showy,
pendulous, tubular flowers.
5 ft
1.5 m
soft, succulent
berry containing
many seeds
spiny fruit funnel-shaped
capsule THORN-APPLE flower
Datura stramonium
f: Solanaceae
Rank-smelling and highly poisonous, this
widespread annual weed is thought to
originate from southern N. America.
231⁄2 in/60 cm

32 in
80 cm
HENBANE CHINESE TOMATO
Hyoscyamus niger LANTERN Solanum lypopersicum
f: Solanaceae Physalis alkekengi f: Solanaceae
Poisonous, narcotic, and f: Solanaceae Probably bred originally
evil-smelling, this annual From S. Europe despite its name, from yellow-fruited
herb inhabits disturbed this relative of Cape gooseberry cherry tomatoes from
ground, especially where is mainly grown for ornament. Peru and Ecuador, this
cattle graze, in Europe, A lantern-shaped papery case short-lived perennial is
Asia, and N. Africa. conceals its berry. 61⁄2 ft/2 m cultivated for its fleshy,
edible berries.

APIALES
35 in/90 cm
The order Apiales is dominated by the highly
distinctive and economically important palmate
carrot family (Apiaceae, often known by the leaf
6½ ft
older name Umbelliferae), with at least 3,500 2m
6½ ft
species. The Araliaceae, consisting of ivies late- 2m
summer
and ginseng, is another large family, while flowers
the parchment-bark family, Pittosporaceae,
of evergreen shrubs and trees is medium- WILD ANGELICA ASTRANTIA HOGWEED
Angelica sylvestris Astrantia major Heracleum sphondylium
sized. Four small families, each with fewer f: Apiaceae f: Apiaceae f: Apiaceae
than 20 species, complete the order. A typical “umbellifer,” this plant A plant of alpine meadows With hollow, hairy stems and
has 15–40 umbrellalike spokes and open woods in S. Europe, umbrella-shaped flowerheads
supporting its flower-bearing stem. astrantia has showy pinkish with 10–20 stalks, hogweed
Angelica grows in grassy places bracts (specialized leaves), under grows on roadsides from
in Europe and temperate Asia. its heads of small flowers. E. Europe to Britain.

16 ft/5 m

compound
umbel
finely 3¼ ft umbel of
divided leaf 8 ft lacy 1m white developing
2.5 m leaf flowers fruit stage
GIANT FENNEL HEMLOCK WILD CARROT
Ferula communis Conium maculatum Daucus carota
f: Apiaceae f: Apiaceae f: Apiaceae
This robust, giant perennial, with A biennial of wet places in Europe, The wild ancestor of cultivated
pungent-smelling, hollow stems, N. Africa, and Asia, widely carrots has just slightly swollen
grows in grassy and heathy places introduced elsewhere, hemlock taproots. It grows throughout
umbels of around the Mediterranean, has hollow, purple-spotted stems. feathery Europe, temperate Asia,
yellow flowers Asia, and N. Africa. All parts are poisonous. leaf and N. Africa.
 10 ft
3m

4 ft
1.2 m
CHILI PEPPER TOBACCO
Capsicum frutescens Nicotiana tabacum
f: Solanaceae f: Solanaceae
Native to Brazil and Bolivia, Originally S. American, this
this small shrub is much herb is the most common
cultivated in tropical Asia of two species grown for mostly spineless
and equatorial America. smoking tobacco. The upper leaf
Its fruits are chilies. leaves contain nicotine.

berrylike
flowers red fruit
3¼ ft and pinnate
1m leaves

POTATO
Solanum tuberosum
f: Solanaceae
Potatoes are a major food crop. The swollen tubers edible
turn green (and poisonous) in sunlight. The plant tubers
was bred from ancestors in the S. American Andes.

33 ft
10 m

egg-shaped
flowerhead

KOHUHU
Pittosporum tenuifolium
f: Pittosporaceae
Growing wild in forests and
scrub on both islands of COMMON HOLLY
New Zealand, this evergreen Ilex aquifolium
tree has almost black f: Aquifoliaceae
branches and honey-scented An evergreen,
spring flowers. woodland understory
shrub or small tree
231⁄2 in/60 cm from Europe, N. Africa,
leathery, and N.W. Asia, holly
spiny leaf 32 in/80 cm 80 ft/24 m develops spiny leaves
to discourage grazers.

SEA HOLLY
Eryngium maritimum
f: Apiaceae
AQU I FOLI A L ES
Leathery leaves help this sturdy perennial The mainly tropical holly family
reduce water loss and repel salt spray (Aquifoliaceae) includes trees and shrubs
in sand dunes around Europe,
N. Africa, and S.W. Asia. with characteristically toothed leaves,
and is the main representative of the
ORIENTAL Aquifoliales order. Other members
GINSENG
IVY Panax ginseng of this small order are an odd family of
Hedera helix f: Araliaceae twining herbs, the Cardiopteridaceae;
f: Araliaceae The root of this Asian herb is used three Asian shrubs in the Helwingiaceae;
An evergreen shrub from as a traditional remedy. Its species four South American shrubs and trees
Europe and S.W. Asia, ivy name, Panax, comes from the
100 ft climbs up other plants or forms Greek word panacea, meaning in the Phyllonomaceae; and a family
30 m creeping carpets in woods. a cure-all for diseases. of tropical trees, the Stemonuraceae.
 ASTERALES ray florets
206 Eleven families make up the Asterales. The
largest is the daisy family (Asteraceae) with some
25,000 species. Typically, each flowerlike head
FLOWERING PLANTS • EUDICOTS

(capitula) has many individual flowers, called

florets, surrounded by showy rays. Some of
the bellflower family (Campanulaceae) show
similar characteristics. This order also includes disk florets
the bogbean (Menyanthaceae) and fan-flower
(Goodeniaceae) families, and seven smaller families.

bract

5 ft SALSIFY 4 ft
1.5 m Tragopogon porrifolius 1.25 m
f: Asteraceae
MICHAELMAS DAISY A biennial herb of grassy places
Symphotrichum novi-belgii around the Mediterranean,
f: Asteraceae salsify has lilac or reddish-purple
Previously classified as a species of Aster, this flower heads, surrounded by
showy, variable garden plant is a native of longer, pointed bracts.
eastern N. America.

COMMON
RAGWORT
Jacobaea vulgaris
3¼ ft
1m
f: Asteraceae 5 ft
1.5 m
Toxic to farm animals and
capitula of disk avoided by rabbits, this
florets which native perennial from Europe
mature into seeds lobed ray florets, and W. Asia has invaded
usually yellow grasslands almost worldwide.
or orange

serrated
leaf
11 ft
3.5 m oval, toothed
leaf
OXEYE DAISY
Leucanthemum vulgare
SUNFLOWER CORNFLOWER f: Asteraceae
Helianthus annuus Centaurea cyanus One of the most common
f: Asteraceae f: Asteraceae small, white white-flowered daisies of Europe
Probably Mexican in origin, this tall Probably native to S. Europe and flower head and W. Asia, this variable
annual is grown as an ornamental as W. Asia, cornflower seeds were inseparable patch-forming perennial
well as commercially for its seeds that from corn seeds, making it a pestilential 30 in quickly seeds into
contain 27–40 percent polyunsaturated agricultural weed before herbicides. 75 cm disturbed ground.
oil and 13–20 percent protein.

upright or
horizontal
5 in leaf
12 cm

20 in
50 cm pink or
yellow purple ray
COMMON flower head florets 4 ft
DAISY DANDELION 1.2 m
Bellis perennis Taraxacum officinale
f: Asteraceae f: Asteraceae PURPLE CONEFLOWER
Originally from Europe The name for this familiar Echinacea purpurea rough,
and W. Asia, this short weed covers an amazing f: Asteraceae toothed
perennial of grazed diversity of minutely Named after the cone shape of its leaf
tall, robust stem grasslands and lawns has differing forms, with central disk florets, this
been transported almost perhaps 1,000 recognized ornamental perennial from eastern
worldwide. It is commonly “microspecies” in Europe N. America is cultivated for herbal
considered a weed. and Asia. remedies against colds and flu.
 8 ft MILK THISTLE
2.5 m Silybum marianum
f: Asteraceae
4 ft
1.2 m 207
This weakly spiny biennial, 6 1/2 ft
with white-blotched leaves, 2m
grows in waste and cultivated
ground around S. Europe, GLOBE THISTLE
N. Africa, and W. Asia. 8 ft Echinops bannaticus
2.5 m f: Asteraceae
CANADIAN GOLDENROD Native to grassy places
Solidago canadensis in S.E. Europe, this hairy- spiny
f: Asteraceae stemmed perennial has a head
Native to N. America but grown spherical head of bluish, CARDOON
in many gardens, this downy tubular “disk” florets. Cynara cardunculus deeply cut
perennial has yellow f: Asteraceae leaf
flower heads borne in curved, The fleshy flower base, eaten as globe
spreading flower spikes. artichoke, comes from the scolymus
variety of cardoon, a wasteground
6 1/2 ft perennial from the E. Mediterranean.
2m
20 in 6 ft
50 cm 1.8 m

COTTONWEED DENSE BLAZING SPEAR THISTLE

Otanthus maritimus STAR Cirsium vulgare
f: Asteraceae Liatris spicata f: Asteraceae
This shrubby perennial f: Asteraceae 5ft Common in disturbed
grows in coastal A plant of damp 1.5m ground in Europe and
habitats in ground in eastern N. W. Asia, this robust
S. Europe, N. Africa, America, this plant is perennial with
BUTTERBUR and S.W. Asia. The named after its dense, spine-tipped
Petasites hybridus common name derives elongated, spike leaves has been
f: Asteraceae from its woolly of rose-purple carried worldwide
early Plants of butterbur have stems with stems and leaves. flower heads. by agriculture.
spring either male or female flowers. It grows
flower in wet meadows and beside streams
throughout Europe to Iran.

61/2 ft/2 m

FRENCH
TARRAGON
Artemisia
dracunculus
f: Asteraceae
Leaves of this bushy,
aromatic herb from
S.E. Europe, Asia,
20 in 23 1/2 in 4 ft and N. America are
50 cm 60 cm 1.2 m used to flavor fish
outer ray and other foods.
florets blue or
YARROW lavender
Achillea millefolium flowers oblong
f: Asteraceae rosette
Found in grassy places
leaf
throughout Europe, W. Asia,
and N. America, this CHICORY COTTON LAVENDER
feathery-leaved perennial has Cichorium intybus Santolina chamaecyparissus
been introduced to Australia f: Asteraceae f: Asteraceae
and New Zealand. Introduced almost worldwide Widely grown in gardens,
from Europe, W. Asia and this strongly aromatic, dwarf,
N. Africa, chicory is cultivated evergreen shrub with
leaf is hairy for salad leaves and for a coffee gray, hairy leaves is native to
on both sides substitute from its roots. rocky places around the 23 1/2 in
W. Mediterranean. 60 cm

8 in/20 cm

20 in
50 cm
GARDEN MARIGOLD STRAND GAZANIA
Calendula officinalis GARDEN GAZANIA Gazania rigens
f: Asteraceae Gazania sp. F: Asteraceae
This species has been cultivated f: Asteraceae This sprawling, mat-forming
so long that its origins are Seventeen species of Gazania grow in perennial often flowers
unknown. Marigold extract southern Africa, most in dry habitats. prolifically on dunes and rocky
from the flowers is used to Cultivars like these can survive in outcrops along the southern
treat skin problems. gardens with little watering. Cape coast of South Africa.
 ASTERALES
208
FLOWERING PLANTS • EUDICOTS

4 ft
1.2 m 28 in/70 cm

perennial flower 16 in bell-shaped oval 5 ft

leaf spike 40 cm flower leaf 1.5 m

GREAT BLUE LOBELIA tubular flower ROYAL BLUEBELL BALLOON FLOWER BUCKBEAN
Lobelia siphilitica Wahlenbergia gloriosa Platycodon grandiflorus Menyanthes trifoliata
f: Campanulaceae f: Campanulaceae f: Campanulaceae f: Menyanthaceae
A showy N. American perennial of The state plant of the Australian The only member of its genus, This far-creeping perennial
woods and meadows from Manitoba Capital Territory, this erect this Asian perennial with produces its beanlike fruits in
to Alabama, this plant was once perennial with deep blue flowers blue or white, bell-shaped bogs and shallow water in
supposed to cure syphilis. grows in mountain grasslands flowers also comes in many N. America, Greenland,
in southern Australia. garden varieties. N. Europe, and Asia.

6 in 5 ft
15 cm 1.5 m

20 in 3¼ ft
50 cm 1m

clustered NETTLE-LEAVED fleshy

flower head BELLFLOWER one-sided leaf
flower
ROUND-HEADED Campanula trachelium SWAMPWEED FRINGED WATER LILY
RAMPION f: Campanulaceae Selliera radicans Nymphoides peltata
Phyteuma orbiculare Found in hedgebanks f: Goodeniaceae f: Menyanthaceae
f: Campanulaceae around Europe, Iran, and This creeping herb is found on Unlike true water lilies, this
With dark blue flowers in rounded N. Africa, this roughly coastal sandflats around Chile, aquatic perennial has smaller
heads, this unbranched perennial hairy perennial has Australia, and New Zealand— yellow flowers with five fringed
grows in chalk grassland from nettlelike leaves and where it also grows alongside petals. It is found from
lanceolate S. England to Greece. mountain streams. Europe to Japan.
leaf blue, bell-shaped flowers.

DIPSACA L ES MUSKROOT
Adoxa moschatellina
The order Dipsacales is found worldwide, but f: Adoxaceae
Inflorescences of this
especially in the northern hemisphere. Its members 6 in delicate woodland 32 in
generally have small flowers, often in compact 15 cm 80 cm
perennial from Europe,
heads, and include many popular ornamental plants. Asia, and N. America
It consists of two families. The moschatel and have five flowers at RED VALERIAN
right angles (with one Centranthus ruber
viburnum family (Adoxaceae) contains about 200 facing upward). f: Caprifoliaceae
species, and the honeysuckle family (Caprifoliaceae) Butterflies appreciate
has some 860 species, including valerians and teasels elder flower head
the flowers of this
that were previously placed in their own families. grayish perennial. It is
found on coastal rocks
and old walls around
the Mediterranean,
lobed leaf and is naturalized
elsewhere.

6½ ft
2m
40 ft/12 m

elder ELDERBERRY
berries Sambucus nigra
f: Adoxaceae
A hedgerow shrub or small tree with
lobed leaves and flat-topped flower heads,
shiny, red elder grows throughout Europe,
berry W. Asia, and N. Africa.

EUROPEAN
CRANBERRYBUSH
Viburnum opulus GARDEN VALERIAN
f: Adoxaceae Valeriana officinalis
Native across Europe to Asia, this f: Caprifoliaceae
deciduous hedgerow shrub has This perennial is found in grassy places
flattened flower heads with large, from N. Europe into Asia. It has flattened
13 ft sterile outer flowers, around heads of pale pink, five-petaled flowers, scented pinnate
4m smaller, fertile ones. which are darker when in bud. flowers leaf
 sweet-scented
flower

flower
stem

oval, pointed
leaf
20 ft
ripe 6m
berries

EUROPEAN
HONEYSUCKLE
Lonicera periclymenum
f: Caprifoliaceae
A deciduous shrub of woods large, red bracts
and hedgerows in Europe and surround and
N. Africa, honeysuckle protect flowers
scrambles over other plants to
reach sunlight before flowering.

HIMALAYAN
6½ ft/2 m HONEYSUCKLE
Leycesteria formosa
f: Caprifoliaceae
funnel-shaped Originally from the Himalayas,
flowers attract bees this deciduous shrub has
for pollination arching stems and crowded
heads of white flowers in
reddish-purple bracts.

26 ft
8m 10 ft/3 m 13 ft/4 m
spiny, oblong
CORAL HONEYSUCKLE SNOWBERRY WEIGELA flowerhead
Lonicera sempervirens Symphoricarpos albus Weigela florida
f: Caprifoliaceae f: Caprifoliaceae f: Caprifoliaceae center of flower
This slender vine scrambles high Native from Alaska to Colorado, Originally from China and Korea, opens first
up forest trees in the E. US. Its but widely planted elsewhere, this this deciduous shrub has clusters
trumpet-shaped red flowers, deciduous, suckering shrub of funnel-shaped flowers that are
with yellow interiors, attract has dense heads of pink flowers, deep rose in color, sometimes
hummingbirds for pollination. ripening to white berries. with a yellow throat.

pincushionlike 6½ ft
flowers 2m

fruiting
head

flower head stalkless

stem leaf
prickly bracts
serrated form cup
pinnate leaf at base
leaf TEASEL
30 in 231⁄2 in 3¼ ft Dipsacus fullonum
75 cm 60 cm
upright, 1m f: Caprifoliaceae
oval leaf This stout biennial of
FIELD SCABIOUS CARMEL DAISY DEVIL’S-BIT SCABIOUS prickly rough ground in
Knautia arvensis Scabiosa prolifera Succisa pratensis flower stem Europe, W. Asia, and
f: Caprifoliaceae f: Caprifoliaceae f: Caprifoliaceae N. Africa has prickly
Flower heads of this hairy perennial The pale yellow flower heads of this The devil supposedly bit off the stumpy stems and leaves, and
have larger outer petals than inner robust E. Mediterranean annual have rootstock of this downy perennial. spiny flower bracts.
ones. It grows in dry pastures large outer petals with smaller inner It grows in damp soils across Seeds are a winter food
from Europe to Siberia. florets. It grows around fields. Europe and N. Africa. source for birds.
fungi
Ranging from mushrooms to microscopic molds, fungi were once
classified as plants. Today, they are recognized as a distinct kingdom of
living things. Growing in or through their food, they digest organic
matter, and often become visible only when they reproduce. Fungi
are both allies and enemies to other forms of life: vital recyclers and
mutually benefiting partners, they also include parasites and pathogens.

212 238 244

MUSHROOMS SAC FUNGI LICHENS
As well as mushrooms, These fungi produce their Living partnerships
this large group includes spores in microscopic between fungi and algae,
bracket fungi, puffballs, sacs, which often form lichens colonize all kinds
and many other species. a felty layer on their fruit of bare surfaces. Some
Their fruit bodies vary in bodies. Many are cup- form flat crusts, while
shape, but all make their shaped, but they also others are like tiny plants.
spores on microscopic include truffles, morels, Most grow slowly, and are
cells called basidia. and single-celled yeasts. exceptionally long-lived.
212 M U S H RO O M S
The phylum Basidiomycota includes the majority of what are
FUNGI • MUSHROOMS

commonly called mushrooms and toadstools. Found in almost

every major habitat type, nearly all share the ability to form
sexual spores externally on special cells called basidia.

PHYLUM BASIDIOMYCOTA Few fungi are as diverse in form as the Basidiomycota.

CLASSES 16 Their fruit bodies (the structures in which spores are
ORDERS 52 produced) have evolved to provide varied and efficient
FAMILIES 177
SPECIES About 32,000
mechanisms for spore dispersal. These fruit bodies
often take the form of a stem, cap, and gills, but
many are simple, crustlike sheets, and others are
more complex, bracketlike forms. More exotic still
are the totally enclosed, spherical types such as the
puffballs and the earthstars. Then there are the
beautiful, corallike structures of the club and coral
fungi, and the often weird, animallike shapes of the
stinkhorns. Each structure supports and produces
special spore-producing cells called basidia, and—
These typical mushrooms show the depending on the fungus concerned—uses a variety
cap with radiating gills on the underside, of mechanisms to disperse these spores. For example,
supported by the central stem.
gilled fungi forcibly eject their spores, to be carried
off by air currents. The puffballs, however, depend on
a combination of wind and raindrops to puff out their
spores. Stinkhorns, on the other hand, use foul odors
and bright colors to attract animals, such as insects
and other invertebrates, to eat their spore mass and
to pass the spores unharmed through their digestive
tracts. In this way the spores are scattered as far as
The microscopic spores of the field the animals travel before excreting them.
mushroom are shown here still attached
to the basidia, or reproductive cells.
fu nga l feedi ng
The main body of the fungus is usually underground,
formed of fine fungal threads called hyphae, which
make up a mycelium (fungal body). These threads
permeate the substance on which the fungus grows, be
it soil, old leaf litter, fallen trees, living plant tissues, or
even dead and decaying animals. The mycelium may
spread over a large area, and in many fungi it also forms
This stinkhorn is covered with a dark
spore mass, which rapidly liquifies and a mutually beneficial relationship with the roots of
emits a foul odor, usually of rotting flesh. plants. In this relationship—called a mycorrhiza—
the mycelium wraps around and penetrates the plant’s
D E B AT E roots, providing the fungus with access to carbohydrates
COLORFUL MYSTERY from the plant. In return, the plant benefits from the
mycelium’s greater ability to absorb water and mineral
Many mushrooms have bright colors:
red, violet, blue, and green. Scientists
nutrients. Other mushrooms break down dead organic
have little idea what advantage these matter, or devour living organisms for food—both
colors give to the mushroom. Unlike processes help to renew the soil and to improve the
flowers, they do not need bright colors growing conditions for other living things.
to attract pollinators. Red and orange
pigments possibly prevent damage
from sunlight, while others may act MUSHROOMS AMONG FALLEN LEAVES
as a warning to predators. Many fungi feed on dead and decaying plant tissues and may
spread in huge numbers where nutrients and moisture occur.
 AGA R ICA L ES FIELD MUSHROOM
Agaricus campestris
214 Many of the most familiar mushrooms and toadstools belong to this order. f: Agaricaceae
Common in meadows of
They include fungi with fleshy, not woody, fruitbodies (the body supporting N. America and Eurasia, this
the spore-forming cells). Many have caps and stems with gills; some also 1½–4 in fungus has a rounded cap;
4–10 cm
MUSHROOMS • AGARICALES

have pores. Other forms include bird’s-nest, brackets, crusts, trufflelike, bright pink gills turning
and puffballs. Most live on leaf litter, soil, or wood; others are parasitic or brown with maturity; and a
short stem with a small ring. 2–4 in
live in association with plant roots (mycorrhizal). 5–10 cm

CULTIVATED MUSHROOM
Agaricus bisporus
slightly THE PRINCE f: Agaricaceae
fibrous Agaricus augustus Selling in the millions, this familiar
surface f: Agaricaceae fungus is common worldwide.
A large, orange-brown, Its cap varies from white to dark
scaly cap and a white, brown with scales.
woolly stem with a floppy
ring characterize this
common species 2–6 in
from Eurasia and 5–15 cm
N. America. 3¼–6 in
8–15 cm FRECKLED
DAPPERLING
Echinoderma asperum
f: Agaricaceae
From woods and
gardens in Eurasia and
N. America, this uncommon
2–43/4 in YELLOW STAINER fungus’s brown cap has
5–12 cm Agaricus xanthodermus pyramidal scales that rub off.
f: Agaricaceae The brown stem has a ring.
This species has a flattened
cap center, chrome-yellow 23/4–6 in
stains on the stem base, 7–15 cm HORSE MUSHROOM
and an unpleasant odor. It Agaricus arvensis
is common in Eurasia and f: Agaricaceae
western N. America. Common in parks in ⅜–1½ in
N. America and Eurasia, 1–4 cm
this species stains dull
brassy yellow. The pendent STINKING
ring on its stem has a DAPPERLING
cogwheellike pattern Lepiota cristata
mature
cap
3/4–2¼ in on its underside. f: Agaricaceae
2–6 cm A rubberlike smell is a
yellow stains characteristic feature
1¼–3¼ in of this species of Lepiota
3–8 cm (most common in woods
and meadows) from
young N. America and Eurasia.
cap
1½–4½ in
4–11 cm
ring
ORANGE-GIRDLED
BLUSHING LAWYER’S WIG DAPPERLING
DAPPERLING Coprinus comatus Lepiota ignivolvata
Leucoagaricus badhamii f: Agaricaceae f: Agaricaceae
f: Agaricaceae This fungus is common on Found in Eurasia, this
Found in Eurasia, this disturbed soils and pathsides in uncommon species has
rare fungus grows in woods 2–3¼ in N. America and Eurasia. Its tall, an orange-margined ring
and gardens with rich soils. 5–8 cm shaggy cap is unmistakable, and set low down on the
White when young, it dissolves into inky fluid. club-shaped, white stem.
bruises blood-red when
handled, and then blackens. WHITE DAPPERLING
Leucoagaricus leucothites
f: Agaricaceae 2–6 in
Common in meadows and grassy 5–15 cm
roadsides in Eurasia and 8–20 in
20–50 cm
N. America, this fungus’s ivory SHAGGY PARASOL
fruitbody turns grey-brown Chlorophyllum rhacodes
with age. Its gills are white f: Agaricaceae
to pale pink. Common in Eurasia and
⅜–1¼ in N. America, this fungus
1–3 cm has a shaggy, brown cap;
thick, double-edged ring;
⅜–2 in reddening flesh; and
1–5 cm inner swollen stem base.
layer
PLANTPOT DAPPERLING GRAY PUFFBALL GIANT PUFFBALL
Leucocoprinus birnbaumii Bovista plumbea Calvatia gigantea
f: Agaricaceae f: Agaricaceae f: Agaricaceae bulbous
Common worldwide, this This ball-shaped fungus is Common in hedgerows, fields, and stem base
species grows on soils of potted smooth when young. As it gardens in N. America and Eurasia,
plants. Its features include a matures, its skin flakes to reveal a this mushroom’s large, smooth,
delicate golden-yellow cap and papery inner layer. It is common white fruit body with a white to
a slender stem with a ring. outer skin in N. America and Eurasia. yellow interior is unmistakable.
 215

¾–1¾ in

MUSHROOMS • AGARICALES
2–4.5 cm

MEADOW PUFFBALL
Lycoperdon pratense ¾–1½ in
f: Agaricaceae 2–4 cm
This short-stemmed species has an
internal membrane separating the
spore-mass from the stem. It is common
in meadows throughout Eurasia.

2–4 in
5–10 cm
COMMON PUFFBALL
Lycoperdon perlatum
f: Agaricaceae
The most common white
⅜–1¼ in puffball, this species is found
1–3 cm in N. America and Eurasia.
contrasting Its granular spines rub off
brown scales 4–12 in PARASOL to leave circular scars in
10–30 cm Macrolepiota procera a regular pattern.
f: Agaricaceae
Common in meadows in
N. America and Eurasia,
this tall species has a
scaly cap and snakeskin-
patterned stem with
a thick ring. SPINY PUFFBALL PESTLE PUFFBALL
Lycoperdon echinatum Lycoperdon excipuliforme
f: Agaricaceae f: Agaricaceae
This species is found in One of the tallest puffballs,
snakeskin beech woods in Eurasia and this fungus has a buff-white
pattern on N. America. It has groups of long stem that is left behind once
the stem spines that meet at their tips. Its the spores are released.
spores are purplish brown. It is found in Eurasia.

⅜–1 in
1–2.5 cm ¾–2 in
2–5 cm
STUMP PUFFBALL
Apioperdon pyriforme
f: Agaricaceae
Common in N. America and Eurasia,
this pear-shaped species grows on
wood. It has prominent cords at its
base, and is firm when young. 2¼–6 in ⅜–1½ in 1¼–2¾ in STINKING PUFFBALL
6–15 cm 1–3.5 cm 3–7 cm Lycoperdon foetidum
f: Agaricaceae
Common in acidic heaths
and woods in Eurasia, this
yellowish brown puffball has
dark brown spines. Its flesh
has an unpleasant smell.

POPLAR FIELDCAP COMMON FIELDCAP SPRING FIELDCAP

Cyclocybe cylindracea Agrocybe pediades Agrocybe praecox
f: Bolbitiaceae f: Bolbitiaceae f: Bolbitiaceae
Found in Eurasia, Usually found in lawns Common in Eurasia and
this rare species grows in Eurasia, this fungus’s N. America during spring,
on poplar wood. The cap smooth, yellowish cap this fungus’s cap may have
bulbous base cracks when dry and and slender stem have a marginal veil fragments. The
the stem has a ring. mealy smell and no veil. stem may have a fragile ring.

2¼–4 in 1¼–2 in ⅜–½ in ⅜–1¼ in

6–10 cm 3–5 cm 1–1.5 cm 1–3 cm

WINTER STALKBALL SANDY STILTBALL MILKY CONECAP YELLOW FIELDCAP

Tulostoma brumale Battarreoides digueti Conocybe apala Bolbitius titubans
f: Agaricaceae f: Tulostomataceae f: Bolbitiaceae f: Bolbitiaceae
Typically found in sandy soils Found in very dry, sandy This species is frequently Growing in meadows, this
in dune slacks in Eurasia and soils in N. America, this tall, found in lawns in Eurasia and fungus is common in
N. America, this fungus has tiny, stalked species emerges from N. America. Its conical, ivory N. America and Eurasia.
whitish yellow, round heads on a leathery “egg.” Its brown cap with orange-brown gills It has a delicate, sticky cap
slender, pale brown stalks. cap contains the spores. sits on a tall stem. that lasts only a day or so.
 AGA RICA LES
216 buff warts

2–4 in 1¼–3¼ in
MUSHROOMS • AGARICALES

5–10 cm 3–8 cm

3¼–8 in 3¼–6 in FALSE DEATHCAP TAWNY GRISETTE

8–20 cm 8–15 cm Amanita citrina Amanita fulva
f: Amanitaceae f: Amanitaceae
Found in Eurasia and Common in woods
eastern N. America, this in Eurasia and
species has a stem with a N. America, this fungus
volva distinctly rimmed, may have veil patches.
rounded bulb, and flesh The stem has a white 2¼–7 in
with a potato odor. Its cap volva (baglike remains of 6–18 cm
is white or pale yellow. an enclosing veil).
CAESAR’S MUSHROOM DEATH CAP
Amanita caesarea Amanita phalloides
f: Amanitaceae f: Amanitaceae BLUSHER
Found under oaks, mainly Common in Eurasia and parts
Amanita rubescens
from the Mediterranean to of N. America, this species has
f: Amanitaceae
C. Eurasia, this mushroom a fragile ring, large volva, and This species is
has an orange cap and stem, sickly smell. Its cap may be common in mixed
and a white volva. greenish, yellowish, or white. woods in Eurasia and
N. America. Its cap
varies from cream
to brown, and its flesh
pure white warts bruises pinkish red.
2–43/4 in
5–12 cm
rounded bulb
FLY AGARIC DESTROYING ANGEL
2¼–6 in Amanita muscaria 2¼–4¼ in Amanita virosa PANTHER CAP
6–15 cm f: Amanitaceae 6–11 cm f: Amanitaceae Amanita pantherina
Common in Eurasia and Found throughout f: Amanitaceae
N. America, this distinctive N. Eurasia, this rare fungus This species has pure white warts
mushroom is especially is pure white with a slightly (remains of its protective veil), a ring
common under birches. sticky, bell-shaped cap on its stem, and a ridged bulb. It is
Its white warts may and a white volva. found in Eurasia and N. America.
wash off after rain.
1¼–4 in
3–10 cm
1½–3¼ in
YELLOW CLUB 4–8 cm
Clavulinopsis helvola club often
f: Clavariaceae flattened
Common in grassland
in Eurasia and N. America,
2–6 in this is one of several yellow 3/4–1½ in
5–15 cm clubs. It can only be 2–4 cm
reliably identified by
its spiny spores.
GOLDEN
SPINDLES
Clavulinopsis corniculata
f: Clavariaceae
This golden-yellow club
with antlerlike branches is VIOLET CORAL WHITE SPINDLES
relatively common in acidic, Clavaria zollingeri Clavaria fragilis
unimproved meadows f: Clavariaceae f: Clavariaceae
and grassy clearings in A rare species found in mossy This fungus forms dense
woods in Eurasia. meadows and woodlands in Eurasia clusters of pure white,
and N. America, this species has simple clubs in woods
violet, corallike branches. and meadows in Eurasia.

⅜–1 in 1½–3¼ in 1¼–3½ in

1–2.5 cm 4–8 cm 3–9 cm
1½–43/4 in ⅜–1¼ in
4–12 cm 1–3 cm
BLUE EDGE LILAC PINKGILL
WOOD PINKGILL MOUSEPEE PINKGILL PINKGILL Entoloma
Entoloma rhodopolium Entoloma incanum Entoloma serrulatum porphyrophaeum THE MILLER
f: Entolomataceae f: Entolomataceae f: Entolomataceae f: Entolomataceae Clitopilus prunulus
Found in Eurasia, this Smelling of mice, this pinkgill This blue-black capped Occasionally found in f: Entolomataceae
pale gray to gray-brown has a hollow stem and a bright fungus has serrated meadows, this fungus Common in mixed woods in
species has a cap with grass-green cap that fades pinkish gills with dark grows in Eurasia. It Eurasia and N. America, this
a domed center and to brown with age. It grows edges. It is found in has a gray-purple fungus has a cap that changes
slender stem. It may in unimproved meadows meadows and parks in fibrous cap and stem, shape with age, starting convex
smell of bleach. in Eurasia. Eurasia and N. America. and pinkish gills. then becoming depressed.
 217
contrasting
cream gills
2–8 in 2–8 in 3¼–8 in 1¼–2¼ in

MUSHROOMS • AGARICALES
5–20 cm 5–20 cm 8–20 cm 3–6 cm

BLUE LEGS CLOUDED FUNNEL ANISEED FUNNEL

Lepista personata Clitocybe nebularis Clitocybe odora
f: Tricholomataceae f: Tricholomataceae f: Tricholomataceae
WOOD BLEWIT The blue legs has a buff cap This fleshy mushroom has Common in Eurasia and
Lepista nuda and a bright blue-lilac stem with crowded gills that often N. America, this fungus
f: Tricholomataceae perfumed flesh. It is common run down the stem. It is unmistakable due to
Common in mixed woods in in open meadows in Eurasia. frequently grows in its strong aniseed odor.
Eurasia and N. America, this large circles in Eurasia Its sea-green color fades
species has a violet cap that and N. America. as it matures
fades to brown, but the stem to grayish green.
and gills remain violet.

3/4–2¼ in
2–6 cm

IVORY FUNNEL
Clitocybe dealbata
f: Tricholomataceae
This species grows in rings in
lawns and meadows in Eurasia
and N. America. It has a
frosted appearance and
slightly decurrent gills.

3/4–3¼ in
2–8 cm

decurrent gills
(running down stem)

SULPHUR KNIGHT
Tricholoma sulphureum
f: Tricholomataceae
Characterized by its bright
sulphur-yellow color and
repulsive smell of coal gas,
43/4–16 in this fungus is common in
inrolled cap (curls 12–40 cm mixed woods in Eurasia
inward at margins) and parts of N. America.
2–43/4 in 1½–4 in
short stem 5–12 cm 4–10 cm
2–10 in/5–25 cm
GIANT FUNNEL
Leucopaxillus giganteus DINGY AGARIC SOAPY KNIGHT PIPE CLUB
f: Tricholomataceae Tricholoma portentosum Tricholoma saponaceum Typhula fistulosa
Often found in huge fairy f: Tricholomataceae f: Tricholomataceae f: Typhulaceae
rings in Eurasia and N. America, Usually found with conifers in northern Variable in color—gray-brown, greenish Often found in large groups in Eurasia
this giant fungus is white, with N. America and N. Eurasia, this species or pinkish gray to mottled—with a soapy and parts of N. America, this species
decurrent gills, an inrolled cap has a smooth, bell-shaped cap and a smell, this species is common in mixed has tall, slender clubs attached to
margin, and a short stem. stout, whitish stem that flushes yellowish. woods in Eurasia and N. America. fallen branches and hardwood litter.
218

F LY AG A R I C
A man it a mu scar ia
The fly agaric is arguably the most famous of all fungi, and it
appears in illustrations in children’s books all over the world.
The bright scarlet cap, usually scattered with white spots of
veil tissue, make it one of the easiest fungi to identify. Found
originally throughout most of Europe, northern Asia, and North
America, it now grows wherever humans have planted the host
trees—usually birch—with which the fungus forms a mutually
beneficial relationship. It is now commonly found in parts of
Africa, India, and Australasia. All parts of the fly agaric are
poisonous, although rarely fatal.

large floppy ring

is easily torn

size Cap diameter 2¼–6 in (6–15 cm)

habitat Birch and pine woods
distribution Almost worldwide
spore color White
 white warts loosely SLICE THROUGH CAP
yellow flesh under attached to cap In cross section, the yellowish orange flesh immediately
outer skin beneath the red skin of the cap is revealed. The gills, or
219
lamellae, are bright white, in contrast.
gills are
pure white

stem flesh

SCALES
The white or pale yellow crowded gills
warty scales are all that
remain of an enveloping
veil of tissue that once
enclosed the young cap.

red skin peels

off easily

FAMILIAR FUNGUS
The fly agaric changes shape
dramatically as it grows and
expands, but the color and key
characters usually remain constant. STEM BASE
The cap may fade to yellow if The base is swollen and GILLS
there has been persistent rain. Its clublike, with warty ridges The fly agaric’s radiating gills are of
common name refers to an old folk encircling the upper part of different lengths, some stopping well
remedy—the red skin of the cap the bulb. Threads of mycelium short of the stem. This allows them
was used in a saucer of milk to (fungal tissue) connect to fine to fill the available space, ensuring
poison annoying houseflies. tree roots below the soil. that spore production is maximized.

GROWTH STAGES OF A FLY AGARIC

warts
dispersed
veil warts ring now
partial veil veil formed fully
beneath cap starting exposed upturned
to tear gills cap

BABY BUTTON BROKEN VEIL GROWING CAP GILLS APPEAR SPORE DISPERSAL OLD AGE
The young button is The stem has started to The cap expands, with The partial veil starts to Spores grow on the At the end of its life the
completely enclosed in grow, and the universal veil the partial veil still tear, forming the ring and exposed gills and are cap coloration fades and
a warty universal veil. on the cap has split apart. attached underneath. exposing the gills above. then released into the air. the cap may turn upward.
 AGA RICA LES
220
1¼–2¼ in 2¼–4 in 2–4 in
3–6 cm 6–10 cm 5–10 cm
MUSHROOMS • AGARICALES

THE GYPSY DAPPLED WEBCAP

2–43/4 in Cortinarius caperatus Cortinarius bolaris 2–3¼ in BLUE AND BUFF WEBCAP
5–12 cm f: Cortinariaceae f: Cortinariaceae 5–8 cm
Cortinarius malachius
From N. Eurasia and N. America, Frequent in beech woods in f: Cortinariaceae
this species, found in conifer Eurasia, this fungus has tiny Found under pines in N. Eurasia,
woods, has a frosted cap with reddish-brown scales on its this uncommon species has a pale
a white veil, and a stem with a cap and stem. The flesh stains brown cap with a lilac tint, and
distinct sheathing ring. yellow-orange in the stem. a stem with bands of white veil.

ELEGANT WEBCAP
Cortinarius elegantissimus
f: Cortinariaceae
Found throughout Eurasia in fine scales and
raised point at beech woods on chalky soil, this fibers cover
cap center fungus has a vivid orange-yellow cap surface
cap and a yellow stem with
a large, marginate bulb.

PEARLY WEBCAP
1¼–3¼ in Cortinarius alboviolaceus
3–8 cm
f: Cortinariaceae
SCALY WEBCAP This species is common in mixed woods
Cortinarius pholideus in Eurasia and N. America. Its silvery-
f: Cortinariaceae white fruitbody has a lilac tint. The
This uncommon species is found gills mature to cinnamon-brown.
under birches in Eurasia. It is scaly ring
easily recognized by its scaly,
brown cap and stem, and violet
gills and stem apex when young.

2¼–4 in
6–10 cm

2¼–4 in
6–10 cm

2–43/4 in
RED AND OLIVE WEBCAP ORANGE WEBCAP 5–12 cm
Cortinarius rufoolivaceus Cortinarius mucosus minute
f: Cortinariaceae f: Cortinariaceae scales
on cap RED BANDED WEBCAP
Found in beech woods throughout Restricted to N. Eurasia and Cortinarius armillatus
Eurasia, this uncommon species N. America, this species is f: Cortinariaceae
has a distinctively copper-colored common in pine woods. Its
cap—with a pink or olive-green orange-brown cap and stout, The cinnabar-red bands of
margin. The bulbous stem is white stem are very slimy. veil on the club-shaped stem
violet to greenish yellow. The gills are rust-brown. distinguish this common species
of birch woods in Eurasia
2¼–6 in and N. America.
3/8–1¼ in 1½–4 in ring zone 6–15 cm
1–3 cm 4–10 cm club-shaped stem
BITTER VIOLET WEBCAP
BIGFOOT WEBCAP Cortinarius violaceus
Cortinarius sodagnitus f: Cortinariaceae
f: Cortinariaceae Found in mixed woodland
1¼–23/4 in Found mainly in S. England throughout Eurasia and
3–7 cm and Mediterranean Eurasia, N. America, this rare species
this uncommon species has a distinctive, intensely
occurs in beech woods violet-colored cap and stem.
on chalky soil. The bulb
on the stem is large. 3¼–6 in
8–15 cm
club-shaped
stem BIRCH WEBCAP
SPLENDID WEBCAP PIXIE WEBCAP Cortinarius triumphans
Cortinarius splendens Cortinarius flexipes f: Cortinariaceae
f: Cortinariaceae f: Cortinariaceae Common under birches
This rare species has a golden-yellow An odor of pelargoniums and delicate in Eurasia, this fungus is
cap, yellow flesh, a sulphur-yellow white hairs on the pointed cap characterize yellow-orange overall, with
veil, and a bulbous stem. It mainly this species. It is common under birches prominent girdles of
occurs in beech woods in Eurasia. in N. Eurasia and N. America. yellow veil on the stem.
 3
⁄16 –1¼ in ⅛–5⁄16 in
221
0.5–3 cm 0.3–0.8 cm

¾–2¾ in TINY PIXIE CAP

2–7 cm Galerina calyptrata
cap (topside) cap (underside) VARIABLE
OYSTERLING f: Hymenogastraceae
PEELING OYSTERLING Crepidotus variabilis Found in Eurasia, this
Crepidotus mollis f: Crepidotaceae fungus is one of many
f: Crepidotaceae One of several similar species only identifiable
This fungus occurs in Eurasia and species in Eurasia and under a microscope. Its
N. America. Its small, fan-shaped, pale N. America, its cap is rounded cap is orange-
cap has a gelatinous skin, which peels dry with fine fibers. It brown and striated.
easily. The stem is absent or very short. usually lacks a stem.
cap not
fully
expanded
2–4¾ in ⅜–2 in
5–12 cm attached 1–5 cm
to wood
ROOTING POISONPIE DEADLY
Hebeloma radicosum GALERINA
f: Hymenogastraceae Galerina marginata
The deeply rooting stem with f: Hymenogastraceae
a large ring, buff cap with flat This species has an orange-brown cap
scales, and strong smell of and a small ring on the stem. It grows on
marzipan characterize this fallen trees in Eurasia and N. America.
Eurasian species.

1¼–2¾ in
3–7 cm

1½–3½ in SPLIT FIBERCAP

4–9 cm Inocybe rimosa
POISONPIE f: Inocybaceae
Hebeloma crustuliniforme Common in mixed
f: Hymenogastraceae woodland in Eurasia
From Eurasia and N. America, this and N. America, its
species strongly smells of radish. Its cap pointed, fibrous cap is
is ivory-white to buff, and slimy when wet. straw-yellow and the
In damp weather, the gills exude droplets. stem is tall and slender.

red band 1¼–2¾ in

3–7 cm
⅜–1½ in
1–4 cm

white
STAR FIBERCAP form
Inocybe asterospora
f: Inocybaceae
A flattened bulb at the stem’s radial
base and star-shaped spores fibers
WHITE FIBERCAP
distinguish this small, brown Inocybe geophylla
fibrous-capped species 1¼–3½ in f: Inocybaceae
from Eurasia. 3–9 cm This is one of the most common
stout stem Inocybe species in woods in Eurasia and
N. America. It has a conical, silky cap
and a slender stem.

violet
5
⁄16 –1½ in
form
⅜–1¾ in
1–4.5 cm 0.8–4 cm

TORN FIBERCAP
Inocybe lacera LILAC LEG
f: Inocybaceae DEADLY FIBERCAP FIBERCAP
Found in Eurasia and Inocybe erubescens Inocybe griseolilacina
N. America, this fungus f: Inocybaceae f: Inocybaceae
is distinguished by its Found in mixed woodland on chalk A common species of
cylindrical spores. It has in Eurasia, this uncommon species beech woods in Eurasia,
a scaly, fibrous cap and a has a fibrous cap that discolors with it has a scaly cap and a pale
slender brown stem. age, and stout stem that bruises red. violet, slender stem.
 AGA RICA LES
222
⅜–1½ in
1–4 cm green stem apex
MUSHROOMS • AGARICALES

gills are
broadly
attached

1–2¼ in ½–2 in
2.5–6 cm 1.5–5 cm

MEADOW WAXCAP SNOWY WAXCAP PARROT WAXCAP

Cuphophyllus pratensis Cuphophyllus virgineus Gliophorus psittacinus
f: Hygrophoraceae f: Hygrophoraceae f: Hygrophoraceae
One of the larger meadow waxcaps, The most common waxcap in all This fungus occurs in Eurasia and
it is found in Eurasia and N. America. types of grassland in Eurasia, its N. America. Its slimy cap is vivid
It has a stout stem and a fleshy cap waxy cap and slender stem are green to orange and the upper
with gills running down the stem. translucent. It has decurrent gills. part of the stem is bright green.
1¼–2¾ in
fleshy, waxy, 3–7 cm
pink gills
⅜–2 in PINK WAXCAP
1–5 cm Porpolomopsis calyptriformis
½–2¼ in 1½–4¾ in f: Hygrophoraceae
1.5–6 cm 4–12 cm This highly recognizable but rare
BLACKENING WAXCAP fungus has a pointed pink cap,
Hygrocybe conica fragile pale stem, and waxy gills.
f: Hygrophoraceae Found in Eurasia, it grows in
Quite common in grasslands unimproved meadows.
and woodlands in Eurasia and
N. America, this species has pale yellow
a conical reddish orange cap margin
and fibrous stem that blacken
with age or bruising. 1½–3¼ in
SCARLET WAXCAP 4–8 cm
Hygrocybe coccinea
f: Hygrophoraceae
The scarlet, waxy cap, gills, and CLUB FOOT
stem make this a very striking Ampulloclitocybe clavipes
fungus of unimproved grasslands f: Hygrophoraceae
in Eurasia and N. America. Common in mixed woods in
late fall, this species from
Eurasia and N. America
½–2¾ in has decurrent gills and a
1.5–7 cm swollen, spongy base.
CRIMSON WAXCAP
GOLDEN WAXCAP Hygrocybe punicea
Hygrocybe chlorophana f: Hygrophoraceae
f: Hygrophoraceae Found in unimproved
Found in Eurasia, this is grassland in Eurasia and
the commonest waxcap N. America, this uncommon
found in meadows. It has species is the largest blood-red
a bright yellow-orange, waxcap. Its stem is dry and
slightly sticky cap. fibrous with a white base. IVORY WOODWAX
Hygrophorus eburneus
f: Hygrophoraceae 1¼–3¼ in
This species is found in 3–8 cm
3
⁄16 –¾ in beech woods in Eurasia and
¾–2 in CONIFERCONE CAP 0.5–2 cm N. America. It has a sticky
2–5 cm Baeospora myosura cap and stem, thick gills,
⅜–2 in
f: Cyphellaceae and a flowery smell.
AMETHYST 1–5 cm Found in Eurasia and
DECEIVER N. America, this is one of a few
Laccaria amethystina fungi growing mostly on pine cones.
f: Hydnangiaceae It has crowded, narrow gills
This highly recognizable, and a velvety stem.
slender fungus from Eurasia 1¼–2 in
and N. America has an 3–5 cm
intense violet coloration pine cone
when fresh, and powdery
spores dusting the gills.
HERALD OF
DECEIVER WINTER
Laccaria laccata Hygrophorus hypothejus
f: Hydnangiaceae f: Hygrophoraceae
Variable in color—from brick-red Appearing in pine woods
to flesh-pink—with a dry cap and after frost in Eurasia and
thick gills, this species is N. America, this sticky
abundant in temperate woods in fungus has an olive-brown
Eurasia and N. America. cap and yellow stem.
cap frequently ST. GEORGE’S
splits as it ages MUSHROOM
Calocybe gambosa 223
f: Lyophyllaceae
3/8–1½ in This ivory-buff species
1–4 cm from Eurasia often grows
in fairy rings at woodland
margins in late spring. The
1¼–4¾ in mushroom smells strongly
3–12 cm of newly ground meal.
3⁄16–½ in
0.5–1.5 cm
PINK DOMECAP
Calocybe carnea
f: Lyophyllaceae
This species grows in
short grass in Eurasia and
N. America. Its smooth cap and
fibrous stem are rose-pink
and the gills are white.
2–4 in
5–10 cm

SILKY PIGGYBACK CLUSTERED DOMECAP

Asterophora parasitica Lyophyllum decastes
f: Lyophyllaceae f: Lyophyllaceae
One of two well-known parasitic Common on roadsides, pathsides,
species to live on rotten fruitbodies of and disturbed soils in Eurasia and
brittle-caps, this fungus from Eurasia N. America, this fungus forms clumps
and N. America has a rounded, silky cap. of tough caps with stout stems.

HORSEHAIR HAIRY PARACHUTE

PARACHUTE Crinipellis scabella
Gymnopus androsaceus f: Marasmiaceae
f: Marasmiaceae 3
⁄16 –½ in
Found on dead stems of grasses,
⅛–3/8 in 0.5–1.5 cm
Found in Eurasia, this fungus has this recognizable fungus is from
0.3–1 cm a distinctive black hairlike stem, Eurasia. It has a small cap and a
while its radially grooved cap long, slender stem covered with
is pale pinkish brown. dense, bristling brown hair.
3
⁄16 –3/8 in
cap depressed FAIRY RING 0.5–1 cm
3/8–2 in
at center 1–5 cm CHAMPIGNON
Marasmius oreades
f: Marasmiaceae
This classic fairy-ring mushroom
is common in open grass in
Eurasia and N. America. It has
fleshy, beige caps with
radial thick gills.
groove
3
⁄16 –¾ in
0.5–2 cm

COMMON BIRD’S NEST

Crucibulum laeve
f: Nidulariaceae
Forming miniature nests with
½–1½ in spore-filled, egglike cases,
1.5–4 cm wiry stem this common species may be
hard to spot on woody debris.
It occurs in Eurasia and
GARLIC PARACHUTE N. America.
Mycetinis alliaceus
f: Marasmiaceae
This species grows in beech COLLARED
woods in Eurasia. It has a PARACHUTE
Marasmius rotula 3
⁄16 –3/8 in
tall, slender, blackish stem f: Marasmiaceae 0.5–1 cm
and a strong odor of
rancid garlic. Like little parachutes, this
mushroom’s rounded, depressed
caps are radially grooved. Its
tough stems are attached to
woody debris. It occurs in
Eurasia and N. America.

FLUTED BIRD’S NEST

Cyathus striatus
f: Nidulariaceae
This fungus is recognized by its hairy, brown, and tall fluted
nests containing 10 –15 egglike cases. It occurs on woody
debris across Eurasia and N. America, but is not common.
 AGA RICA LES
224
¾–2 in ⅜–1¼ in
SILVERLEAF FUNGUS 2–5 cm 1–3 cm
Chondrostereum purpureum
f: Cyphellaceae wavy margin
MUSHROOMS • AGARICALES

Common on cherry and

plum trees, this fungus is ⅜–2¼ in
found in Eurasia and 1–6 cm
N. America. Violet below
when young, it darkens to
purple-brown with age.

SAFFRONDROP
BONNET COMMON BONNET
Mycena crocata Mycena galericulata
f: Mycenaceae f: Mycenaceae
⅛–⅜ in Common in woods with Abundant in temperate woods
0.3–1 cm chalky soils in Eurasia, this in Eurasia and N. America, this
species exudes a bright fungus is variable in color. Its
purple underside saffron-orange juice pinkish-gray gills have linking
when it is broken. veins and cross-ridges.
ORANGE BONNET 3⁄16–1 in ⅜–1½ in
Mycena acicula 0.5–2.5 cm 1–4 cm
domed
f: Mycenaceae cap center
Common in leaf litter and debris YELLOWLEG BONNET
in broadleaf woods in Eurasia Mycena epipterygia
and N. America, this fungus has f: Mycenaceae
a tiny, translucent cap that is Occurring in Eurasia and
striped almost to the center. N. America, this fungus grows
on acidic soils in woods and
heaths. Its cap and stem have
a sticky, peelable layer.

CLUSTERED BONNET
Mycena inclinata
1¼–2¼ in f: Mycenaceae
3–6 cm BLACKEDGE BONNET Found in dense clusters
Mycena pelianthina on wood in Eurasia and
f: Mycenaceae N. America, this fungus has ROSY BONNET ¾–2¼ in
Found in Eurasia, this fungus a cap with toothed margins Mycena rosea 2–6 cm
is characterized by a strong and a strong, soapy smell. f: Mycenaceae
smell of radish. It has purplish Found in beech woods
black-edged gills and a pale in Eurasia, this common
lilac to gray-brown cap. OLIVE OYSTERLING species has a robust pink
Sarcomyxa serotina cap and stem, and smells
f: Mycenaceae strongly of radish.
From Eurasia and
N. America, this mushroom
fruits in winter, often near
water on hardwood tree trunks. 1¼–4 in club-shaped
Its cap is slimy when wet. 3–10 cm stem

SPINDLE TOUGHSHANK WOOD WOOLLYFOOT

Gymnopus fusipes Gymnopus peronatus
f: Omphalotaceae f: Omphalotaceae
1½–3¼ in Abundant in Eurasian oak woods The common name of this
4–8 cm from early summer onward, this 1–2¼ in species from Eurasia refers
fungus forms large clumps of tough 2.5–6 cm to the stiff, fuzzy hairs at the
fruit bodies on tree roots. base of the stem.

2–6 in
5–15 cm

1½–4 in
4–10 cm

1¼–2¼ in
SPOTTED 3–6 cm
TOUGHSHANK
Rhodocollybia maculata BUTTER CAP
f: Omphalotaceae Rhodocollybia butyracea
Common in mixed f: Omphalotaceae JACK O’LANTERN
woods across Eurasia and This fungus is abundant in Omphalotus illudens
N. America, this fungus woods in Eurasia and N. America. f: Omphalotaceae
has a white cap, stem, and Varying from blackish or reddish Found across Eurasia and N. America, this
gills. The crowded gills brown to dark ocher, its cap is bright orange, poisonous fungus is famous for gills
stain rust-red with age. greasy to the touch. that glow in the dark with an eerie, greenish light.
 ¾–6 in 1–4 in
2–15 cm 2.5–10 cm
225

⅜–2¼ in PORCELAIN ROOTING SHANK

MUSHROOMS • AGARICALES
1–6 cm FUNGUS Hymenopellis radicata
Mucidula mucida f: Physalacriaceae
VELVET SHANK f: Physalacriaceae Found across Eurasia
1¼–4 in 1–4 in Flammulina velutipes Usually found on beech logs and N. America, this
3–10 cm 2.5–10 cm f: Physalacriaceae in Eurasia, this fungus has deep-rooted fungus has a
HONEY FUNGUS Growing throughout winter gray-white caps that are stiff, tall stem. The cap is
Armillaria lutea WRINKLED PEACH in Eurasia and N. America, slimy when wet. The tough slimy when wet, and the
f: Physalacriaceae Rhodotus palmatus this fungus is characterized stem has a thin ring. gills are widely spaced.
Found in woods across f: Physalacriaceae by its often sticky cap and
Eurasia, this species is Found on fallen logs of mainly velvety stem.
usually found on the elm, this uncommon species of
ground. It is a weak Eurasia and N. America has an
parasite on the trees unusual, peach-pink, wrinkled caps often
around which it grows. cap and a fruity odor. overlap

BRANCHING OYSTER
Pleurotus cornucopiae
f: Pleurotaceae
Growing in clusters on fallen logs, usually
2¼–8 in
of elm, this fungus has trumpet-shaped 6–20 cm
caps with gills running down the short,
frequently branching stems. It occurs
in Eurasia and N. America.

1½–4¾ in
4–12 cm

tiny glistening granules

OYSTER MUSHROOM
Pleurotus ostreatus
f: Pleurotaceae
From Eurasia and N. America,
this species is found on trees
and logs. Its shelflike caps vary
from blue-green to pale buff. white veil
The stem is almost absent. breaks into
¾–1¼ in patches
2–3 cm

GLISTENING INKCAP
Coprinellus micaceus
f: Psathyrellaceae
Usually found in clusters, this
mushroom’s rounded, grooved caps 1–3 in
2.5–7.5 cm
are dusted with micalike flakes of
veil. It is common on wood in
Eurasia and N. America.
3⁄16–⅜ in
0.5–1 cm WEEPING WIDOW
Lacrymaria
FAIRIES’ BONNETS lacrymabunda
Coprinellus disseminatus f: Psathyrellaceae
f: Psathyrellaceae Common on pathsides
Often found in clusters on rotted and disturbed soils in
stumps in Eurasia and N. America, Eurasia and N. America,
½–2¾ in
it has tiny, umbrellalike caps with it is named after its black
1.5–7 cm deeply fluted grooves. The gills gills that weep droplets
are blackish when mature. from their edges.

1–3¼ in 2–3¼ in
2.5–8 cm 5–8 cm

5⁄16–1½ in
0.8–4 cm

PALE BRITTLESTEM CLUSTERED MAGPIE INKCAP

Psathyrella candolleana BRITTLESTEM Coprinopsis picacea
f: Psathyrellaceae Psathyrella multipedata COMMON INKCAP f: Psathyrellaceae
Usually found in clusters on f: Psathyrellaceae Coprinopsis atramentaria Found on chalky soils in
woody debris in early summer, The densely clustered stems f: Psathyrellaceae Eurasian woodlands, this
this fungus has fragile, slender of this species are joined at Occurring in Eurasia and N. America, this uncommon species has white,
stems and pale or buff caps. It the base. It is usually found species has egg-shaped caps that dissolve to fluffy scales that contrast with
occurs in Eurasia and N. America. in open grass across Eurasia. a blackish fluid as the spores are released. the dark gray-brown cap.
 AGA RICA LES
226 ⅜–2¼ in
4–10 in
10–25 cm
1–6 cm
1½–4 in BEEFSTEAK FUNGUS
4–10 cm Fistulina hepatica
f: Schizophyllaceae
MUSHROOMS • AGARICALES

YELLOW SHIELD
DEER SHIELD Pluteus chrysophlebius 1–3¼ in Resembling a fleshy steak,
Pluteus cervinus f: Pluteaceae 2.5–8 cm with bloodlike red juice,
f: Pluteaceae A golden to greenish-yellow this species is particularly
Found in Eurasia and cap, yellow gills turning common in warmer parts
N. America, this species is variable in pink, and a whitish stem of Eurasia and N. America.
color, with a radially fibrous cap, often characterize this species. WILLOW SHIELD
dome-centered; pink gills not attached Found in Eurasia, it grows Pluteus salicinus
to the stem; and a fibrous stem. on decayed wood. f: Pluteaceae
Common in broad-leaved
woodlands in Eurasia, this
species can be identified by
fine, silky-hairy the bluish-gray staining at the
surface base of its slender stem. ⅜–2 in
1–5 cm

cap (underside)
COMMON PORECRUST
Schizophyllum commune
SILKY ROSEGILL 4–10 in f: Schizophyllaceae
Volvariella bombycina 10–25 cm Found in Eurasia and
f: Pluteaceae N. America, this fan-shaped
Growing on deciduous trees in Eurasia and fungus is identified by the
N. America, this rare species has a white to pale gill-like spore surface on
lemon cap and a whitish stem with a baglike, cap
(upper side) the underside of the cap.
thin veil enclosing the base of the stem.
greenish orange cap
yellow gills center

2¼–5½ in
6–14 cm

STUBBLE ROSEGILL
Volvopluteus gloiocephalus
f: Pluteaceae
Quite common in fields and old
stubble, and on woodchip mulch,
in Eurasia and N. America, this
species has a sticky, gray cap.

1¼–2¾ in 1¼–2¾ in
3–7 cm 3–7 cm 2–4 in
5–10 cm
CONIFER TUFT
Hypholoma capnoides SULPHUR TUFT BRICK TUFT
f: Strophariaceae Hypholoma fasciculare Hypholoma lateritium
Found in Eurasia f: Strophariaceae f: Strophariaceae
and N. America, this Abundant in temperate woods This species is characterized by a
uncommon species in Eurasia and N. America, this fleshy cap with veil fragments and
occurs on conifer wood. species has greenish yellow gills pale yellow gills that mature to
Its whitish gills mature that mature to dark purple, and lavender. It grows on hardwoods
to grayish lilac. are easy to recognize in the field. in Eurasia and N. America.
 SHEATHED ALDER SCALYCAP
WOODTUFT Pholiota alnicola
Kuehneromyces mutabilis f: Strophariaceae 227
¾–2¾ in f: Strophariaceae Found in Eurasia, this
2–7 cm Often confused with the deadly species is characterized by
Galerina marginata, this species ½–2¼ in a sticky cap and tufted
1.5–6 cm

MUSHROOMS • AGARICALES
ring on can be identified by its sticky habit. Despite its common
stem cap, scaly stem, and brown gills. name, it is usually seen
It occurs in Eurasia and at the base of birches.
N. America. 1¼–4¾ in
3–12 cm
REDLEAD ROUNDHEAD 1¼–2¾ in
Leratiomyces ceres 3–7 cm
f: Strophariaceae
Previously known as Stropharia
aurantiaca, this species has a bright
red cap and reddish, flushed stem.
It grows among woodchips in
Eurasia and N. America.

2–6 in
5–15 cm GOLDEN SCALYCAP
DUNG
3⁄16–1½ in ROUNDHEAD Pholiota aurivella
0.5–4 cm Protostropharia SHAGGY SCALYCAP f: Strophariaceae
semiglobata Pholiota squarrosa This rather common species is found
f: Strophariaceae f: Strophariaceae on beech logs or trunks in Eurasia
Occurring on animal dung Growing in Eurasia and and N. America. Its sticky golden
or in grass where animals N. America, this fungus has a caps have dark orange-brown scales.
graze, this species is found distinctive dry, sharply scaly
in Eurasia and N. America. cap and stem; pale yellow gills;
3⁄16–¾ in 1¼–2¾ in
It has a sticky cap and a and a smell of corn or radish.
0.5–2 cm 3–7 cm slender stem with a ring.

LIBERTY CAP BLUE-GREEN scaly, furry

Psilocybe semilanceata SLIMEHEAD surface
f: Strophariaceae Stropharia cyanea
This well-known agaric f: Strophariaceae
has a conical, dull yellow This fungus has a
blue-green cap that 2¼–6 in
cap with a distinct point. 6–15 cm
Common in meadows in fades to yellow. It is
late fall, it occurs in found in Eurasia and
Eurasia and N.America. N. America.
1½–2¾ in 2–4 in
4–7 cm 5–10 cm

cap margin
with veil
remnants
PLUMS AND CUSTARD
2–6 in WHITELACED SHANK COMMON CAVALIER Tricholomopsis rutilans
5–15 cm Megacollybia platyphylla Melanoleuca polioleuca f: Incertae sedis
f: Incertae sedis f: Incertae sedis Common on pine stumps, this
Found in Eurasia and Common in grass in species has a reddish purple
N. America, this species Eurasia, this species has a cap and stem, and golden-
has a radially fibrous, pale gray-brown cap with white yellow gills. It
gray-brown cap; widely gills. Its stem has blackish grows in Eurasia and
spaced, deep gills; and a stem flesh at the base. N. America.
base with rootlike strands.
SPECTACULAR
RUSTGILL
Gymnopilus junonius ⅜–1½ in
f: Incertae sedis 1–4 cm
Occurring in Eurasia, this 2–8 in
fungus is found in clumps, PETTICOAT 5–20 cm
usually at the base of a tree. MOTTLEGILL
It has a dry cap with crowded,
Panaeolus papilionaceus
shallow yellowish gills.
f: Incertae sedis root attaches
to conifer wood
The minute toothed remains of ¾–3¼ in
the veil at the cap margin and 2–8 cm
⅜–2¼ in mottled black gills characterize 1¼–2¾ in
1–6 cm this species. It is found in 3–7 cm
Eurasia and N. America. CINNABAR
EGGHEAD MOTTLEGILL POWDERCAP TROOPING FUNNEL
Panaeolus semiovatus GOBLET Cystodermella cinnabarina Infundibulicybe geotropa
f: Incertae sedis Pseudoclitocybe cyathiformis f: Incertae sedis f: Incertae sedis
Found on animal dung, this f: Incertae sedis Found in Eurasia and This fungus has a
species occurs in Eurasia and This distinctive species has a very dark N. America, this species has a funnel-shaped, fleshy cap;
N. America. It is characterized coloration and a tall, fibrous stem. brick-red cap and pale cream a tall stem; and pale
by a sticky, gray cap and a Found in Eurasia, it is common in gills. The cap and stem leather-brown coloring.
ring around its tall stem. late fall and winter. have a granular surface. It occurs in Eurasia.
 BOL E TA L E S wrinkled
cap surface
228 This order contains fleshy fungi and includes
those with both pored and gilled fruit bodies,
most with cap and stem, but some are crusts,
MUSHROOMS • BOLETALES

puffballs, or trufflelike. The majority live in 4–10 in 2¾–6 in

10–25 cm 7–15 cm
association with trees (mycorrhizal), but some
feed on dead wood and cause a brown rot, while
orange-brown
others are parasitic. The spore-producing layer— cap
the hymenium—is easily loosened 1½–6 in
from the flesh. 4–15 cm BAY BOLETE
Imleria badia PENNY BUN SUMMER BOLETE
f: Boletaceae Boletus edulis Boletus reticulatus
cylindrical A common species f: Boletaceae f: Boletaceae
stem found among conifers or Found worldwide, this species A cracking, matte, brown cap
beech trees in Eurasia has fine white markings on its and a stem with a fine white net
and N. America, its color stem, unchanging cream flesh, extending to its base distinguish
varies from orange- and white pores that this species from Eurasia and
brown to bay. mature to yellow. eastern N. America.
sticky cap
PARASITIC BOLETE
Pseudoboletus parasiticus
2¼–5½ in BITTER BEECH BOLETE f: Boletaceae
6–14 cm Caloboletus calopus This small mushroom from
f: Boletaceae Eurasia and N. America
From Eurasia and western grows only on the common
N. America, its cap varies from white to earth-ball, causing its host
buff. Its yellow pores and cream to become hollow.
flesh bruise blue.

black scales
on face
2¼–6 in
6–15 cm

1¼–2 in
3–5 cm

BITTER BOLETE
Tylopilus felleus
f: Boletaceae ¾–2¾ in
PEPPERY BOLETE This species, found in 2–7 cm
Chalciporus piperatus Eurasia and N. America, is
f: Boletaceae characterized by pores that
Common under conifers turn pink with age and a
and also found with fly agaric strongly netted stem.
under birches in Eurasia and
N. America, it has cinnamon
pores and yellow flesh.
2–4 in
5–10 cm
common
earthball

OLD MAN OF
THE WOODS
Strobilomyces strobilaceus
f: Boletaceae
Found in Eurasia and
N. America, this rare species
is unmistakable with black
woolly scales on its cap and
stem, and white tubes.
 2–39 in 2–3½ in
5–100 cm 5–9 cm
229
3¼–6 in 2¼–6 in ⅜–¾ in
8–15 cm 6–15 cm 1–2 cm

MUSHROOMS • BOLETALES
ORANGE BIRCH BROWN BIRCH
BOLETE BOLETE
Leccinum versipelle Leccinum scabrum
f: Boletaceae f: Boletaceae
Found in Eurasia, this species One of many similar species,
has a yellow-orange fleshy this fungus is from Eurasia and
cap. The stem has black, N. America. Its flesh may flush
woolly flecks and the pink when cut, and the cap WET ROT
flesh stains lilac-black. is sticky when wet. STALKED
PUFFBALL-IN-ASPIC Coniophora puteana BAROMETER EARTHSTAR
Calostoma cinnabarinum f: Coniophoraceae Astraeus hygrometricus
f: Calostomataceae Found worldwide, this fungus f: Diplocystidiaceae
From N. America, this species’ forms a brown sheet of tissue, Frequent in Eurasia and N. America,
bright cinnabar-red ball on often warty or wrinkled, on this species’ starlike arms peel back
a stem emerges from wet lumber. It causes serious to reveal the inner spore-filled ball,
a gelatinous layer. damage to buildings. but close again in dry weather.
2–3¼ in ½–2 in
5–8 cm 1.5–5 cm

COPPER SPIKE BROWN ROLLRIM

Chroogomphus rutilus Paxillus involutus
f: Gomphidiaceae f: Paxillaceae
CORNFLOWER Common under pines Common in mixed woods in
BOLETE ROSY SPIKE in Eurasia and western Eurasia and N. America, the
Gyroporus cyanescens Gomphidius roseus N. America, this species downy, inrolled cap margin
f: Gyroporaceae f: Gomphidiaceae has a pointed, coppery 2¼–6 in and soft yellow gills staining
Growing on acidic soils Growing in association brown cap. 6–15 cm brown characterize this species.
in Eurasia and eastern with the bolete Suillus bovinus
N. America, this uncommon under pines in Eurasia, this
species is distinguished by fungus has a slimy, rose-pink
its brittle, hollow stem. cap with grayish gills. 1½–3¼ in
4–8 cm

¾–3¼ in FALSE CHANTERELLE

1½–4 in 2–8 cm Hygrophoropsis aurantiaca
4–10 cm f: Hygrophoropsidaceae
Sometimes mistaken for a
chanterelle, this species from
Eurasia and N. America is
distinguished by its crowded,
¾–2 in multi-forked, soft gills.
COMMON EARTH-BALL 2–5 cm
Scleroderma citrinum 4–12 in
f: Sclerodermataceae 10–30 cm
This potatolike fungus is common POTATO EARTHBALL
in damp woodland in Eurasia and Scleroderma bovista
N. America. Its thick skin has dark f: Sclerodermataceae DYEBALL VELVET ROLLRIM
scales and a spore-filled black interior. Common in woods in Pisolithus arhizus Tapinella atrotomentosa
Eurasia and N. America, f: Sclerodermataceae f: Tapinellaceae
its smooth outer skin cracks This species is found Common on pine stumps
into a fine mosaic. The inner worldwide on poor, sandy in Eurasia and N. America,
purple-black spore-mass soils associated with pines. it has a cap with an inrolled
dries brownish. The egglike spore-sacs in margin; soft, thick gills;
2–4 in its interior are embedded and a velvety stem.
cap skin 5–10 cm in blackish jelly.
peels off

1½–4 in yellow-brown cap

4–10 cm surface is dry and rough

VELVET BOLETE
Suillus variegatus 2¾–5 in
f: Suillaceae 7–13 cm
dots can Frequent under pines in
weep milky Eurasia, this species has a
latex felty-scaly cap, with dark
2–4 in 1¼–2¾ in 2–4 in cinnamon-brown pores,
5–10 cm 3–7 cm 5–10 cm and a ringless stem.

BOVINE BOLETE LARCH BOLETE

SLIPPERY JACK Suillus bovinus Suillus grevillei
WEEPING BOLETE Suillus luteus f: Suillaceae f: Suillaceae
Suillus granulatus f: Suillaceae Common under pines in Confined to larch woods
f: Suillaceae Found on pines in Eurasia and Eurasia and N. America, in Eurasia and N. America,
Common under pines in Eurasia N. America, this fungus has a slimy this species has a sticky this species is yellow-orange
and N. America, its ringless stem cap and yellow pores. The large ring cap and angular, to brick-red in color. The
is covered with glandular dots. on its stem has a lavender underside. irregular pores. stem has an ascending ring.
 CANTHARELLALES CRESTED CORAL
Clavulina coralloides
230 The species of the Cantharellales order may look like agarics (members of f: Clavulinaceae
Very common in woods in
the Agaricales order), but differ in several important respects. They may have Eurasia and N. America, this
fleshy fruitbodies with a cap and stem, but they lack true gills, having instead fungus forms a corallike
a smooth, wrinkled, or folded gill-like spore-producing surface on the mass of white branches,
underside. The spores are smooth and usually white to cream. each dividing into fine points.

1¼–3¼ in
⅜–2¼ in 3–8 cm
1–6 cm

3⁄16–¾ in
0.5–2 cm TRUMPET CHANTERELLE
Craterellus tubaeformis
f: Cantharellaceae
Often found in large troops
HORN OF PLENTY in mixed woods in Eurasia and N.
Craterellus cornucopioides America, this fungus has a
f: Cantharellaceae variable color and shallow, blunt
A distinctive species found veins rather than gills.
throughout Eurasia, this thin
trumpet grows in troops in
beech leaf litter. It produces
a white spore deposit.
cap often
depressed inrolled margin
at center ¾–4¾ in
2–12 cm
CHANTERELLE
2–6 in Cantharellus cibarius
5–15 cm f: Cantharellaceae
Found in Eurasia and
N. America, this species
has blunt-edged gills with
numerous crossveins and
a smell of apricots.

tapered stem

WOOD
HEDGEHOG
irregular, often Hydnum repandum
lobed cap f: Hydnaceae
Occurring in Eurasia
and N. America, this dusky
orange species is irregular
in shape, with tiny spines
underneath the cap.

GEASTRALES STRIATE EARTHSTAR COLLARED

Geastrum striatum EARTHSTAR
The Geastrales, or earthstars, share the f: Geastraceae 1¼–2½ in collar below Geastrum triplex
common character of a thick outer layer—the Found in Eurasia, this 3–6.5 cm spore sac f: Geastraceae
is one of the smaller Growing in Eurasia
peridium—which splits apart and peels back earthstars. Its pale gray and N. America, this
to form starlike arms. These reveal a central spore sac is stalked and is the most common
spore sac, like a puffball, from which the dark has a sharply pointed, earthstar. Its rays crack
brown, warted spores exit via a pore at the striated opening. to leave a cuplike collar
around the spore sac.
apex. Earthstars are found in leaf litter and 1½–4¾ in
bare, sandy soils—for example, in sand dunes. 4–12 cm

1¼–2¼ in 2–3¼ in
3–6 cm 5–8 cm

2¾–6 in
7–15 cm

SESSILE EARTHSTAR ARCHED EARTHSTAR PEPPER POT

Geastrum fimbriatum Geastrum fornicatum Myriostoma coliforme
f: Geastraceae f: Geastraceae f: Geastraceae
Found in Eurasia and N. America, This earthstar’s arms stay attached to a Found on dry, sandy soils in Eurasia
this fungus’s globe-shaped, pale brown disk of tissue on the soil, and its stalked and N. America, this rare species
fruit body splits open with 5–9 arms. Its spore sac has a prominent opening. It has a distinctive, large spore sac
grayish spore sac has a fringed opening. occurs in Eurasia and N. America. with several pore openings.
GOMPHALES ¾–2¼ in
2–6 cm stains green
Although some species were included with the
with age 231
2–4 in
chanterelles in Cantharellales, DNA analysis 5–10 cm
of the Gomphales fungi suggests that they are

MUSHROOMS • HYMENOCHAETALES
more closely related to the stinkhorn fungi in SCALY VASE
CHANTERELLE
the Phallales. They often form large fruit bodies, Gomphus floccosus
varying in shape from simple clubs (in genus f: Gomphaceae
Clavariadelphus) to trumpet- or chanterellelike Common in N. America,
structures with a complex spore-producing surface this fungus resembles a
fleshy, trumpet-shaped
(in genus Gomphus). vase with scales at the
top and wrinkled “gills”
on its underside.
GIANT CLUB
Clavariadelphus pistillaris
f: Clavariadelphaceae ½–1½ in
1.5–4 cm
A rare species of Eurasia and
N. America, this fungus forms a
large, swollen club, with a smooth
to slightly wrinkled surface, which
bruises purple-brown.
2¾–6 in
7–15 cm
1¼–3¼ in
3–8 cm UPRIGHT CORAL ROSSO CORAL
Ramaria stricta Ramaria botrytis
f: Gomphaceae f: Gomphaceae GREENING CORAL
A fairly common species in Eurasia Found in beech woods in Phaeoclavulina abietina
and N. America, it is always Eurasia and N. America, this f: Gomphaceae
attached to decaying wood or uncommon coral fungus has Found in conifer woods in Eurasia and
woodchip mulch. The branches pinkish-white branches N. America, this yellow-olive fungus has
are pale brown, bruising reddish. with deep red tips. densely packed branches that bruise green.

GLOEOPHYLLALES H Y M E NO C H A E TA L E S
This is an order of wood-decay fungi that is This group contains a number of diverse types of fungi
characterized by the ability to produce a brown including some crust fungi, polypores, such as Mensularia
rot of wood. The order Gloeophyllales has a single and Phellinus, as well as several agaric species, such as
family, the Gloeophyllaceae, which includes the Rickenella. The Hymenochaetales are defined through
genus Gloeophyllum. Some well-known bracket molecular studies and have few uniting physical
fungi on conifer wood are members of this genus. characteristics. Many feed on wood and may cause ⅜–2¼ in OAK CURTAIN CRUST
1–6 cm Hymenochaete rubiginosa
a white rot of wood. f: Hymenochaetaceae
Found across Eurasia, this fungus forms
4–16 in overlapping brackets and crusts mainly
ANISE MAZEGILL 10–40 cm on fallen oaks. Its tough fruit bodies have
Gloeophyllum odoratum concentric markings.
f: Gloeophyllaceae
This fungus, which grows
on decayed conifer wood, is WILLOW BRACKET
found in Eurasia and Phellinus igniarius
N. America. It has irregular f: Hymenochaetaceae
2–8 in brackets with yellow pores This gray to almost black
5–20 cm and an aniseed odor. perennial bracket from
Eurasia and N. America
grows over many years.
It is hoof-shaped and
DAC RY M YC E TA L E S extremely woody.
This order forms very simple, rounded or branched
gelatinous fruit bodies, usually with bright orange thick,
coloration. Smooth or wrinkled, the Dacrymycetales pale margin 1¼–3¼ in
3–8 cm
have unusual spore-producing cells (basidia), which
typically have two stout stalks ALDER BRACKET
(sterigmata), each bearing a Mensularia radiata
spore. They mainly feed f: Hymenochaetaceae
This deep reddish-brown
on dead wood. fungus with a paler margin
often forms vertical chains
of brackets on alder and other
trees in Eurasia and N. America.
⅛–⅜ in
YELLOW 0.3–1 cm
STAGSHORN
Calocera viscosa TIGER’S EYE ORANGE MOSSCAP
f: Dacrymycetaceae Coltricia perennis Rickenella fibula
Attached to conifer wood, this f: Hymenochaetaceae f: Incertae sedis in Rickenella clade
fungus is found in Eurasia and Frequently found in acidic heathlands This tiny species is common in mossy
3
⁄16 –1½ in ¾–4 in
N. America. Its clubs usually 0.5–4 cm 2–10 cm of Eurasia and N. America, this grasslands of Eurasia and N. America.
divide into branches with a fungus’s goblet-shaped, thin fruitbodies Its bright orange cap is radially
gelatinous, rubbery texture. are concentrically zoned. marked with a darker center.
 P O LY P O R A L E S bracket grows
in layers
6–12 in
15–30 cm
232 The Polyporales form a large group of diverse
fungi. Most of these are polypores—wood
decomposers whose spores are formed in tubes
MUSHROOMS • POLYPOR ALES

4–10 in
(rather like the tubes of the boletes) or sometimes 10–25 cm
on spines. Most lack fully developed stems and
grow shelf-, bracket-, or crustlike fruit bodies RED BELTED POLYPORE
on wood, but some have more or less central Fomitopsis pinicola
stems and grow at the base of trees. A few f: Fomitopsidaceae
Polyporales appear to grow from the soil. Occurring in Eurasia and N. America,
this hoof-shaped, woody bracket
is usually found on pines, and
sometimes on birches.
upper
surface

4–12 in
10–30 cm
underside
DYER’S MAZEGILL hairy
OAK MAZEGILL Phaeolus schweinitzii surface
Daedalea quercina f: Fomitopsidaceae
f: Fomitopsidaceae
Usually found at the base of conifers
Found on fallen oaks in Eurasia in Eurasia and N. America, this large,
and N. America, this perennial furry, cushionlike bracket is used
4–20 in
species has tough brackets with long, 10–50 cm
for the manufacture of dye.
mazelike pores on the underside.

RAZORSTROP FUNGUS 2–12 in CHICKEN OF THE WOODS

Fomitopsis betulinas 5–30 cm Laetiporus sulphureus
f: Fomitopsidaceae f: Fomitopsidaceae
This large, kidney-shaped bracket These large brackets, found
is pale brown to white. A damaging in Eurasia and N. America, usually
parasite of birches, it is found grow on oaks and sometimes
in Eurasia and N. America. on other trees.

4–23½ in LACQUERED
10–60 cm BRACKET
Ganoderma lucidum
f: Ganodermataceae
This deep reddish to 4–12 in
purple-brown bracket 10–30 cm
with a shiny surface may
have a long, lateral stem.
It is found in Eurasia
and N. America.
ARTIST’S BRACKET
Ganoderma applanatum
f: Ganodermataceae
Found in Eurasia and N. America, this
woody perennial bracket grows for GIANT POLYPORE
many years to reach huge sizes. Its fallen Meripilus giganteus
spores are a rich cinnamon-brown. f: Meripilaceae margin is wavy
One of the largest polypores, its and lobed
overlapping brackets are thick and fleshy.
It grows around beech and other trees
in Eurasia and N. America.

4–8 in JELLY ROT 1½–6 in

10–20 cm Phlebia tremellosa 4–15 cm
f: Meruliaceae
Found on logs in Eurasia,
this species has a pale,
velvety upper side and a
yellow-to-orange underside
with dense ridges.

1¼–2¾ in
3–7 cm
ZONED ROSETTE SMOKY BRACKET
Podoscypha multizonata Bjerkandera adusta
f: Meruliaceae f: Meruliaceae
Growing on soil from buried This common bracket, found
oak roots, this rare species in Eurasia and N. America,
occurs in Eurasia. It has can be identified by the
4–20 in surface densely packed lobes that ash-gray pore surface
10–50 cm bruises black form a circular mass. on its underside.
 2–12 in TINDER BRACKET 3¼–6 in BLUSHING BRACKET BIRCH MAZEGILL
5–30 cm Fomes fomentarius 8–15 cm Daedaleopsis confragosa Lenzites betulina
f: Polyporaceae f: Polyporaceae f: Polyporaceae 233
This gray-brown, hoof-shaped, One of the most common Usually found on birches in
perennial bracket grows on brackets, especially on willows, Eurasia and N. America, this
birches and other deciduous in Eurasia and N. America, this tough, leathery species has
trees. It is found in Eurasia semicircular species has cream pores that can be extremely
and N. America. pores that bruise pinkish red. elongated in gill-like ridges.

4–23½ in 1¼–4 in
10–60 cm upper surface 3–10 cm

DRYAD’S SADDLE
Cerioporus squamosus underside
f: Polyporaceae funnel-shaped
Found in early summer in cap
Eurasia and N. America, these
circular or fan-shaped brackets
have concentric scales and
pores on the underside. 2–8 in BLACK FOOTED TUBEROUS
5–20 cm POLYPORE POLYPORE
Polyporus durus Polyporus tuberaster
velvety f: Polyporaceae f: Polyporaceae
stem base This fungus has a funnel-shaped, Growing on fallen branches in
leathery cap and its stem is 2–8 in Eurasia and N. America, this
black at the base. It grows on 5–20 cm species may root into the
fallen beech logs in Eurasia ground and form a large,
and N. America. tuberous mass.

1¼–4 in ¾–1½ in
1¼–3¼ in 3–10 cm 2–4 cm
3–8 cm
WINTER POLYPORE PURPLEPORE
Lentinus brumalis BRACKET
f: Polyporaceae Trichaptum abietinum
This small species grows f: Polyporaceae
on fallen branches in Found on fallen conifers
Eurasia and N. America. It in Eurasia and N. America,
has rather large, decurrent CINNABAR BRACKET this fan-shaped bracket has
(running down the stem) Pycnoporus cinnabarinus concentric zones of pale gray,
pores. The stem is central f: Polyporaceae often tinged green by algae,
or off-center. Growing on dead deciduous with a purple margin.
timber in Eurasia and
N. America, this rare species
has a bright reddish-orange, 2–4¾ in
leathery, annual bracket. 5–12 cm

HAIRY BRACKET
Trametes hirsuta
f: Polyporaceae
This semicircular bracket
is covered in minute hairs.
It grows on dead deciduous
wood or sometimes on old
gorse stems in Eurasia
and N. America.
TURKEYTAIL ¾–2¾ in
Trametes versicolor 2–7 cm
f: Polyporaceae
Found in Eurasia and N. America in a variety of
colors, this species has brackets with concentric
fleshy, bright zones of different colors and a white pore surface.
yellow-orange
bracket
LUMPY BRACKET
Trametes gibbosa
f: Polyporaceae
This cream-colored bracket is
often stained green from algae.
4–12 in Its pores may be long and small-lobed
10–30 cm
mazelike. It grows on fallen bracket
4–16 in deciduous logs in Eurasia
10–40 cm and N. America.
¾–2¼ in
2–6 cm
WOOD
CAULIFLOWER
Sparassis crispa HEN OF THE WOODS
f: Sparassidaceae Grifola frondosa
Growing at the base of f: Sparassidaceae
conifers in Eurasia and Found in Eurasia and N. America, this
N. America, this species has species grows at the base of oak, often
cream lobes that are flattened where lightning has struck. It has a basal stem
and fleshy like a cauliflower. dense cluster of small brackets.
 RU SSU L A L E S
234 The best-known genuses within this order are Russula 1¼–2¼ in LIVER MILKCAP
and Lactarius, which, although resembling typical 3–6 cm Lactarius hepaticus
f: Russulaceae
mushrooms, are not related at all to the true Agaricales. Associated with pines in Eurasia,
MUSHROOMS • RUSSULALES

Apart from the cap-and-stem shapes, Russulales this mushroom has a smooth
produce fruit bodies in a wide range of forms. Most cap with a grooved margin. The
Russulales have spores with warts that stain blue-black gills when cut bleed a white latex
that stains yellow.
in iodine solutions. When cut, the Lactarius species
produce a latex, which may be white to colored.
1½–3¼ in OAKBUG MILKCAP
4–8 cm Lactarius quietus
f: Russulaceae UGLY MILKCAP
BEECH MILKCAP Common under oaks Lactarius turpis
Lactarius blennius in Eurasia, its reddish f: Russulaceae
f: Russulaceae brown cap has darker A common species
This is a common species zones, and the flesh has of birch woods in
growing with beech in Eurasia. a sweet, oily smell. N. America and Eurasia, it is
Slimy when wet, the gray-green olive-green to almost black
1½–3½ in cap has spots around the margin. 2–6 in in color and has a slimy cap.
4–9 cm 5–15 cm

depressed
cap center

2–6 in
5–15 cm

SAFFRON MILKCAP
Lactarius deliciosus
f: Russulaceae
Found with pines in
N. America and Eurasia, this
fungus has an orange-zoned
and spotted cap that stains
green with age. The flesh
bleeds orange-red latex.

grooved
cap margin

2–6 in 1¼–2¼ in
5–15 cm 3–6 cm

WOOLLY MILKCAP CURRY MILKCAP

Lactarius torminosus Lactarius camphoratus
f: Russulaceae f: Russulaceae
Common under birches in As the fruit body of this fungus
N. America and Eurasia, dries, it gives off a smell of
this mushroom has a dark curry, which persists for many
pink, hairy cap that is weeks. It is found in Eurasia
strikingly zoned. and N. America.
hairy
and rolled-in
margin

3¼–8 in
8–20 cm

2¼–4 in SOOTY MILKCAP PEPPERY MILKCAP

6–10 cm Lactarius fuliginosus Lactifluus piperatus
f: Russulaceae f: Russulaceae
Found in deciduous woods This uncommon species from mixed
in Eurasia, this uncommon woods in N. America and Eurasia has a
species has a dark brown cap funnel-shaped cap with extraordinarily
and stem that bleed a white crowded, narrow gills. It exudes
latex that rapidly turns pink. white latex when cut.
 GREEN BRITTLEGILL
Russula aeruginea
dry, smooth
cap surface
f: Russulaceae 235
Common under birches in
N. America and Eurasia, this
mushroom’s pale olive to

MUSHROOMS • RUSSULALES
2–6 in 2–4 in grass-green cap has tiny
5–15 cm 5–10 cm rust-colored spots. Its
1½–3½ in
4–9 cm spores are pale cream.

CHARCOAL BURNER BLOODY BRITTLEGILL

Russula cyanoxantha Russula sanguinaria
f: Russulaceae f: Russulaceae 1¼–2¾ in 2–4 in
Growing in mixed woods in Associated with pines in 3–7 cm 5–10 cm
N. America and Eurasia, this N. America and Eurasia,
mushroom’s cap varies from this species has a scarlet BEECHWOOD YELLOW SWAMP
purple-lilac to uniformly cap and red-streaked stem. SICKENER BRITTLEGILL
green. The gills are forked, Its spores are pale ocher. Russula mairei Russula claroflava
flexible, and greasy. f: Russulaceae f: Russulaceae
Exclusively found with A common species on
beech in Eurasia, this moss in boggy birch woods
species has a scarlet cap in N. America and Eurasia,
and bluish white gills. its cap, yellowish cream
gills, and white stem
all bruise gray-black.

2–4¾ in dry and matte

5–12 cm cap surface
1¼–3¼ in
3–8 cm

1½–4¾ in
4–12 cm
OCHER BRITTLEGILL SICKENER
Russula ochroleuca Russula emetica
f: Russulaceae f: Russulaceae
One of the commonest species found Growing in wet pine
1½–4 in in Eurasia, it is identifiable by its woods in N. America and
4–10 cm matte yellow-ocher or greenish Eurasia, the sickener
yellow cap and white gills. has a bright scarlet cap
that contrasts with its pure
white gills and stem.
PRIMROSE BRITTLEGILL
Russula sardonia
f: Russulaceae gills may have
Found in Eurasia, this species grows red edges
in association with pine. Variable in
color, from purple to green or ROSY BRITTLEGILL
yellow, it has a fruity odor. 3¼–6 in STINKING Russula lepida
8–15 cm BRITTLEGILL f: Russulaceae
Russula foetens stem often Found in Eurasia, this fungus
f: Russulaceae flushes red has a carmine-red, dry, hard
2¼–6 in This large, orange-brown fungus cap that fades rapidly. Its
6–15 cm
with a grooved, lumpy cap margin stem may also be red
CRAB BRITTLEGILL grows in N. America and Eurasia. and the flesh has a smell
Russula xerampelina It has a sour, rancid odor. similar to cedarwood.
f: Russulaceae
Found in N. America and Eurasia,
this fungus is one of many related
species that can only be identified
under a microscope or by habitat. EARPICK FUNGUS 3⁄16–¾ in
Auriscalpium vulgare 0.5–2 cm 2–10 in 4–16 in
f: Auriscalpiaceae 5–25 cm 10–40 cm
¾–2¼ in Growing on pine cones
2–6 cm in N. America and Eurasia,
this unique mushroom looks
like a bent-over spoon with
minute spines hanging down
from the small, furry cap.
4–20 in
10–50 cm

BLEEDING
BROADLEAF CRUST
HAIRY CURTAIN CRUST Stereum rugosum ROOT ROT CORAL TOOTH
Stereum hirsutum f: Stereaceae Heterobasidion annosum Hericium coralloides
f: Stereaceae Growing in N. America f: Bondarzewiaceae f: Hericiaceae
Found in N. America and Eurasia, and Eurasia, this species Usually parasitic on Usually found on beech trees
this mushroom varies in shape forms small crusts, and conifers in N. America in N. America and Eurasia, this
from fully crustlike to small occasionally small brackets and Eurasia, this species endangered species has white
overlapping brackets. It is hairy on wood. Its upper surface has a pale brown crust fruit-body branches with pendent
above and smooth below. bleeds red where cut. that darkens with age. spines on the lower surfaces.
 AU R IC U L A R I A L E S WITCHES’ BUTTER
Exidia nigricans
236 Although often grouped with other jelly fungi, the f: Auriculariaceae
Found in temperate Eurasia
Auriculariales are separated by their unusual and N. America, frequently on
basidia (spore-producing cells). These vary in shape hardwood trees, it resembles a
MUSHROOMS • AURICULARIALES

but all are partitioned by dividing membranes into wrinkled mass of gelatinous
four divisions, with each division producing a spore. tar. It shrivels when dry to
a hard, black mass. 3/8–31/4 in
3/4–4 in 1–8 cm
2–10 cm
broad, thick,
jellylike bracket

11/2–43/4 in
4–12 cm

JELLY EAR JELLY TOOTH

TRIPE FUNGUS 11/2–6 in Auricularia auricula-judae Pseudohydnum gelatinosum
Auricularia mesenterica 4–15 cm f: Auriculariaceae f: Incertae sedis soft, peglike spines
f: Auriculariaceae Common on dead wood of deciduous This species from Eurasia and N. America on underside,
Found in Eurasia, this species is common on dead wood, trees in Eurasia and N. America, this has a pale translucent gray to pale brown on which spores
especially elm. It resembles a small bracket fungus from the fungus has thin, elastic “ears” that are coloration. It is occasionally found are formed
top, and has a wrinkled, rubbery, grayish purple underside. velvety outside and wrinkled inside. on conifer stumps.

THELEPHORALES PHALLALES
This diverse group includes bracket fungi, crust Named for the phallic shape of many of the
fungi, earthfans, and toothed fungi. Many of these species in this group, such as the stinkhorns,
have tough, leathery flesh, and commonly feature this order also contains some false truffles.
knobbed or spiny spores. The group was only The stinkhorns “hatch” from an egglike
2–4 in
identified as a result of molecular studies as the 5–10 cm structure, often in just a few hours.
Thelephorales have few physical features in common. DRAB TOOTH
Bankera fuligineoalba
f: Bankeraceae
An uncommon species of conifer
woods in Eurasia, this fungus has
a short, stout stem. Its cap has an
underside covered with tiny,
grayish-white spines.

spores on
inside of cage
11/4–4 in
3–10 cm
stout,
velvety 4 in
stem 10 cm
thick, fleshy
scales 11/2–51/2 in
4–14 cm

BITTER TOOTH BLACK TOOTH cage bursts

Hydnellum scabosum Phellodon niger from “egg”
f: Bankeraceae f: Bankeraceae
Found in mixed woods in Eurasia and Growing in mixed woods in
blue-green N. America, this rare species has a centrally Eurasia and N. America, this
stem base
depressed cap with irregular scales. The uncommon species smells
spines beneath are pale buff. of fenugreek when dry. The RED CAGE FUNGUS
irregularly shaped cap is gray Clathrus ruber
to purplish black with gray 1–51/2 in f: Clathraceae
2.5–14 cm Found in parks and gardens in Eurasia,
ragged spines underneath.
margin this rare species has a red cage with
black, foul-smelling spores. The cage
hatches from a small, pale “egg.”
11/4–6 in 11/2–4 in
3–15 cm 4–10 cm
EARTHFAN
DEVIL’S TOOTH Thelephora terrestris
Hydnellum peckii f: Thelephoraceae DEVIL’S FINGERS
f: Bankeraceae Quite common in woods or on Clathrus archeri
This locally common species heaths in Eurasia and N. f: Clathraceae
from conifer woods in Eurasia America, this fungus grows on Introduced from Australasia, this
and N. America has a flattened, soil or woody debris. Its species is found mainly in S. Eurasia,
knobbly, woolly cap that often fan-shaped fruit bodies overlap, where it remains rare. Its red arms
exudes blood-red droplets. The forming clumps with paler, emerge from a white “egg” and have
underside has pale brown spines. fringed margins. blackish spores that smell fetid.
E XOBA SI DI A L E S gall on
leaf PUCCINIALES
This small group consists mainly of One of the largest orders of fungi with over 7,000 species,
237
gall-forming plant parasites whose the rust fungi include numerous serious parasites of crop plants.
spore-producing cells form a layer on EXOBASIDIUM VACCINII They can have very complex life cycles with multiple hosts and

MUSHROOMS • PUCCINIALES
the leaf surface. Some cause disease f: Exobasidiaceae produce different types of spores at different stages in their life.
in cultivated plants of the Vaccinium 3/8–3/4 in
Found in Eurasia and N. America, this
genus, which includes the 1–2 cm common fungus infects cowberry and black, powdery spores
eventually turns the leaves bright red. The in yellow spots on
common blueberry. leaves may become distorted and gall-like. lower leaf surface
RASPBERRY
black, powdery YELLOW RUST
U ROC YST I DI A L E S 1⁄16–5⁄32 in spores on Phragmidium rubi-idaei
2–4 mm leaf surface f: Phragmidiaceae
This order contains some well-known smut fungi, in This rust from Eurasia and
particular species of the genus Urocystis. These are parasites N. America causes pustules
of flowering plants such as anemone, onions, wheat, and to form on the upper surface
of leaves. It survives winter
rye, and often cause serious injury to the host plant. with the help of black spores
on the underside of leaves.
ANEMONE SMUT
Urocystis anemones
f: Urocystidaceae
This smut fungus from Eurasia and N. America orange rust
forms dark brown, powdery, raised pustules on damaging
the leaves of anemones and some other plants. rose stem
ROSE RUST
Phragmidium tuberculatum
DOG STINKHORN f: Phragmidiaceae
foul-smelling This common rust is found in
spore mass Mutinus caninus
on cap f: Phallaceae N. America and Eurasia. It causes
This common stinkhorn is found in orange pustules on the undersides
mixed woods in Eurasia and N. America. of leaves and on distorted stems. The
Its tip, covered in greenish black spores, pustules turn black in late summer.
is joined to the spongy stem that emerges
from a white “egg.”
ALEXANDERS RUST
hollow, 3/8–43/4 in 2–8 in Puccinia smyrnii
spongy stem 1–12 cm 5–20 cm f: Pucciniaceae
Found across Eurasia, this
common rust fungus forms raised
plaques or warts on the leaves of
white skirt Alexanders (Smyrnium olusatrum).
drops down
from cap
yellow warts leaf surface
of rust fungus spotted with
rust fungus
raised
blister of
rust fungus
HOLLYHOCK RUST
Puccinia malvacearum
f: Pucciniaceae
This serious pest of hollyhocks is
found in Eurasia and N. America. It
covers the leaves with small pustules.
Older leaves die and fall off.

PUCCINIA ALLII
f: Pucciniaceae
Common on onions, garlic, and
PHALLUS MERULINUS leeks in Eurasia and N. America, this
f: Phallaceae species forms pustules on infected
leaves. These break open to release
This tropical species is mainly found dustlike airborne spores.
in Australasia. It hatches from a white
“egg.” There are many similar species,
some with brightly colored “skirts.” rounded bright powdery
orange-yellow pustules pustules on leaf
large, erupt through leaf surface
white “egg”

2–8 in
5–20 cm STINKHORN HYPERICUM RUST FUCHSIA RUST
Phallus impudicus Melampsora hypericorum Pucciniastrum epilobii
f: Phallaceae f: Melampsoraceae f: Pucciniastraceae
Common in mixed woods, this fungus This common species A parasite of fuchsias and
is found in Eurasia. Its spore-covered, from Eurasia is visible as fireweed in Eurasia, this
honeycomblike cap hatches in a few scattered, raised pustules species infects the leaves
hours from an “egg.” Its foul odor is on the undersides of and forms pustules on the
often evident many yards away. Hypericum leaves. undersides of the leaves.
238 SAC F U N G I
The Ascomycota, or sac fungi, are fungi that produce their spores in
FUNGI • SAC FUNGI

rounded to elongate sacs called asci located on the fertile surface of

the fruit body. They are the largest group of fungi, and include many
cup- or saucer-shaped species.

PHYLUM ASCOMYCOTA Ranging in size from the microscopic to about

CLASSES 7 8 in (20 cm) in height, the sac fungi occupy a wide
ORDERS 56 variety of habitats, growing on dead, dying, and
FAMILIES 226
SPECIES About 33,000
living tissue, and floating in fresh and saltwater
environments. Many species are parasitic, and include
some of the most serious crop pests. Others form
mutually beneficial relationships, called mycorrhizae,
with plants. Sac fungi include some of the most
important fungi in the history of medicine, such as
the source of penicillin, while others are serious
pathogens—Pneumocystis jirovecii, for instance, can
cause lung infections in people with a weak immune
system. This phylum also contains the yeasts, which,
Many sac fungi display vivid colors, being vital components in the production of alcohol
although it is uncertain what biological and bread, have played a pivotal role in human history.
function such bright colors may have.
The Ascomycota display a wide range of fruit-body
shapes, including cup-shaped, club-shaped, potato-
like, simple crusts or sheets, pimplelike, corallike,
shield-shaped, or stalked with a spongelike cap.
Depending on the type of fruit body, the spore-
producing asci may grow externally on a special
fertile layer or be contained internally. Not all species
have a sexual stage; in fact, many use asexual methods
Spore-producing asci are shown here of reproduction. Most yeasts grow and rapidly
under a microscope. Arranged in dense
layers, the asci contain eight spores each. colonize new areas as a result of asexual division,
or budding, where a small bud forms on the outside
of a yeast cell, then separates, becoming a new cell.

th e cu p fu ngus
The common name of cup fungus refers to one of the
most conspicuous shapes of fruitbody associated with
the Ascomycota. Its open top (which is sometimes
disk- or saucer-shaped) allows wind and rain to scatter
Many species form their asci in special
protective chambers called perithecia; the spores that line its inner surface. In some varieties,
from here they discharge their spores. water is absorbed by the asci, which contain the spores,
so that pressure builds up and forcibly ejects the
D E B AT E spores up to 12 in (30 cm) from the fruit body. Close
HEROES OR VILLAINS? examination of surfaces such as rotting logs, fallen
branches, or leaves may reveal a fascinating world
Sac fungi form beneficial associations
with plants, algae, and even arthropods
of almost microscopic cups. In the larger cup species,
such as beetles. But equally they disturbance of the cup can produce such a vigorous
include some of the world’s most ejection of spores that their release is not only visible
damaging pathogens: Cryphonectria, for as a faint cloud of spores, but also audible.
example, is responsible for the recent
death of millions of chestnut trees.
Perhaps no other group of fungus has ORANGE PEEL CUP
such contrasting effects on our world. The orange peel fungus is a good example of the simple
cup-shaped fruit body adopted by many fungi species.
 HYPOCREALES XYLARIALES
240 The fungi in this order are usually distinguished by Members of this order often have their spore-producing
¾–3¼ in
2–8 cm
their brightly colored, spore-producing structures. cells in chambers, which are embedded in a woody
These are usually yellow, orange, or red. The Hypocreales growth called stroma. Although many species live
SAC FUNGI • HYPOCREALES

are often parasitic on other fungi and also on insects. on wood, some also occur on animal dung,
The best known among them is the genus Cordyceps fruit, leaves, and soil, or are associated
with club- or branchlike fruit bodies (bodies with insects. The order includes
that support the spore-forming cells). many economically important
Some species have medicinal uses. plant parasites.
tips covered in
powdery spores
1¼–2¼ in 2–5 in 3/8–1½ in
3–6 cm 5–13 cm 1–4 cm

false truffle 3/8–½ in

parasitized 1–1.5 cm
by truffleclub CANDLESNUFF DEAD MAN’S
FUNGUS FINGERS
SCARLET Xylaria hypoxylon Xylaria polymorpha
CATERPILLAR CLUB SNAKETONGUE f: Xylariaceae f: Xylariaceae
Cordyceps militaris TRUFFLECLUB Common on dead Growing on dead wood
f: Cordycipitaceae Tolypocladium ophioglossoides wood in Eurasia and in Eurasia and N. America,
Found in Eurasia and f: Ophiocordycipitaceae N. America, this species this fungus forms brittle
N. America, this fungus is Parasitic on buried false truffles, resembles a snuffed-out black clubs with a rough
parasitic on moth pupae. this species is found in Eurasia candle with a velvety surface, tiny pores,
The head of its club bears and N. America. It forms yellow black stem. and thick, white flesh.
tiny spore-producing clubs with an elongated, NAIL FUNGUS
structures. greenish black heads. Poronia punctata stemless fruit body has
f: Xylariaceae hard, brittle surface
purple infected Found on horse dung in Eurasia
rounded, ergot on bolete and N. America, this species is
cushionlike flower heads fruit body in decline. Its flattened disk dead trunk ¾–4 in
of ash tree 2–10 cm
fruitbodies has numerous tiny holes from
which spores are released.

¼–3/8 in
0.5–1 cm

½ in
1.5 cm

8–12 in
20–30 cm

CORAL SPOT ERGOT BOLETE EATER

Nectria cinnabarina Claviceps purpurea Hypomyces
f: Nectriaceae f: Clavicipitaceae chrysospermus BEECH WOODWART
Abundant on damp wood in This species has caused f: Hypocreaceae Hypoxylon fragiforme CRAMP BALLS
Eurasia and N. America, this outbreaks of mass This common mold, f: Xylariaceae Daldinia concentrica
fungus forms pink “pimples” poisoning. Found in found in N. America and Occurring in clusters on beech logs, f: Xylariaceae
when sexually immature Eurasia and N. America, Eurasia on bolete, turns this European and N. American species Found in Eurasia and N. America, this fungus has rounded
and red-brown clusters it is parasitic on grass bright golden yellow, forms hard, rounded fruit bodies with fruit bodies that reveal concentric, whitish zones when cut
when mature. and cereal crops. with a fluffy texture. tiny spore-releasing chambers. in half. They eject black spores from their outer layer.

E RYS I P H A L E S mildew CAPNODIALES

patches
Parasitic on leaves and fruit of flowering plants, the Commonly called sooty molds, these sac
Erysiphales are powdery mildews. The hyphae fungi are frequently found on leaves. They
(filaments) of their mycelium (vegetative part feed on the honeydew excreted by insects
of the fungus) penetrate the cells of the host or on liquid exuded from the leaves. Some
plant and take up nutrients. affected cause skin problems in humans.
apple leaf

feltlike white mycelium

covers leaf surface APPLE POWDERY MILDEW CLADOSPORIUM
POWDERY MILDEW Golovinomyces cichoracearum CLADOSPORIOIDES
OAK POWDERY MILDEW Podosphaera leucotricha f: Erysiphaceae f: Davidiellaceae
Erysiphe alphitoides f: Erysiphaceae Found in Eurasia and This mold, common in
f: Erysiphaceae Common on apple leaves in N. America, this species Eurasia and N. America,
Growing on oaks in Eurasia Eurasia and N. America, this appears on members of grows on damp bathroom
and N. America, this fungus mildew appears first on the Asteraceae (the sunflower walls. It may cause an
covers young leaves, causing underside of leaves as whitish family), causing patches on allergic reaction in
them to shrivel and blacken. patches, before spreading rapidly. leaves, which eventually die. some people.
HELOTIALES BROWN CUP
Rutstroemia firma
The fungi of this order are distinguished by their disk- or f: Rutstroemiaceae 241
3⁄16–½ in Consisting of a light brown cup and a
cup-shaped fruit bodies, unlike some similar cup fungi. Their 0.5–1.5 cm narrow stem, this fungus grows on fallen
saclike spore-producing cells, or asci, do not have an apical branches, especially of oaks, in Europe.

SAC FUNGI • HELOTIALES

lid (a flap through which they open). Most members live on It turns the host wood black.
humus-rich soil, dead logs, and other organic matter. This order
also includes some of the most damaging plant parasites.

ROSE BLACK SPOT

Diplocarpon rosae
f: Dermateaceae
Commonly found on
rose leaves in Eurasia and
N. America, this fungus
causes black spots, which
merge together to cause
merged large black patches.
spore- black spots
producing spots
1¼–2¾ in
TROCHILA ILICINA spore-producing smooth surface 3–7 cm
f: Dermateaceae inner surface of cup
Abundant on fallen holly leaves,
this fungus is found in Eurasia and SCALY EARTHTONGUE
N. America. It forms spore-producing Geoglossum fallax
spots on the leaf surface. f: Geoglossaceae
Growing in meadows in Eurasia and
N. America, this uncommon fungus
is one of several species with blackish,
flattened clubs, which can only be 1⁄16–3/8 in
separated using a microscope. 0.2–1 cm

3⁄16–1½ in 3⁄16–1¼ in 3⁄16–1¼ in

0.5–4 cm 0.5–3 cm 0.5–3 cm
BLACK BULGAR
Bulgaria inquinans
f: Bulgariaceae
Found in Eurasia and N. America, this species has
a brown outer surface. Its spore-producing inner ANEMONE CUP BEECH JELLYDISK
surface is smooth, black, and rubbery. Dumontinia tuberosa Neobulgaria pura
f: Sclerotiniaceae f: Helotiaceae
Parasitic on the tubers of Frequently found on fallen beech logs in Eurasia, this
anemones, this common translucent jellydisk varies from pale pink to pale lilac.
buff-colored Eurasian species has a long, The disks are often distorted from crowding together.
pustules on
infected fruit black stem and a simple
brown cup.

APPLE BROWN ROT 1⁄32–1/8 in LEMON DISCO

Monilia fructigena 1–3 mm Bisporella citrina
f: Sclerotiniaceae f: Helotiaceae
Very common in Eurasia, this fungus occurs Very common on dead hardwood in Eurasia, this
primarily on apples and pears, although it is species occurs in clusters. Its golden-yellow
also found on Prunus sp., causing brown fruit rot. disks sometimes cover entire branches.

1/8–3/8 in 1⁄16–3/8 in 3⁄16–¾ in

0.3–1 cm 0.2–1 cm 0.5–2 cm

JELLY
BABIES
Leotia lubrica
f: Leotiaceae GREEN ELFCUP LARGE PURPLE DROP
Frequently found in Chlorociboria aeruginascens Ascocoryne cylichnium BOG BEACON
mixed woodland in f: Helotiaceae f:Helotiaceae Mitrula paludosa
Eurasia and N. America, Found in Eurasia and N. America, Quite common on fallen beech logs in f: Helotiaceae
this species has a lobed the blue-green mature cups are rather Eurasia, this species attaches itself centrally This species, found in Eurasia and N. America, grows
head with a margin that uncommon, but the green stains they to the wood and forms gelatinous, irregular on plant remains in shallow water in spring and early
rolls back on itself. cause on fallen oak logs are easily visible. disks when sexually mature. summer. It has a rounded to tongue-shaped head.
 PEZIZALES 3⁄16 –¾ in
0.5–2 cm
tall 3⁄16 –3/8 in
0.5–1 cm
242 Members of this order produce
orange cups

spores inside saclike structures,

or asci, which typically open by
rupturing to form an operculum 2–4 in
(terminal lid) and eject the 5–10 cm 3⁄16 –½ in
0.5–1.5 cm
spores. The order includes a
number of species of economic COMMON EARTHCUP HARE’S EAR COMMON EYELASH TOOTHED CUP
importance, such as morels, Geopora arenicola Otidea onotica Scutellinia scutellata Tarzetta cupularis
truffles, and desert truffles. f: Pyronemataceae f: Pyronemataceae f: Pyronemataceae f: Pyronemataceae
Common in Eurasia, this This common species is This is one of many A common species in
fungus is difficult to spot as it often found in clusters similar species; its fruit body alkaline soils in woods of
lies buried in sandy soils. It has in broadleaved forests in is a scarlet cup with a fringe Eurasia and N. America,
dark brown, a smooth, spore-producing Eurasia and N. America. of black hairs. This fungus is its gobletlike cup has
wrinkled cap inner surface. Its tall cups are split common on wet, rotten wood a short stem.
down on one side. in Eurasia and N. America.
irregular ridges
and pits

ORANGE PEEL FUNGUS

Aleuria aurantia
f: Pyronemataceae
Often found along
gravelly dirt tracks in
Eurasia and N. America,
this is an unmistakable
species, with thin,
orange caps.
¾–4 in
2–10 cm

1½–4 in
4–10 cm

BLEACH CUP
FALSE MOREL Disciotis venosa
Gyromitra esculenta f: Morchellaceae
2–6 in f: Discinaceae Growing in spring in damp
5–15 cm This poisonous species is found woodlands in Eurasia and
throughout N. America and N. America, this short-stemmed
Eurasia, and usually grows under species has a chlorinelike smell. Its
conifers in spring. The shiny brown inner surface is brown and wrinkled
cap looks like a wrinkled brain. and the outer surface is pale.

HALF-FREE MOREL
Morchella semilibera
f: Morchellaceae 2–6 in 2–8 in
This hollow morel looks like 5–15 cm 5–20 cm
a dark, ridged thimble on a
scurfy, pale stem. It is common
smooth cap surface in spring in mixed woods in
Eurasia and N. America. hollow stem
THIMBLE MOREL
Verpa conica BLACK MOREL MOREL
f: Morchellaceae Morchella elata Morchella esculenta
An uncommon species, this fungus
2–4 in 2–6 in
f: Morchellaceae f: Morchellaceae
grows in woods and hedges on Common in woods This prized species grows
chalky soils in Eurasia and 5–10 cm 5–15 cm
during spring in Eurasia and in spring in calcareous
N. America. Its smooth, thimble- N. America, this species has a woodland in N. America
like cap sits on a hollow stem. pinkish buff to black cap, with and Eurasia. It has a
cross-connected black ridges, spongelike, hollow cap
and a hollow stem. and a hollow stem.

½–2¾ in
1.5–7 cm

1¼–4 in 1–3 in
3–10 cm 2.5–7.5 cm

BLISTERED CUP CELLAR CUP BAY CUP

Peziza vesiculosa Peziza cerea Peziza badia
f: Pezizaceae f: Pezizaceae f: Pezizaceae
Common in Eurasia and Found across Eurasia and One of many similar species, this
N. America, this fungus typically N. America, this fungus often common fungus of Eurasian and
occurs in clusters on compost, occurs on damp brickwork. N. American woods has a spore-
straw, or manure. Its brittle It is dark ocher on the inside producing inner surface. Its brown
cups can have ragged edges. and pale outside. cup develops an olive tint with age.
 ¾–2¾ in
2–7 cm
¾–3¼ in
2–8 cm
E U ROT I A L E S R H Y T I SM ATA L E S
Better known for the blue and green molds, Commonly called tar spot fungi, the species in this
243
this order includes the fungi Penicillium (famous order infect plant matter, such as leaves, twigs, bark,
for producing penicillin, the first discovered female conifer cones, and occasionally berries. Many

SAC FUNGI • PLEOSPOR ALES

antibiotic) and Aspergillus (a significant cause species attack the needles of conifers, causing needle
of disease in humans). drop. The tar spot of maple leaves
is perhaps the most frequently seen.
PERIGORD WHITE TRUFFLE powdery
TRUFFLE Tuber magnatum interior brown spots on
Tuber melanosporum f: Tuberaceae upper leaf surface
f: Tuberaceae Prized in Italy and
This highly prized truffle France, this expensive
grows underground truffle of alkaline soils 1–2 in
around oaks in the in S. Europe can be 2.5–5 cm
Mediterranean region. cultivated on suitable host
The truffles are found trees such as oak and
using dogs or pigs. poplar, by inoculation.

3/8–¾ in
SUMMER TRUFFLE 1–2 cm
Tuber aestivum ½–1¾ in TAR SPOT YELLOW FAN
f: Tuberaceae 1.5–4.5 cm Rhytisma acerinum Spathularia flavida
Found across FALSE TRUFFLE f: Rhytismataceae f: Cudoniaceae
S. and C. Europe, this Elaphomyces granulatus Abundant on acer or maple trees This common species grows
highly valued truffle f: Elaphomycetaceae in N. America and Eurasia, this in wet, mossy conifer woods
grows underground, Common in sandy soils below conifers in Eurasia and fungus causes irregular spots, of Eurasia and N. America. It
near a variety of N. America, this reddish-brown truffle has a roughened with paler yellow margins, has flattened, pale to darker
broadleaved trees. surface and purple-black inner spore-mass. which disfigure the leaves. yellow, rubbery heads.
¾–2 in/2–5 cm

TA PH R I NA L E S PEACH LEAF CURL

Taphrina deformans
5½–16 in
14–40 cm
This order contains many plant parasites, with most f: Taphrinaceae
This fungus infects most
species in the genus Taphrina. All species have two varieties of peach and nectarine
growth states: in the saprophytic state, they are trees in Eurasia and N. America.
yeastlike and propagate by budding; but in the The affected leaves curl and
parasitic state, they emerge through crinkle, and frequently
turn reddish purple.
plant tissues, causing distorted
leaves and galls.

3/8–3¼ in BIRCH BESOM

1–8 cm Taphrina betulina
f: Taphrinaceae
SCARLET ELFCUP Found across Eurasia, this 8–37 in
Sarcoscypha austriaca common species causes 20–95 cm
f: Sarcoscyphaceae witches’ broom, a disease
Found from winter to early spring on fallen branches in of birch trees that causes red patches
Eurasia and N. America, this species has a scarlet cup clumps of narrow twigs to caused by
contrasting with its pale outer surface. grow at branch ends. infection

¾–2¼ in
2–6 cm
PLEOSPORALES PEAR SCAB
Venturia pyrina
Typical members of this order develop f: Venturiaceae
Found in pear orchards in
their asci within a flasklike fruiting Eurasia and N. America, this
body. The asci have two wall layers: at common parasite causes the fruit
maturity, the inner wall protrudes beyond to distort, discolor, and even drop
the outer wall, ejecting the spores. Many before it is ripe.
species grow on plants; some form lichens.

2–6 in
5–15 cm dark,
dark cones sunken
spots

LEPTOSPHAERIA
ACUTA
WHITE SADDLE ELFIN SADDLE f: Leptosphaeriaceae BOEREMIA
Helvella crispa Helvella lacunosa Found in Eurasia and HEDERICOLA
f: Helvellaceae f: Helvellaceae N. America, this common f: Didymellaceae
Possibly poisonous, this Found in mixed woods in species infects the dead Found in Eurasia and
species is common in Eurasia and N. America, stems of the stinging nettle, N. America, this species
mixed woods in Eurasia this common species Urtica dioica. It forms tiny causes circular white
and N. America. Its thin, has a lobed, dark cap cones that push through the lesions on the leaves
saddlelike cap sits on on a gray, fluted, and stem surface of the host in of the ivy plant, which
a fragile, ribbed stem. columned stem. order to eject its spores. turn brown and die.
244 LICHENS
From outcrops of rock exposed to the sea to desert areas where
FUNGI • LICHENS

the only place they can grow is inside the rock itself, lichens
survive in the harshest parts of the world. They are some of nature’s
pioneers, creating foundations on which others can live.

PHYLA ASCOMYCOTA A lichen is not a single organism, but a composite

BASIDIOMYCOTA consisting of a green alga or cyanobacterium and
CLASSES 10
ORDERS 15
a fungus, living together in a mutually beneficial
FAMILIES 40 association. The algal partner provides nutrients
SPECIES About 18,000 through photosynthesis, while the fungal partner aids
the alga by retaining water and capturing mineral
nutrients. The fungus is normally a member of the
Ascomycota (the sac fungi), or, more rarely, of
the Basidiomycota (mushrooms)—lichen classification
reflects the type of fungus involved. Typically, the
fungus surrounds the photosynthetic cells of the alga,
enclosing them within special fungus tissues unique
to lichens. It appears that although neither partner
Asexual soredia are bundles of fungal is capable of surviving on its own, together they can
hyphae (filaments) and algal cells. Here endure the most extreme conditions. Lichens have even
they are budding off and awaiting dispersal.
been found some 250 miles (400 km) from the South
Pole, but they also grow in familiar places, such as
dry-stone walls, rocks, and bark.
Lichens broadly fall into three types, based on
shape: foliose lichens, which have leaves; crustose
lichens, which form a crust; and fruticose lichens,
which have branches. However, there are some that
defy this categorization, such as filamentous (hairlike)
Asexual isidia are tiny peglike cells on and gelatinous (water-absorbent) lichens.
the edge of the lichen, which break off to
form new colonies of lichen.
r e p ro d u c t i o n
Many lichens reproduce sexually by means of spores.
These are produced by the fungal half of the partnership
and are usually formed in special cuplike or disklike
structures called apothecia. These spores, once ejected,
must land next to a suitable algal partner if they are to
form another lichen and survive. Other lichens produce
spores inside special chambers called perithecia, which
Lichen cells are shown here in
microscopic cross-section. Spore- are like microscopic volcanoes, releasing their spores
producing asci rise from algal cells. through a hole at the top. Alternatively, lichens may
reproduce asexually by budding or breaking off
D E B AT E specialized parts of their body. These soredia or isidia
U N L I K E LY P A R T N E R S ? contain a mix of fungal and algal cells, which, if they
land in a suitable habitat, will go on to form new lichen
Lichen evolution remains a mystery.
Scientists are still trying to understand
colonies. Rocky shorelines in North America are home
how and why fungi and algae came to to vast colonies of lichens several miles in length.
live together. It may have started as These will have taken hundreds or even thousands
an attack on one by the other, which of years to spread to such a huge size.
developed into a partnership. Not all
lichen associations are necessarily
mutually beneficial—some may be SPREADING MAP LICHEN
parasitic rather than a partnership. Typical of lichens, Rhizocarpon is able to colonize harsh
environments such as dry, exposed rock surfaces.
 LICHENS 1–3 in 1–3 in
2.5–7.5 cm 2.5–7.5 cm
246
FUNGI • LICHENS

rounded lobes
that lift at edges

COMMON
2–4 in GOLDEN-EYE LICHEN WALL LICHEN
5–10 cm Teloschistes chrysophthalmus Xanthoria parientina
f: Teloschistaceae f: Teloschistaceae
ORANGE LICHEN A critically endangered species in Growing on trees, walls,
Caloplaca verruculifera America, Eurasia, and the tropics, this and roofs in N. America,
f: Teloschistaceae lichen grows on shrubs and small trees in Eurasia, Africa, and Australia,
This lichen has radiating lobes with apothecia old orchards and hedgerows. Its branched this lichen has rounded,
(spore-producing disks) at the center. It is lobes produce large, orange disks. yellow-orange lobes.
found on rocks in coastal areas in N. America
and Eurasia, often near bird perches.
1–3 in
2.5–7.5 cm
2–6 in/5–15 cm

1–3 in
REINDEER MOSS HAMMERED SHIELD LICHEN 2.5–7.5 cm
Flavocetraria nivalis Parmelia sulcata
f: Parmeliaceae f: Parmeliaceae
Found on mountain heaths and upland This lichen’s flattened lobes are gray-green, 1–3 in HOARY ROSETTE
moors in N. America and Eurasia, this with rounded tips and powdery, reproductive 2.5–7.5 cm LICHEN
species has brownish, flattened leafy structures on the surface. It is commonly
Physcia aipolia
fronds with spiny margins. found on trees in N. America and Eurasia.
f: Physciaceae
Growing on tree bark in the
Americas and Eurasia, this gray
1–3 in 1–3 in to brownish-gray species forms
2.5–7.5 cm 2.5–7.5 cm rough patches with a lobed
margin. It has black apothecia.

3/8–2 in
1–5 cm

DEVIL’S MATCHSTICK
Cladonia floerkeana
f: Cladoniaceae
This species is common on peaty soil
in N. America and Eurasia. It forms a
BEARD LICHEN POWDER-HEADED HOODED crust of greenish gray scales from
Usnea filipendula TUBE-LICHEN TUBE-LICHEN which stalks emerge, tipped with
f: Parmeliaceae Hypogymnia tubulosa Hypogymnia physodes scarlet apothecia.
Found mainly in northern regions, f: Parmeliaceae f: Parmeliaceae
this species forms green-gray, Common on twigs and tree trunks, Found worldwide on trees, rocks, and
pendent clumps on trees. Spiny this lichen has lobes that are gray-green walls, this lichen has pale gray-green lobes
apothecia form at the tips. above and dark underneath. It is found with wavy margins. The rare apothecia are
in N. America and Eurasia. red-brown with gray margins.

1–4 in 1–4 in/2.5–10 cm

2.5–10 cm

1–5 in 1–4 in
2.5–12.5 cm 2.5–10 cm

CARTILAGE CORAL LICHEN 1–4 in

LICHEN Sphaerophorus globosus BLACK SHIELDS 2.5–10 cm
STONEWALL
Ramalina fraxinea f: Sphaerophoraceae Tephromela atra RIM-LICHEN REINDEER LICHEN
f: Ramalinaceae Found on rocks in mountainous f: Mycoblastaceae Lecanora muralis Cladonia portentosa
Growing on trees in areas of N. Eurasia and This lichen has pale gray crustose f: Lecanoraceae f: Cladoniaceae
N. America and Eurasia, N. America, this fungus forms lobes that look like dried porridge. This lichen often grows on concrete One of several reindeer lichens,
this species forms flattened, dense cushions of pinkish Its apothecia are black. The lichen and on rocks. Its gray-green lobes this species is common on heath and
gray-green branches brown branches with is found on exposed rocks in radiate outward. It is found in moorland in N. America and Eurasia. Its
dotted with apothecia. globular apothecia. N. America and Eurasia. N. America and Eurasia. thin, hollow branches repeatedly divide.
 247
1–2 in
2.5–5 cm

2–20 in
5–50 cm

BLACK TAR LICHEN DOG LICHEN 8–12 in BLISTERED JELLY LICHEN

Verrucaria maura Peltigera praetextata 20–30 cm Collema furfuraceum
f: Verrucariaceae f: Peltigeraceae f: Collemataceae
Occurring on rocks along seashores Found on rocks in Eurasia and This lichen has flat, gelatinous,
in N. America and Eurasia, this N. America, this lichen has large wrinkled lobes. It is found on rocks
species has a dark gray, cracked gray-black lobes, with paler margins and trees in areas with high rainfall
crust that contains apothecia. and reddish brown apothecia. in N. America and Eurasia.

green lobes spread

out from center

2–4 in 2–6 in
5–10 cm 5–15 cm
TREE LUNGWORT
Lobaria pulmonaria
f: Lobariaceae
Mainly found on tree bark in coastal areas in
black, slitlike N. America, Eurasia, and Africa, this species
2–8 in 1–3 in openings is declining due to habitat loss.The branching
5–20 cm 2.5–7.5 cm release spores lobes are pale orange underneath.
ROCK TRIPE PETALED ROCK-TRIPE COMMON
Lasallia pustulata Umbilicaria polyphylla SCRIPT LICHEN
f: Umbilicariaceae f: Umbilicariaceae Graphis scripta 1–4 in
This N. American and Eurasian A common species on rocks f: Graphidaceae 2.5–10 cm
lichen forms colonies on nutrient- in mountains in N. America and Often seen on tree bark in
rich rocks in coastal or upland areas. Eurasia, it has smooth, broad N. America and Eurasia, this
It has a gray-brown upper surface lobes that are dark brown above lichen forms a thin, gray-green
with many oval pustules. and black beneath. crust with slitlike openings.

2–8 in 1–3 in
5–20 cm 2.5–7.5 cm

PERTUSARIA PERTUSA LECIDEA LICHEN

f: Pertusariaceae Lecidea fuscoatra
Common on tree bark f: Lecideaceae
dome-shaped in N. America and Eurasia, This species is common on
apothecia this lichen forms gray crusts siliceous rocks and old brick
with a pale margin. The crusts walls in N. America and Eurasia.
are covered with groups of It forms a gray, cracked crust,
CRAB-EYE LICHEN warts with tiny openings. with sunken, black apothecia.
Ochrolechia parella
f: Ochrolechiaceae
This lichen forms patches on walls and
rocks in N. America and Eurasia. Its surface
usually has many pink-brown apothecia.
MAP LICHEN 3⁄16 –2½ in
Rhizocarpon geographicum 5–65 mm
f: Rhizocarpaceae
Common on rocks in
mountains in northern regions
and Antarctica, this lichen
forms a flat patch bordered
by a black line of spores.
Groups of these lichens have
a patchwork appearance.

1–5 in
2.5–12.5 cm

BAEOMYCES
RUFUS
f: Baeomycetaceae
Forming gray-green crusts
on sandy soil and rocks,
this lichen has brown ball-like
apothecia on stalks a few
millimeters high. It is found
in N. America and Eurasia.
anima
ls
Animals make up the largest kingdom of
living things. Driven by the need to eat food, and to
escape being eaten, they are uniquely responsive
to the world around them. The vast majority are
invertebrates, but mammals and other chordates
often outclass them in size, strength, and speed.

250 322
INVERTEBRATES CHORDATES
Hugely varied in shape, Most of the world’s large
invertebrates also have animals are chordates.
a wide range of lifestyles. Externally, they can be
Insects make up the covered in fur, feathers, or
largest group, but others overlapping scales, but
include jellyfish, worms, internally nearly all have a
and animals protected backbone, which makes up
by a hard shell. part of a bony skeleton.
250
I N V E RT E B R AT E S
With about 1.4 million species identified, animals make
A NIM ALS • IN VERTEBR ATES

up the largest kingdom of living things on Earth. The vast

majority of them are invertebrates—animals without a
backbone. Invertebrates are extraordinarily varied; many
are microscopic, but the largest are over 33 ft (10 m) long.
Invertebrates were the first animals to be remarkably successful, allowing
to evolve. Initially, they were small, arthropods to invade every natural
soft-bodied, and aquatic—features habitat in water, on land, and in the air.
that many living invertebrates still Invertebrates also include animals that
share. During the Cambrian Period, have a shell, and ones that are reinforced
which ended about 485 million years by mineral crystals or hard plates. Only
ago, invertebrates underwent a vertebrates, however, have an internal
spectacular burst of evolution, skeleton made of bone.
developing a huge range of body forms
and some very different ways of life. spli t li v es
This evolutionary explosion produced Most invertebrates begin life as an egg.
almost all the major phyla, or groups, of Some look like miniature versions of
invertebrates that exist today. their parents when they hatch, but SPONGES
Among the simplest animals, sponges have a sievelike body,
many start life with a very different with an internal skeleton of mineral crystals. Classified as
g r e at d i v e r s i t y body form. These larvae change shape, the phylum Porifera, there are more than 9,000 species.
There is no such thing as a typical food sources, and feeding methods as ARTHROPODS
invertebrate, and many of the phyla they grow up. For example, sea urchins The phylum Arthropoda is the biggest in the animal kingdom,
with more than 1.2 million species identified. It includes
have very little in common. The have drifting larvae, which filter food insects, crustaceans, arachnids, centipedes, and millipedes.
simplest kinds do not have a head or from the sea, while the adults scrape
a brain, and usually rely on internal algae from rocks. The change in shape—
fluid pressure to keep their shape. At metamorphosis—can be gradual, or it
the other extreme, arthropods have can take place abruptly, with the young
a well-developed nervous system and animal’s body being broken down,
elaborate sense organs, including and an adult body being assembled
complex eyes. Crucially, they also have in its place. Metamorphosis enables
an external body case, or exoskeleton, invertebrates to exploit more than one
with legs that bend at flexible joints. food source, and it also helps them to
This particular body plan has proved spread, often over great distances.

INVERTEBRATE MOLLUSKS ARTHROPODS

TREE RIBBON SEGMENTED
WORMS
WORMS
FLATWORMS WATER VELVET
LAMPSHELLS BEARS WORMS

CNIDARIANS
BRYOZOANS ECHINODERMS ROUNDWORMS
SPONGES Invertebrates are not a true taxonomic
group. They range from simple sponges
INVERTEBRATES to insects—the most successful animals
continues to chordates on Earth in terms of numbers.
CNIDARIANS FLATWORMS SEGMENTED WORMS
Members of the phylum Cnidaria are soft-bodied Numbering about 30,000 species, the phylum With about 18,000 species, the phylum Annelida contains
invertebrates that kill their prey with stinging cells. Of Platyhelminthes contains animals with a flat, worms that have a sinuous body, which is divided into
the 11,947 known species, the vast majority are marine. thin body, and a distinguishable head and tail. ringlike segments. It includes earthworms and leeches.

CRUSTACEANS MOLLUSKS ECHINODERMS

Mainly aquatic, crustaceans are arthropods that breathe One of the most diverse invertebrate groups, the phylum Recognizable by their five-fold symmetry, members of the
with gills. Classified as the subphylum Crustacea, there Mollusca contains almost 72,000 species. It includes phylum Echinodermata have a skeleton of small, chalky
are about 70,000 species, including crabs and lobsters. gastropods, bivalves, and cephalopods. plates set in their skin. There are about 7,450 species.
252 SPONGES
Simple, mostly marine animals, adult tiny pores in the sponge’s surface. Special collar cells PHYLUM PORIFERA
sponges live permanently attached to lining the water canals trap and engulf bacteria and
INVERTEBRATES • SPONGES

CLASSES 4

rocks, corals, and shipwrecks. A few other small plankton for food, and waste water exits ORDERS 32

species live in fresh water. through large openings called oscula. FAMILIES 144

Many sponges are classically “spongy,” others SPECIES More than 9,000

The Porifera vary in size and shape, from thin are rock hard, soft, or even slimy depending on
sheets to huge barrels, but all have the same basic the supporting skeleton, which is made up of tiny
structure: different types of specialist cells, but no spicules of silica or calcium carbonate. Spicules
organs. A system of water canals branches through vary in shape and number between species and
the sponge, and water is drawn into them through are an important feature in identification.

CA LCA REOUS SPONGES DEMOSPONGES

The skeleton of Calcarea is composed of densely packed, mostly More than 85 percent of all sponges belong to
the Demospongiae. Although they are varied 3 1/4 ft
starlike, calcium carbonate spicules, each with three or four pointed 1m
rays. Variable in shape and crunchy to the touch, most species are small in appearance, most have a skeleton made up
of scattered silicon dioxide spicules and a BLUE SPONGE
and lobe- or tube-shaped. Haliclona sp.
flexible organic collagen called spongin. f: Chalinidae
A few encrusting species have no skeleton One of very few blue-colored
and some have only spongin. sponges, this species commonly
grows on the tops of corals and
rocks off N. Borneo.
3 1/4 ft
1m

31/4 in 3/8–11/2 in
8 cm 1–4 cm
LEMON SPONGE
Leucetta chagosensis 14 in
f: Leucettidae CLATHRINA CLATHRUS 35 cm
Growing on steep coral reefs in the f: Clathrinidae
W. Pacific, this saclike sponge provides Made up of many tubes, MEDITERRANAEAN BATH
a splash of bright color. each only a few millimeters BROWN TUBE SPONGE SPONGE
wide, this Mediterranean Agelas tubulata Spongia (Spongia) officinalis
sponge is a distinctive f: Agelasidae f: Spongiidae
yellow color. This sponge consists of uneven This sponge has an elastic skeleton and no
brown tubes arranged in clusters. It harsh scpicules, so after it has been cleaned
spicules is common on deep reefs in the and dried, it is ideal to use for washing.
surrounding Caribbean and Bahamas.
osculum GOLF BALL
1 3/4–2 in 2–4 in SPONGE
2–5 cm 5–10 cm Paratetilla bacca
f: Tetillidae
PURSE SPONGE One of many tropical
Sycon ciliatum ball-shaped sponges, this
f: Sycettidae species grows on sheltered
This simple, hollow sponge coral reefs in the
of N.E. Atlantic coasts has W. Pacific.
spiky, calcareous spicules
surrounding its osculum. 43/4 in
12 cm

4 in 3 1/4 in ELEPHANT HIDE SPONGE

tiny pores allow
10 cm 8 cm Pachymatisma johnstonia
water entry
f: Geodiidae
Mounds of this tough sponge can
cover large areas of rocks and
wrecks in clear coastal waters
of the N.E. Atlantic.

12–16 in
30–40 cm
LEUCONIA SP. RED PURSE SPONGE
f: Baeriidae Grantessa sp. RED TREE SPONGE
The shape of this f: Heteropiidae Negombata magnifica
N.E. Atlantic sponge Shaped like a small gourd, f: Podospongiidae
varies from lobes and this delicate sponge grows Attempts to farm this beautiful
cushions to crusts. between coral heads Red Sea sponge are meeting with
It grows in areas of high on shallow reefs in some success. It contains chemicals
water movement. Malaysia and Indonesia. of potential medical importance.
 20 in 6–12 in 12–16 in
50 cm 15–30 cm 30–40 cm
253

BORING SPONGE BORING SPONGE

Cliona celata Cliona delitrix
f: Clionaidae f: Clionaidae
Although it appears as yellow lumps, Water leaves this Caribbean sponge
much of this European sponge via a number of large oscula (one
remains hidden as strands growing is shown here). This sponge bores
through shells and calcareous rocks. through corals by secreting acid.

3/8 in/1 cm 3/8–3/4 in

1–2 cm

YELLOW FINGER
SPONGE
Callyspongia
(Callyspongia) nuda
f: Callyspongiidae
The bright color of this
tropical Pacific sponge is due
to the chemicals it contains.
Sponge extracts are used by
the pharmaceutical industry.
BREADCRUMB SPONGE SPIRASTRELLA
Halichondria panicea CUNCTATRIX
f: Halichondriidae f: Spirastrellidae
This N.E. Atlantic sponge grows One of many colorful encrusting sponges
as crusts on rocky shores and in found in the ocean, this species covers 21/2–61/2 ft 1 1/2 ft
shallow water. Its color comes inshore rocks in the Mediterranean 0.8–2 m 45 cm
from symbiotic algae. and North Atlantic.

BARREL SPONGE
Xestospongia testudinaria
f: Petrosiidae
Small fishes and many invertebrates
live on and in this huge Indo-Pacific AZURE VASE
sponge, which grows big enough to SPONGE
fit a person inside. Callyspongia plicifera
f: Callyspongiidae
A common Caribbean
APLYSINA ARCHERI species, this sponge colors
f: Aplysinidae reefs with its pale blue to
The graceful long tubes of this sponge purple vases. Its surface is
sway gently with the current on the sculpted with ridges and
61/2 ft/2 m Caribbean reefs where it grows. valleys.

H O M O S C L E RO M O R P H A S P O N G E S GLASS SPONGES
This small class of just under 130 sponges is found in both temperate and Hexactinellida are a small group of sponges that live mostly
tropical waters. They are mostly soft, as they have tiny silica spicules or in the deep ocean, where some form reeflike mounds up to 14 in
35 cm
none at all. Their planktonic larvae are 65 ft (20 m) high. The six rayed, starlike, silica spicules of
unique in form. the skeleton are usually fused together as a rigid lattice.
After death, the skeleton remains as a ghostly outline.
VENUS’S FLOWER
BASKET
8 in Euplectella aspergillum
CHICKEN LIVER SPONGE 20 cm
Plakortis lita f: Euplectellidae
f: Plakinidae Growing in tropical oceans
The common name of this BOLOSOMA below about 500 ft (150 m),
sponge reflects its look and feel. GLASS SPONGE this sponge was collected and
Dark brown inside and out and Bolosoma sp. displayed by Victorians for its
smooth to the touch, it is found
f: Euplectellidae delicate silica skeleton.
on coral reefs in the W. Pacific. Looking like fat fungi,
Bolosoma sponges live in
soft coral polyp the deep sea and are
held up into the water rigid lattice of
4–6 in silica spicules
10–15 cm currents by a long,
thin stalk.
254 C N I DA R I A N S
This phylum includes jellyfish, corals, this phylum. These stinging organs are called PHYLUM CNIDARIA
and anemones. They have stinging cnidocysts (or cnidocytes), and it is from these
INVERTEBRATES • CNIDARIANS

CLASSES 6

tentacles to catch live prey, which that cnidarians get their name. Cnidocysts are ORDERS 22

they digest in a simple saclike gut. concentrated on the tentacles, and are triggered FAMILIES 278

by touch or chemical signals, both when the animal SPECIES 11,947

All cnidarians are aquatic and most are marine. comes into contact with potential prey and when
They have two body plans: a free-swimming, it is under attack. Each cnidocyst contains a
bell-like shape called a medusa—as seen in microscopic sac of venom and discharges a tiny
jellyfish—and a static polyp, typical of anemones. coiled “harpoon” to inject the venom into flesh.
Neither medusae nor polyps have a head or front Some stings can penetrate human skin and inflict
end. Their tentacles encircle a single gut opening, severe pain, but the vast majority of cnidarians
used to take in food and to eliminate waste. are harmless to humans.
A cnidarian’s nervous system consists of a simple
network of fibers; there is no brain. This means a lt e r n at i n g l i f e c yc l e
that the animal’s behavior is usually simple. Many cnidarians have a life cycle that alternates
Although they are carnivorous, cnidarians cannot between medusa and polyp—usually with one
actively pursue their prey, with the possible of these forms dominating, though in some
exception of the box jellyfish. Instead, most groups one or other form is missing. The free-
species wait for swimming animals to blunder swimming medusa is normally the sexual stage.
into the reach of their tentacles. In most species fertilization is external: sperm
and eggs are released in open water, forming
stings planktonic planula larvae that resemble tiny
Cnidarians’ outer skins—and in some species the flatworms. These settle in order to grow into
inner skin as well—are peppered with tiny stinging polyps. Specialized polyps then produce new This microscopic view shows
cnidocysts (stinging cells) that have
capsules of a kind that is unique to animals of medusae to complete the cycle. been triggered to discharge their
venom-laden harpoons.

BOX STA L K E D T RU E J E L LY F I S H
J E L LY F I S H J E L LY F I S H The familiar jellyfish bell is the medusa stage
Members of the class Cubozoa are The adult is the only easily seen of the Scyphozoa life cycle. The polyp is
found in tropical and subtropical phase in the life cycle of members reduced, or in some deep-sea forms, is
waters. They differ from true jellyfish of the Class Staurozoa. Unlike other missing. Jellyfish polyps undergo strobilation,
in having a greater ability to control the jellyfish it does not swim, but is a process that produces new tiny medusae by
direction and speed of their movement, attached to the substrate by a stalk. budding. Rhizostome jellyfish lack a fringe of
rather than drifting with the currents. Eight arms radiate from the mouth, tentacles around the bell.
A flapping skirt at the bottom of the with clumps of tentacles at their
bell powers them at tips. The other life stages are the
considerable speed. extremely small planula larvae and
Box jellyfish have polyps (called stauropolyps).
eyes, set in 8–16 in 51/2–61/2 in
20–40 cm 14–16 cm
clusters on the
sides of their
transparent bell. SPOTTED LAGOON
They can see MOON JELLYFISH
enough to steer JELLYFISH Mastigias papua
Aurelia aurita f: Mastigiidae
past obstacles and f: Ulmaridae Like other rhizostome
to spot prey. This globally widespread genus jellyfish, this algae-containing
11/2 in has four long “arms” as well as species traps planktonic food
4 cm small marginal tentacles. It with mucus. It enters South
bell swarms to breed near coasts Pacific lagoons, including
and polyps settle in estuaries. those of oceanic islands.
1–10 ft
0.3–3 m 8–12 in
20–30 cm
UPSIDE-DOWN
JELLYFISH
SEA WASP STALKED JELLYFISH Cassiopea andromeda
Chironex fleckeri Haliclystus auricula f: Cassiopeidae
f: Chirodropidae f: Haliclystidae Superficially resembling an
This Indo-Pacific species— This species is found attached to anemone, this Indo-Pacific
the largest box jellyfish— algae and seagrasses in shallow rhizostome jellyfish lives at the
delivers a particularly painful cooler waters of the Atlantic and bottom of lagoons, mouth
sting and has caused Pacific, where it catches food using upward, with its bell pulsating to
human fatalities. stinging cells on the tentacles. circulate water.
H Y DROZOA NS 3/8 in
10 mm
3/8 in
10 mm
Most animals of the class Hydrozoa live as branching sea-dwelling
255
colonies forming miniature polyp-bearing forests. Some anchor
themselves to surfaces by creeping horizontal stems, called stolons.

INVERTEBRATES • CNIDARIANS
Colonies are supported by a horny translucent sheath and some of
the polyps produce sexual medusae. Freshwater hydra lack medusae:
instead, sex organs develop directly on solitary polyps.
11/4–2 in
colony of 3–5 cm
snailfur on shell 3/8 in
10 mm
COMMON OBELIA EIGHT-RIBBED CUP HYDROMEDUSA
Obelia geniculata HYDROMEDUSA Phialella quadrata
f: Campanulariidae Melicertum octocostatum f: Phialellidae
This is a worldwide hydrozoan, f: Melicertidae Like related Obelia, branching
abundant on intertidal seaweeds. This hydrozoan from the North colonies of this globally
1⁄16–1/8 in It grows in zig-zag colonies of Atlantic and North Pacific oceans widespread hydrozoan release
2–3 mm cup-encased polyps that sprout belongs to a family related to those free-living medusae, which
from creeping horizontal stems. with cup-encased polyps. It is reproduce sexually.
known mainly from medusae.
SNAILFUR BLUE BUTTONS
Hydractinia echinata Porpita porpita PORTUGUESE 16–23½ in
f: Hydractiniidae f: Porpitidae MAN OF WAR 4–6 cm
One of a family of colonial This colonial, jellyfishlike Physalia physalis
spiny hydrozoans, this N.E. Atlantic hydrozoan of tropical f: Physaliidae
species grows over the surface oceans is sometimes regarded Resembling a jellyfish,
of snail shells containing as a highly modified this is really an
hermit crabs. individual polyp. ocean-going colony of
hydrozoans. Stinging
2–4 in PURPLE LACE “CORAL” tentacles and specialized
5–10 cm HYDROID polyps are suspended
Distichopora violacea from a gas-filled float.
f: Stylasteridae
Like some other hydrozoans,
colonies of this Indo-Pacific lace FEATHER HYDROID
coral have polyps specialized for Aglaophenia cupressina
different functions, such as gas-filled f: Aglaopheniidae
feeding and stinging. float Feather hydroids and the
closely related sea ferns
belong to a group of
16 in 33–165 ft colonial hydrozoans with
40 cm 10–50 m cup-encased polyps. This
is an Indo-Pacific species.

1½–6½ ft
0.5–2 m

5
⁄32 –½ in
4–15 mm

HULA SKIRT
SIPHONPHORE
LION’S MANE JELLYFISH Physophora hydrostatica
Cyanea capillata f: Physophoridae
f: Cyaneidae This widespread free-floating
COMMON HYDRA
A large species of Arctic waters, this hydrozoan colony has a
jellyfish has many tentacles arranged smaller float than its relative, Hydra vulgaris
in dense groups. It has a powerful the Portuguese man of war, f: Hydridae
and carries prominent Lacking the medusa stage,
umbrellalike sting and its prey includes fishes. hydras have small freshwater
bell swimming bells.
polyps that can reproduce
4–8 in asexually by budding. This
10–20 cm is a globally widespread
cold-water species, shown
here in false color.

3/4–1 in
2–2.5 cm

4–8 in 16–20 in
10–20 cm 40–50 cm

FIRE “CORAL”
PINK-HEARTED HYDROID FRESHWATER
DEEP SEA CORONATE HYDROID Millepora sp. JELLYFISH
JELLYFISH Tubularia sp. f: Milleporidae Craspedacusta sowerbii
Periphylla periphylla f: Tubulariidae Like others of its family, this f: Olindiidae
f: Periphyllidae Tubularia hydrozoans have fiercely stinging colonial Sporadic in freshwater
This is one of many little-known polyps on long stems with two hydrozoan has a calcified ponds, lakes, and streams
deep-water species in a group of whorls of tentacles, one around skeleton and is a reef- worldwide, minute polyps
jellyfish characterized by a bell the polyp base and one builder. It is distantly related of this animal develop into
oral arm with an encircling groove. around its mouth. to true stony corals. dominant medusae.
 ANEMONES AND CORALS 31/4–5 ft
1–1.5 m
polyp cluster
256 Unlike other cnidarians, the Anthozoa lack the free-swimming COMMON
medusa stage altogether and most remain static throughout SEA FAN
their life cycle. The polyps, many of which have a flowerlike Gorgonia ventalina
f: Gorgoniidae
appearance, produce sperm and eggs. Members of this class Sea fans, such as this
include solitary anemones, colonial sea pens, soft corals, and Caribbean species, have
tropical reef-forming stony corals. upright fanlike colonies
supported by a central
slender red or axis, which is stiffened
8–12 in yellow lobe with a horny substance
20–30 cm called gorgonin.

8–12 in
20–30 cm

3½–20 ft
1–6 m
4–6 in
SEA 10–15 cm CARNATION
TOADSTOOL
LEATHER CORAL STRAWBERRY CORAL
Sarcophyton trocheliophorum Gersemia rubiformis. Dendronephthya sp.
f: Alcyoniidae f: Nephtheidae f: Nephtheidae
This soft coral can form enormous This stalked soft coral forms bright Typical of a family of stalked soft
leathery colonies on tropical reefs of pink lumpy colonies. It occurs in corals with polyps arranged in
the Indo-Pacific. It is fast-growing, northern parts of the Pacific clusters, this colorful species occurs
nourished by photosynthetic algae. and Atlantic Oceans. on Indo-Pacific tropical reefs.

white
polyps 20–40 in
50–100 cm 16–20 in
flexible
body 40–50 cm

4–8 in
10–20 cm

RED DEAD-MAN’S RED CORAL

FINGERS Corallium rubrum
Alcyonium glomeratum f: Coralliidae
f: Alcyoniidae Colonies of this 14–16 in
35–40 cm
This is a more slender, Mediterranean sea fan
upright ally of dead-man’s (not a true coral) are
fingers, found on sheltered supported by a skeleton
rocky European coastlines. of tiny, enmeshed, KIDNEY SEA PEN ORANGE SEA PEN
It varies from yellow COMMON DEAD-MAN’S calcified needles. It is Sarcoptilus grandis Ptilosarcus gurneyi
to red. FINGERS valued for jewelry. f: Pennatulidae f: Pennatulidae
Alcyonium digitatum Widespread in temperate- One of many brightly colored
f: Alcyoniidae waters, this sea pen has sea pens, this species from the
A typical soft coral, this thickly lobed kidney-shaped branches Pacific coast of N. America
European species has colonies of polyps, that occur in rows along retracts into its burrow when
which are carried on a fleshy mass each side of its body. disturbed by predators.
1½–6½ ft unsupported by a rigid skeleton.
0.5–2 m calcareous
tubes 6–12 in
15–30 cm

20–40 in
50–100 cm
20–40 in
50–100 cm

tentacle
WHITE SEA WHIP RED SEA WHIP ORGAN PIPE CORAL BLUE CORAL
Junceella fragilis Ellisella sp. Tubipora musica Heliopora coerulea
f: Ellisellidae f: Ellisellidae f: Tubiporidae f: Helioporidae
Sea whips are threadlike relatives of Ellisella sea whips form two-prong This Indo-Pacific soft coral has polyps Despite its hard, calcified skeleton,
sea fans, with their supporting horny branching colonies, some of which may in upright, calcified tubes, which are this species is more closely related to
axis reinforced with calcium. This is form dense underwater bushes. They connected to the colony by a rootlike soft coral than stony coral. It is the
an Indonesian reef species. occur in tropical and temperate waters. network at the base. only member of its order.
 4–8 in 13–16 ft 31/4–61/2 ft 4–4¾ in 31/4–9¾ ft
10–20 cm 4–5 m 1–2 m 10–12 cm 1–3 m
257

INVERTEBRATES • CNIDARIANS
FLOWERPOT CORAL LOBE CORAL LOBED BRAIN CORAL DAHLIA ANEMONE STAGHORN CORAL
Goniopora columna Porites lobata Lobophyllia sp. Urticina felina Acropora sp.
f: Poritidae f: Poritidae f: Lobophylliidae f: Actiniidae f: Acroporidae
This species has daisylike One of the most common Massive colonies of this brain Sticky swellings on this anemone The branching staghorn corals
polyps that are greatly reef-builders of the Indo-Pacific, coral—either flattened or may carry so much debris that, are among the largest tropical
lengthened when fully extended. this coral forms large encrusting dome-shaped—occur on reefs with tentacles retracted, it reef-builders. Nourished by
It is an Indo-Pacific relative of colonies in places buffeted by in tropical waters of resembles a small pile of gravel. photosynthetic algae, these
the lobe coral. strong wave action. the Indo-Pacific. It occurs around the North Pole. species are fast-growing.

31/4 ft 31/4–10 ft 4–8 in

1m 1–3 m 10–20 cm

DAISY CORAL LARGE-GROOVED MUSHROOM CORAL

Goniopora sp. BRAIN CORAL Fungia fungites
f: Poritidae Colpophyllia sp. f: Fungiidae
Long-reaching polyps of f: Faviidae This is one of many tropical MAGNIFICENT
The hemispherical brainlike 20–40 in
Goniopora stony corals on mushroom corals that are not 50–100 cm SEA ANEMONE
tropical reefs typically have 24 structure of this coral is typical of reef-building, but live among Heteractis magnifica
tentacles. They are among the its family. It is a tropical reef-builder other species as solitary polyps 1–6 in f: Stichodactylida
most flowerlike of corals. that contains photosynthetic algae. crawling over the ocean floor. 2.5–15 cm This giant anemone is from
Indo-Pacific reefs, where it lives in
close partnership with a variety of
300–600 ft/100–200 m fish species, including clown fishes.

3/8–1/2 in 2–2¾ in
10–15 mm 5–7 cm

ATLANTIC COLD
WATER CORAL
Desmophyllum pertusum
f: Caryophylliidae
Unlike many deep sea corals
lacking nutrient-supplying PLUMOSE ANEMONE SNAKELOCKS
Metridium senile ANEMONE
algae, this North Atlantic f: Metridiidae Anemonia viridis
species forms extensive reefs, f: Actiniidae
although only very slowly. This is a globally widespread
member of a family of The conspicuously long
anemones that is characterized tentacles of this common
by a fuzzy mass of tentacles. It European intertidal anemone
can split to form genetically rarely retract, even when
identical populations. exposed at low tide.

20–40 in
50–100 cm

4–6 in
10–15 cm

DEVONSHIRE CUP CORAL

Caryophyllia (Caryophyllia) smithii COMMON TUBE ANEMONE ATLANTIC
f: Caryophylliidae Cerianthus membranaceus BLACK CORAL
This N. E. Atlantic species belongs to a f: Cerianthidae Antipathes sp.
family containing cold water corals, Tube anemones burrow through f: Antipathidae
some with large, anemonelike polyps. sediment, using unique non-stinging Mostly found in deep waters,
Barnacles frequently attach to it. cnidocysts to build feltlike tubes from black corals consist of spiny polyp
mucus. This species occurs on colonies encased in slender,
European offshore mud. horny exoskeletons.
258 F L AT WO R M S
Structurally among the simplest of even without a blood stream to circulate PHYLUM PLATYHELMINTHES
animals, flatworms live wherever a the nutrients. Some free-living flatworms
IN VERTEBR ATES • FL AT WOR MS

CLASSES 6

moist habitat can provide their thin, are detritus-eaters, gliding along on a bed ORDERS 41

flat body with oxygen and food. of microscopic, beating hairs. Others prey FAMILIES About 420

on other invertebrates. SPECIES About 30,000

Platyhelminthes, commonly known as flatworms,

are a major phylum of animals found in various i n t e r n a l pa r a s i t e s
D E B AT E
forms in the ocean, in freshwater ponds, or even Tapeworms and flukes are parasites. Inside A NEW PHYLUM?
within the bodies of other animals. Superficially the host animal, their flattened body is
resembling leeches, flatworms are, in fact, far perfectly suited for absorbing nutrients. Many Traditionally, the tiny, gutless, and
simpler animals. With no blood system and no have complex ways of passing from host to brainless marine animals called acoels
host and may infect more than one type of are viewed as flatworms. Controversial
organs for breathing, they use their entire body
studies suggest that they belong in a
surface to absorb oxygen and release carbon animal. They enter the body of the host in new phylum of their own, and that they
dioxide. Flatworm species that have no gut also infected food or sometimes by penetrating are the most primitive living animals to
absorb their food in this way. In other species the skin. Once inside, a parasite may go deeper show bilateral symmetry—in contrast
the gut has one opening but branches frequently, into the body—burrowing through the gut to radial (round) cnidarians.
so that digested food can still reach all tissues, wall to lodge in vital organs.

GIANT 6½–23 ft
head cone INTESTINAL 2–7 m
with suckers FLUKE
Fasciolopsis buski
¾–2 in f: Fasciolidae
2–5 cm This E. Asian species
is one of the largest
flukes to infect humans.
It lives only in the
upper intestine
reproductive and does not enter
organs other organs.
COMMON
LIVER FLUKE divisible body
3 in sections each
Fasciola hepatica 7.5 cm
f: Fasciolidae contain sex
Flukes typically have complex, organs
parasitic life cycles. This species
infects grass-eating sheep or
cattle via freshwater snails, and
feeds from the host’s liver.

CATTLE-SNORING
BLOOD FLUKE
⅜ in
Schistosoma nasale
10 mm f: Schistosomatidae PORK
This parasite is transmitted TAPEWORM
3⁄16–¼ in through freshwater snails. Taenia solium
5–6 mm It infects cattle, causing f: Taeniidae
nasal growths that Tapeworms are intestinal
make the cattle snore parasites, but at an intermediate
JACK FLUKE when breathing. stage can invade muscle as cysts.
Provitellus turrum This species’ intermediate host
f: Monorchiidae is pigs, and may pass to humans
Provitellus flukes infect in infected meat.
the guts of fishes living
on tropical reefs. This
species, shown here in
false color, has been
recorded in jack fishes. worms on
coral surface
3⁄16 in
5 mm
EPIZOIC
FLATWORM 3¼– 4 in 1½–2 in
Waminoa sp. 8–10 cm 4–5 cm CANDY STRIPE
f: Convolutidae FLATWORM
This coral-living GIANT LEAF FLATWORM Prostheceraeus vittatus
species is one of Kaburakia excelsa f: Euryleptidae
many marine f: Stylochidae This animal belongs to a group of mainly
micro-flatworms that Like other leaf flatworms, this N. American marine, free-living, cross-fertilizing
resemble planktonic intertidal species is mainly carnivorous. It smothers hermaphrodites called polyclads.
larvae of cnidarians. its prey with a mouth-bearing extension of its gut. It is an Atlantic Ocean species.
 RO U N DWO R M S
BLACK AND YELLOW
FLATWORM
Pseudoceros dimidiatus 259
f: Pseudocerotidae
Marine polyclads are large,
free-living flatworms. Many The simple, cylindrical roundworms are remarkably
successful—surviving almost everywhere, resisting

INVERTEBR ATES • ROUNDWORMS

are strikingly colored, warning
predators of their nasty
taste. This species is from drought, and reproducing rapidly.
Indo-Pacific waters.
2¾–3¼ in
The phylum Nematoda is PHYLUM NEMATODA
7–8 cm ubiquitous. There can be millions CLASSES 2
of roundworms in a square yard ORDERS 17
of soil, and they also live in FAMILIES About 160

freshwater or marine habitats. SPECIES About 26,000

Many are parasites. These

undulating worms, or nematodes, can be extraordinarily prolific, producing
margin hundreds of thousands of eggs per day. When environmental
SHOVEL-HEADED
LAND FLATWORM conditions deteriorate they can survive heat, frost, or drought by
Bipalium kewense producing resistant stages and becoming dormant.
f: Bipaliidae
Land flatworms are mostly
8–12 in
20–30 cm
Roundworms have a muscle-lined body cavity and two openings
tropical and need damp conditions. to their gut—the mouth and anus. Their cylindrical body is
This Asian species has been
accidentally introduced to coated in a tough layer, called a cuticle—functionally similar to
greenhouses around the world. that of arthropods—which they molt periodically as they grow.
11/2–31/2 in
4–9 cm COMMON SOIL
7⁄32–1/2 in 1/32 in
7–11 mm ROUNDWORM 1 mm
FLOWERS’ FLATWORM Caenorhabditis elegans
Pseudobiceros flowersi f: Rhabditidae
f: Pseudocerotidae stained green
This widespread, colorless,
Like many reef-dwelling polyclads, soil-living roundworm is
this Indo-Pacific species glides with an used in research to further the
undulating margin. It has been found understanding of genetics
under rubble in lagoons. NEW WORLD and development.
HOOKWORM
Necator americanus
f: Uncinariidae stained orange
2¾–3¼ in Hookworm larvae
7–8 cm penetrate skin and
travel in the blood to
the lungs of the host. ¹/32 in
1 mm
GOLD-SPECKLED FLATWORM They crawl to the
Thysanozoon nigropapillosum throat, are swallowed,
f: Pseudocerotidae and mature in the gut.
Many Thysanozoon polyclads are coated with This species, shown SOYBEAN
1/2– ¾ in pimply swellings, which are yellow-tipped here in false color, CYST ROUNDWORM
1.5–2 cm on this otherwise velvety black species of parasitizes humans, Heterodera glycines
the Indo-Pacific. dogs, and cats. f: Heteroderidae
4–61/2 in This colorless, soil-living roundworm infects the
10–17 cm roots of leguminous plants around the world,
DRAB FRESHWATER NEW ZEALAND causing cysts and significantly reducing yields.
FLATWORM LAND FLATWORM
Dugesia lugubris Arthurdendyus
f: Planariidae triangulatus
Triclads are flatworms with f: Geoplanidae
a triple-branched gut. This This large, soil-living
European species is one of species is native to
many triclads found in New Zealand, but has
freshwater habitats; others invaded Europe. It preys
are marine. on earthworms. 1¼–2 in
3–5 cm

6–14 in
15–35 cm

3/8–1/2 in ¾–1¼ in
1–1.5 cm 2–3 cm

STREAM COMMON LARGE INTESTINAL

BROWN FLATWORM WHIPWORM ROUNDWORM
FRESHWATER Dugesia gonocephala Trichuris trichiura Ascaris lumbricoides
FLATWORM f: Planariidae f: Trichuridae f: Acarididae
Dugesia tigrina Many of the freshwater Like many other gut parasites, A common parasite of humans
f: Planariidae Dugesia flatworms, such the mainly tropical whipworm in regions of poor sanitation
This species is native to as this European species infects humans who eat food where people get infected by
freshwater habitats of of running water, have contaminated with feces. It eating food contaminated with
N. America, but has been earlike flaps for detecting completes its life cycle in feces, with adult worms then
introduced to Europe. water currents. the intestines. living in the host intestine.
260 S E G M E N T E D WO R M S
With more complex muscle and organ Marine segmented worms—the predatory PHYLUM ANNELIDA
systems than flatworms, many species ragworms and their filter-feeding relatives—
INVERTEBRATES • SEGMENTED WORMS

CLASSES 4

of this phylum, called Annelida, are carry bundles of bristles along their body, typically ORDERS 17

accomplished swimmers or burrowers. borne on little paddles that help with swimming, FAMILIES About 130

burrowing, or even walking. This group of SPECIES About 18,000

Segmented worms include earthworms, ragworms, bristled, segmented worms are called polychaetes
and leeches. For most, their blood circulates in or bristle worms.
vessels and they have a firm sac of fluid (the coelom) Land-living earthworms are more sparsely
running the length of their body, which keeps bristled detritus-feeders, and are important
motion of the gut separate from motion of the recyclers of dead vegetation and aerators of soil.
body wall. The coelum is split into sections, with Many leeches are specialized further, and carry
each one corresponding to a body segment. Each suckers to extract blood from a host. Their saliva
body segment contains a set of muscles, and contains chemicals that stop blood clotting.
coordination of these sets can send a wave of Other leeches are predatory. Both earthworms
contraction down the length of the body, or make and leeches have a saddle-shaped, glandular
it flex back and forth. This makes many annelids structure around their body called a clitellum,
highly mobile both on land and in the water. which they use to make their egg cocoons.

3/4–23/4 in SLUDGE WORM 20 in MEGADRILE

2–7 cm Tubifex sp. 50 cm EARTHWORM
f: Naididae Glossoscolex sp.
This widely distributed worm can f: Glossoscolecidae
be seen in mud polluted by sewage, Glossoscolex are large
the front end buried, the rear end earthworms from tropical
wiggling to extract oxygen. C. and S. America. Many are
found in rainforest habitats.
6–10 in
15–25 cm
4–6 in 11/2–23/4 in
10–15 cm 4–7 cm
clitellum CHRISTMAS TREE
or “saddle” TUBE WORM
TIGER WORM Spirobranchus giganteus
Eisenia foetida COMMON EARTHWORM f: Serpulidae
f: Lumbricidae Lumbricus terrestris Characterized by its spiral
This European inhabitant of rotting f: Lumbricidae whorls of tentacles, used for
vegetation secretes defensive pungent fluid This earthworm—native to Europe but filter feeding and extracting
and, like other earthworms, has a “saddle” introduced elsewhere—drags leaves into oxygen, this species is
for producing cocoons of eggs. its burrow at night as a source of food. widespread on tropical reefs.

V E LV E T WO R M S
These soft-bodied worms, cousins of rainforests of tropical America, Africa, and PHYLUM ONYCHOPHORA
arthropods, lumber slowly along dark Australasia; they shun the open, preferring CLASS 1

forest floors like giant caterpillars, but to hide in crevices and leaf litter. They come ORDER 1

they are extraordinary hunters. out at night or after rainfall to hunt other FAMILIES 2

invertebrates. Velvet worms capture their prey SPECIES About 200

These animals have the body of an earthworm in a unique way: they immobilize their victims
and the multiple limbs of a millipede, but belong by spraying them with sticky slime which is
in a phylum of their own: Onychophora. Velvet produced by glands that open through pores
worms are rarely seen in their warm native straddling the mouth.

skin covered SOUTHERN 4–8 in CARIBBEAN

in fine hairs AFRICAN 10–20 cm VELVET WORM
VELVET WORM Epiperipatus broadwayi
Peripatopsis moseleyi f: Peripatidae
f: Peripatopsidae This species is one of a family
4 in This species belongs of equatorial velvet worms,
10 cm to a family with a which typically have more
globally southern legs than those of the
distribution. southern continents.
 21/4–12 in SEA MOUSE 1–11/4in 43/4–10 in 31/4–13 ft
6–30 cm Aphrodita aculeata 2.5–3cm 12–25 cm 1–4 m
f: Aphroditidae
This mud-burrowing LUGWORM HONEYCOMB
FIRE WORM scale worm is found in Arenicola marina WORM
Hermodice carunculata the shallow waters of f: Arenicolidae Sabellaria alveolata
f: Amphinomidae N. Europe. Its scales SEA CUCUMBER An earthwormlike f: Sabellaridae
are covered with SCALE WORM
A tropical Atlantic offshore species, Gastrolepidia clavigera polychaete, the lugworm This tubeworm builds its tubes
this polychaete preys on corals, a hairlike coat. lives in burrows on beaches from sand and shell fragments.
f: Polynoidea
sucking out the soft flesh from the This Indo-Pacific polychaete and mudflats, where it Dense populations form
hard skeleton. Its body extensions 4–8 in ingests sediment and feeds honeycomblike reefs in the
10–20 cm has flattened back scales. It is
have painfully irritating bristles. parasitic on sea cucumbers. on detritus. Atlantic and Mediterranean.

2–6 in 31/4–4 in
5–15 cm 8–10 cm

GREEN PADDLE WORM PACIFIC FEATHERDUSTER

Eulalia viridis WORM
f: Phyllodocidae Sabellastarte sanctijosephi
Paddleworms are active carnivores f: Sabellidae
with leaf-shaped flaps on their body A tropical Indo-Pacific tubeworm,
extensions. This European species this species is common along
lives among intertidal rocks and kelp. coastlines, including coral reefs
and tidal pools.
10–16 in
25–40 cm

2–23/4 in
5–7 cm

KING RAGWORM
Alitta virens
f: Nereididae RED TUBE
Ragworms, close relatives WORM
of paddleworms, have Serpula vermicularis
two-pronged body extensions. f: Serpulidae
This Atlantic mud-burrowing Serpulid tubeworms
species can deliver a painful bite. build hard, chalky tubes.
Like most serpulids, this 1⁄16–1/8 in
widespread species has 2–3mm
GIANT TUBE WORM tentacles modified to plug
Riftia pachyptila the tube after retraction. NORTHERN SPIRAL
f: Siboglinidae TUBE WORM
This tubeworm lives in the body used Spirorbis borealis
hot, sulfur-rich darkness as anchor f: Serpulidae
of ocean-floor volcanic 61/2–73/4 ft The small coiled tube of
2–2.4 m
vents in the Pacific. The red this tubeworm is cemented
plume harbors bacteria to brown Fucus seaweeds
that make its food using and kelps along North
chemicals from these vents. Atlantic coastlines.

WA T E R B E A R S
Visible only through a microscope, algae, where many use their needlelike jaws PHYLUM TARDIGRADA
stubby-legged, agile water bears share to pierce the cells of this vegetation and suck its CLASSES 3

their aquatic community with microbes sap. Many water bear species are known only as ORDERS 5

and much simpler invertebrates. females, which reproduce asexually by spawning FAMILIES 20
offspring from unfertilized eggs. If its habitat SPECIES About 1,000

Tiny water bears, or tardigrades (meaning dries up, a water bear is able to survive
“slow-stepped”) clamber through miniature by going into a kind of suspended
forests of water weed, four pairs of short legs animation called cryptobiosis, shriveling
hanging on with clawed feet. Most are less than into a husk, sometimes for years at a
a millimeter long. They abound among moss or time, until rainfall revives it.

MOSS 0.25mm clawed feet

WATER BEAR
Echiniscus sp. SEAWEED WATER BEAR 0.25mm
f: Echiniscidae Echiniscoides sigismundi
Many water bears live in moss, f: Echiniscoididae
but their ability to survive in a One of many little-known marine water bears,
dried state has helped a number of this species has been recorded among seaweed
species to disperse worldwide. in coastal locations around the world.
262 A RT H RO P O D S
Jointed legs and flexible armor have in millipedes and centipedes. In other groups, PHYLUM ARTHROPODA
helped a phylum that includes winged various segments have fused together into
INVERTEBRATES • ARTHROPODS

CLASSES 19

insects and underwater crustaceans to discrete body sections. Insects are divided into ORDERS 123

evolve unrivaled diversity. a sensory head, a muscular thorax with legs and FAMILIES About 2,300

wings, and an abdomen containing most of SPECIES About 1.2 million

There are more species of arthropods known the internal organs. In arachnids and some
to scientists than all the other animal phyla put crustaceans, the head and thorax are fused D E B AT E
together—and doubtless many more await into one single section. CAMOUFLAGE
discovery. They have an extraordinary range of AND MIMICRY
lifestyles including grazing, predation, filtering g e t t i n g ox yg e n
particles of food in water, and drinking fluids Aquatic arthropods, such as crustaceans, Many arthropods have adapted to their
environment by blending imperceptibly
such as nectar or blood. breathe with gills. The bodies of most land
into it. Stick insects, for example, look
Arthropods are coated by an exoskeleton made arthropods—insects and myriapods—are so like twigs that they are very difficult
of a tough material called chitin. It is sufficiently permeated by a network of microscopic, for predators to spot. In contrast,
flexible around the joints and between segments air-filled tubes called tracheae. These open out arthropods, such as wasps, have
to allow for mobility, but resists stretching. As through pores known as spiracles on the sides evolved bright colors that warn others
arthropods grow, they must periodically molt their of the body, usually one pair per body section. they are distasteful or dangerous. The
exoskeleton, which is replaced by a slightly larger Small muscles inside the spiracles regulate the hornet moth is completely harmless
one. The exoskeleton serves as protective armor flow of air by means of valves. In this way, but avoids predation because it looks
but also helps reduce water loss in very dry habitats. oxygen can reach all cells of the body and does and sounds exactly like a hornet.
Despite their similar appearance, other
not need to be transported in blood. Some
anatomical features separate them into
b o dy pa rt s arachnids breathe with tracheae; others with different orders. The hornet moth
Arthropods evolved from a segmented ancestor, book lungs in their abdomen that evolved from belongs to the order Lepidoptera and
perhaps a wormlike creature. Body segmentation the gills of their aquatic ancestors. Most use a the hornet to the order Hymenoptera.
persists in all arthropods, but is especially obvious combination of the two.

MILLIPEDES AND
CENTIPEDES segmented
exoskeleton
head

Millipedes and centipedes form a group of

multi-segmented arthropods called myriapods.
Millipedes typically have two legs per body 11/2–2 in
segment; centipedes have only one per segment. 4–5 cm
Whereas millipedes are vegetarian, centipedes stout sensory
are predatory carnivores. antenna

1⁄16–1/8 in
2–3 mm BROWN GIANT PILL MILLIPEDE
Zephronia sp.
f: Zephroniidae
AMERICAN SHORT- Giant pill millipedes, such as this Bornean species,
HEADED MILLIPEDE 3/8–3/4 in differ from their smaller northern hemisphere
Brachycybe sp. 1–2 cm relatives in that they have 13 body sections
f: Andrognathidae rather than 12.
BRISTLY MILLIPEDE This is a N. American representative of a BLACK GIANT PILL
family of small, flat millipedes that live 11/4–11/2 in
Polyxenus lagurus 3–4 cm MILLIPEDE
f: Polyxenidae in rotting wood and leaf litter. Zoosphaerium sp.
One of a family of millipedes f: Sphaerotheriidae
armed with defensive One of many southern
bristles, this tiny species of overlapping plates hemisphere giant pill
the northern hemisphere millipedes, this species
lives under the bark of trees is native to Madagascar.
and in leaf litter.
frontal
shield

WHITE-RIMMED PILL
MILLIPEDE 1/4–3/4 in
Glomeris marginata 0.6–2 cm
f: Glomeridae
Pill millipedes, such as this
European species, have fewer
body segments than other
millipedes and can roll into
a ball when disturbed.
 263
3/4–21/4 in
BLACK SNAKE MILLIPEDE 2–6 cm
Tachypodoiulus niger
f: Julidae
1/2–11/4 in Unlike its close relatives, this distinctive,
1.5–3 cm white-legged millipede from W. Europe 3–5 in
spends most of its time above ground— 7.5–13 cm
even climbing trees and walls.
AMERICAN GIANT MILLIPEDE
Narceus americanus
f: Spirobolidae
This large, Atlantic coast millipede belongs to a family
8–15 in of mainly American, cylindrical millipedes. Like its
20–38 cm
relatives, it releases noxious chemicals in defense.
BROWN SNAKE MILLIPEDE
Julus scandinavius
f: Julidae AFRICAN GIANT MILLIPEDE BORING MILLIPEDE
A member of a large family Archispirostreptus gigas Polyzonium germanicum
of cylindrical millipedes with f: Spirostreptidae f: Polyzoniidae
ring-encased segments, this species One of the largest of all millipedes, this This somewhat primitive millipede
occurs in European deciduous species occurs mainly in tropical E. Africa and is has a scattered distribution in Europe. 3⁄16–3/4 in
woodland, especially on acid soils. one of many species that release irritating It lives in woodland and resembles 0.5–1.8 cm
chemicals to defend themselves. beech bud scales when coiled.

EASTERN FLAT- YELLOW EARTH

BACKED MILLIPEDE CENTIPEDE
Polydesmus complanatus Geophilus flavus
f: Polydesmidae 1/2–21/4 in f: Geophilidae 3/4–13/4 in
Polydesmids have projections 1.5–6 cm Eyeless geophilids have 2–4.5 cm
on their exoskeleton that more segments (and
make them look flat-backed. therefore legs) than other
This E. European species centipedes. This European
is a fast runner. soil-dweller has been
introduced to the
Americas and Australia.

11/2–21/4 in 3/4–11/4 in 1–2 in

4–6 cm 2–3 cm 2.5–5 cm
TANZANIAN FLAT-BACKED
MILLIPEDE
Coromus diaphorus
f: Oxydesmidae
The shiny, pitted surface typical
of many of the eyeless flat-backed
millipedes is especially prominent 3/4–11/4 in
in this species from tropical Africa. 2–3 cm
BROWN STONE
CENTIPEDE
Lithobius forficatus
BANDED STONE f: Lithobiidae
CENTIPEDE Found sheltering beneath bark
Lithobius variegatus and rock, stone centipedes
f: Lithobiidae characteristically have 15 body
Once thought to be confined to Britain, segments. This globally
this centipede is now known in continental widespread species frequents
Europe, where native populations have forests, gardens, and seashores.
unbanded legs.

HOUSE
CENTIPEDE
Scutigera coleoptrata
f: Scutigeridae
An unmistakable centipede,
this long-legged animal
with compound eyes is one
of the fastest invertebrate
runners. Native to the
Mediterranean region, it has
“ fangs” inject venom been introduced elsewhere.
8–10 in
20–25 cm

TIGER GIANT BLUE-LEGGED CENTIPEDE

CENTIPEDE Ethmostigmus trigonopodus
Scolopendra hardwickei one pair of f: Scolopendridae
f: Scolopendridae legs per body This close relative of the giant
Many of the giant Scolopendra centipedes have segment Scolopendra centipedes is widespread
vibrant warning colors. This Indian species 4–6 in throughout Africa and is one of
is one of several with a “tiger” pattern. 10–15 cm several species with blue-tinted legs.
264 A R AC H N I D S
A class within the phylum Arthropoda, chelicerae as fangs to inject venom and many PHYLUM ARTHROPODA
arachnids include predatory spiders first ensnare prey by spinning webs. Scorpions
INVERTEBRATES • AR ACHNIDS

CLASS ARACHNIDA

and scorpions, as well as mites and inject venom with the stinger in their tail. ORDERS 12

blood-sucking ticks. Between their legs and chelicerae, arachnids FAMILIES 661

also have a pair of limblike pedipalps. These SPECIES About 103,000

Arachnids and related horseshoe crabs are are modified into grasping pincers in scorpions
chelicerates—arthropods named for their or sperm-transferring clubs in male spiders.
clawlike mouthparts (chelicerae). The head and
thorax of chelicerates are fused into one body m i c ro s c o p i c d i v e r s i t y
section, which carries the sensory organs, brain, Many mites are too small to see with the naked
and four pairs of walking legs. Unlike other eye. They abound in almost every habitat, where
arthropods, chelicerates lack antennae. they scavenge on detritus, prey on other tiny
invertebrates, or live as parasites. Some lead
e f f e c t i v e p r e dat o r s innocuous lives in skin follicles, feathers, or
Scorpions, spiders, and their relatives are land- fur; others cause disease or allergy. One group—
This female wasp spider waits at the
living predators that have evolved quick ways of the ticks—are blood-suckers and can spread center of her dewy orb web to ensnare
immobilizing and killing prey. Spiders use their disease-causing microbes. any insect prey that might fly in.

CHILEAN BURROWING 1⁄16–1⁄8 in

SCORPION 1.5–2.5 mm
Centromachetes pococki
f: Bothriuridae
1½–2 in A burrowing scorpion from the southern 1/8 in 1/8 in
4–5 cm hemisphere, this species occurs in 3 mm 3 mm
temperate forests in S. America,
often under rotting logs.

MARITIME FINGER-CLAWED BROAD-HEADED

PSEUDOSCORPION PSEUDOSCORPION PSEUDOSCORPION
Neobisium maritimum Dactylochelifer latreillei Chthonius ischnocheles
f: Neobisiidae f: Cheliferidae f: Chthoniidae
A common W. European Pseudoscorpions are tiny Pseudoscorpions, such as this
coastal pseudoscorpion, this arachnids that kill their prey European species, sometimes
species lives under stones and with venomous pincers. This find small prey in animal nests by
seaweed in the splash and European species lives at the hitchhiking in the fur or feathers
upper intertidal zones. base of seashore grasses. of the parent animal.
stinger
6–10 in
vesicle contains 15–25 cm AFRICAN ROCK SCORPION
two poison sacs Hadogenes phyllodes
f: Hemiscorpiidae
Typical of a family of rock scorpions,
this southern African species has a
IMPERIAL broad, flat body for squeezing into
SCORPION cracks between rocks.
Pandinus imperator
pair of eyes on top f: Scorpionidae 4–7 in
10–18 cm
of the cephalothorax One of the largest of all scorpions,
this African forest species is more
docile and has a milder sting than
many of the desert-dwellers.
23/4–4 in 3/8 in
7–10 cm 1 cm
COMMON
EUROPEAN SCORPION
Buthus occitanus
f: Buthidae
Found throughout N. Africa and
around the Mediterranean, the
toxicity of this species varies
across its range, being most
potent in the south.
YELLOW SAY’S
FAT-TAIL SCORPION HARVESTMAN
Androctonus amoreuxi Vonones sayi
f: Buthidae f: Cosmetidae
Most fat-tail scorpions are Harvestmen have an undivided
21/4–31/4 in small but have potent venoms. body and are non-venomous.
6–8 cm This large species from the Sahara Many, including this American
pedipalps modified and Middle East has caused species, produce distasteful
as pincers human fatalities. chemicals to deter predators.
 pair of
small eyes AMERICAN SUN SPIDER
massive
Eremobates sp. 265
chelicerae
f: Eremobatidae
Eremobates are sun spiders with 31/4–4 in
particularly large jaws. This 8–10 cm

INVERTEBRATES • AR ACHNIDS
nocturnal species is found in warm
parts of N. and C. America. PAINTED SUN SPIDER
Metasolpuga picta
f: Solpugidae
Sun spiders, also called camel
31/4–6 in spiders, are fast-running, desert-living
8–15 cm relatives of true spiders. This is a
leglike diurnal species from Namibia.
pedipalp
segmented 1–2 in
abdomen 2.5–5 cm WIND SPIDER
Galeodes arabs
f: Galeodidae
1⁄64 in A common species in the largest
0.3–0.6 mm genus of sun spiders, this Middle
Eastern species is named for its
1⁄32 in 1/8–3⁄16 in ability to tolerate sand storms.
1 mm 3–5 mm

1⁄64 in 5⁄16–3/8 in
0.4 mm 8–10 mm

characteristic white
spot on body
FLOUR MITE AUTUMN CHIGGER COMMON VELVET MITE RED SPIDER MITE
Acarus siro Neotrombicula autumnalis Trombidium holosericeum Tetranychus urticae
f: Acaridae f: Trombiculidae f: Trombidiidae f: Tetranychidae
This is an important pest Adults of chigger mites This is a widespread Eurasian Spider mites are a
that feeds on stored cereal are vegetarian, but their species of velvet mite. When family of mites that suck
products. Like many larvae feed on the skin of young, it lives parasitically off the sap of plants. They
other species, its presence other animals, including other arthropods, but becomes weaken the plant and can
can cause allergic humans. Their chewing predatory when mature. transmit viral diseases.
reactions in humans. causes intense irritation.

1⁄32–1⁄16 in 1⁄10–1⁄64 in 1⁄64–1⁄32 in 1⁄64 in

1–2 mm 0.3–0.5 mm 0.5–1 mm 0.5 mm

LONE STAR TICK

Amblyomma americanum
f: Ixodidae
Like other blood-sucking
ticks, this species—common
in woodlands in the US—
can transmit a number of
disease-causing microbes.
VARROA MITE MANGE MITE CHICKEN MITE PERSIAN FOWL TICK
Varroa sp. Sarcoptes scabiei Dermanyssus gallinae Argas persicus
f: Varroidae f: Sarcoptidae f: Dermanyssidae f: Argasidae
Parasitic on honeybees, young This tiny mite burrows into the A blood-sucking parasite of A blood-sucking parasite of fowl,
of this mite feed on bee larvae. skin of various mammal species, poultry, this mite lives—and including domestic chickens, this pincerlike
When mature, they attach to completing its life cycle there. completes its life cycle—in oval-shaped, soft-bodied tick can 3⁄4–11/2 in
2–4 cm palps
and feed on adult bees and It causes scabies in humans and crevices away from its host, spread disease between birds and
spread to other hives. mange in carnivores. but emerges at night to feed. can cause paralysis.
very long
front legs

3/8–1/2 in 5⁄32–11⁄32 in 11/4–11/2 in

1–1.5 cm SPINY 4–9 mm 3–4 cm
HARVESTMAN
Discocyrtus sp.
f: Gonyleptidae
This S. American harvestman
has spiny hind legs as a possible
defense against predators.
It lives under stones and
logs in forests.
TAILLESS WHIP SPIDER WHIP SCORPION
HORNED HARVESTMAN Phrynus sp. Thelyphonus sp.
Phalangium opilio f: Phrynidae f: Thelyphonidae
f: Phalangiidae Whip spiders are not true Tropical whip scorpions such
A common harvestman of Eurasia spiders. They have long, as this species have a whiplike
and N. America, the male of this whiplike front legs and pincers tail, but no stinger or venom.
species has a jaw ornamented for grabbing prey, but lack However, they can spray acetic
with projecting horns. venom. All are tropical. acid from their abdomen.
 A R AC H NI DS ORANGE BABOON
SPIDER
266 2–3 in Pterinochilus murinus
5–7.5 cm f: Theraphosidae
pedipalp African baboon spiders are
named because their leg
INVERTEBRATES • AR ACHNIDS

segments resemble baboon

3/4–2 in 2–3 in fingers. Like other baboon
2–5 cm 5–7.5 cm spiders, this species excavates
burrows using its fangs
CHACO TARANTULA and pedipalps.
Acanthoscurria insubtilis
f: Theraphosidae
Many large theraphosids,
such as this S. American
FUNNEL-WEB SPIDER MEXICAN RED-KNEED species, live in abandoned 2–21/2 in
Atrax robustus TARANTULA rodent burrows. They are 5–6 cm
f: Atracidae Brachypelma smithi ambush predators, lying
Females of this aggressive f: Theraphosidae in wait for prey.
Australian spider live in This is one of many
burrows with funnellike, theraphosids: large-bodied
spiders popularly called forward-pointing
silk-lined entrances. The chelicerae carrying fangs
dangerous bite most bird-eaters or tarantulas.
frequently comes from males They prey on large insects,
looking for mates. rarely on small vertebrates.
eight small eyes
3/8–3/4 in 1
⁄16 in
1–2 cm 1.5–2 mm

3/8–1/2 in
1–1.5 cm
NORTH AMERICAN SIX-EYED SPIDER WOODLOUSE SPIDER hairy, brown
TRAPDOOR SPIDER Oonops domesticus Dysdera crocata body
Ummidia audouini f: Oonopidae f: Dysderidae
f: Halonoproctidae This tiny, pink, six-eyed This nocturnal European
This N. American spider builds spider occurs in warm spider has huge fangs that
a corklike trapdoor set with parts of Eurasia, but catch and break into the hard
trip lines. Stumbling prey alert further north—including exoskeletons of woodlice. It
the predator waiting in a in Britain—it is found occurs in damp places where
silk-lined burrow beneath. only in houses. woodlice are abundant.

1/4–1/2 in white bands spinnerets

0.6–1.6 cm on legs extrude silk

1/8 in
3 mm
LADYBUG DWARF
SPIDER SPIDER
Eresus kollari Gonatium sp.
f: Eresidae f: Linyphiidae 1/8–1/4 in
Only males of this Eurasian Typical of a very large 3–6 mm
spider have the eponymous family of tiny “money
ladybug pattern. This spiders,” this northern NORTHERN
species catches prey from hemisphere species builds 5⁄32–1/2 in
SPITTING SPIDER 4–13 mm
within silk-lined burrows sheet webs and can travel Scytodes thoracica
on heathland slopes. on the wind by ballooning f: Scytodidae EUROPEAN GARDEN SPIDER
on silken threads. Spitting spiders are sluggish Araneus diadematus
spiders that spray gummy, f: Araneidae
poisonous fluid to immobilize A weaver of orb webs from the northern
prey before biting. This species hemisphere, this spider of woodland, heath,
spans the northern hemisphere. and gardens is marked by a white cross on
a variable background color.

1/4–13/4 in 1⁄16–11⁄32 in
0.6–4.5 cm 2–9 mm
DADDY LONG-LEGS CRABLIKE SPINY
SPIDER AMERICAN GOLDEN ORB-WEAVER
Pholcus phalangioides SILK ORB-WEAVER Gasteracantha cancriformis
f: Pholcidae Trichonephila clavipes f: Araneidae
egg sac Spindly-legged pholcids vibrate f: Araneidae One of many American
their webs when disturbed. This is the only American orb-weavers with defensive
Many are cave dwellers, but species of a group of tropical spines, this species inhabits
7⁄32–3/8 in this cosmopolitan species golden orb-weavers. It is the S. US and the
7–10 mm inhabits houses. Females carry distinguished by feathery Caribbean. Colors of the
eggs in their jaws. tufts on its legs. body and spines vary.
 5⁄32–1/2 in 3/4–11/4 in NURSERY WEB
4–13 mm 2–3 cm SPIDER
Pisaura mirabilis 267
f: Pisauridae
Females of this Eurasian
spider carry their egg sac
beneath the body by their
jaws, before weaving a
nursery “tent” of silk to
3⁄16–5⁄16 in protect hatching young.
5–8 mm

BLACK TARANTULA WOLF SPIDER EUROPEAN WOLF SPIDER

WIDOW SPIDER Lycosa tarantula Pardosa amentata egg sac
3/8–1/2 in
Latrodectus mactans f: Lycosidae f: Lycosidae carried by jaws
1–1.5 cm
f: Theridiidae This species is the only true tarantula: A typical wolf spider, this furry
A small spider with venom a large member of the wolf spider brown spider hunts on the ground
unusually dangerous to humans, family from Mediterranean Europe, without a web. Females carry an
females of this N. American not to be confused with hairy- egg sac and young ride on
species frequently eat the smaller bodied “bird-eaters.” the mother’s back.
male after mating.
distinctive
pale stripes
3/8 in 7⁄32–7⁄10 in 3/8–3/4 in on body 3/8–7/8 in
1.2 cm 7–18 mm 1–1.8 cm 1–2.2 cm

silvery abdomen
is due to
trapped air dark,
velvety body

egg sac
WATER SPIDER GIANT HOUSE SPIDER RAFT SPIDER
Argyroneta aquatica Eratigena duellica Dolomedes fimbriatus
CAVE SPIDER f: Dictynidae f: Agelenidae f: Pisauridae
Meta menardi The only routinely aquatic spider, A member of a group of A European swamp spider,
f: Tetragnathidae this Eurasian pond species builds spiders that build sheet webs this species preys on
Many spiders of the genus Meta, such as a submerged, air-filled chamber, with tubular retreats, this small fishes, which it can
this European species, inhabit caves, where inside which it eats its prey, species frequents buildings attract by vibrating the water
they suspend their drop-shaped egg sacs. including small fishes. across the northern hemisphere. surface with its legs.

3⁄16–3/8 in
5–10 mm
GOLDENROD
CRAB SPIDER
Misumena vatia
11/4–2 in f: Thomisidae 3/4–11/4 in
3–5 cm Females of this northern 2–3 cm
hemisphere spider can slowly
BRAZILIAN change their color from white
WANDERING SPIDER to yellow for camouflage in
Phoneutria nigriventer flowers before ambushing
f: Ctenidae nectar-feeding insects.
Wandering spiders get their BROWN HUNTSMAN
SPIDER
name from their nocturnal Heteropoda venatoria
roaming habits. This species large forward-facing f: Sparassidae
from the Brazilian forests is eyes for judging distance
believed to be the world’s Widespread throughout tropical and
most venomous spider. subtropical regions, this large but
harmless spider is sometimes welcomed
in homes for preying on cockroaches.

thickened front legs

ELEGANT JUMPING
SPIDER
Chrysilla lauta BROWN JUMPING
f: Salticidae 1/8–3/8 in SPIDER
Jumping spiders are 3–9 mm Evarcha arcuata
especially diverse in the f: Salticidae
tropics. Many, such as this This Eurasian grassland species
3⁄16–7⁄32 in
E. Asian species, are 5–7 mm is one of thousands of species
strikingly colored, of jumping spiders with acute
sometimes to aid binocular vision and complex
communication. courtship behavior.
268 MEXICAN
RED-KNEED
TA R A N T U L A
INVERTEBRATES • AR ACHNIDS

Brachy pelma smithi

With its stout, furry body, the Mexican red-kneed tarantula
might seem more mammallike than spiderlike. The female
can live up to 30 years, which is unusual for an invertebrate, but the life
span of the male is only up to six years. Brachypelma smithi is one of a group of
spiders that gets the nickname of bird-eater because of their size, and although
most of their diet consists of fellow arthropods, they can and do take down size Body length 2–3 in (5–7.5 cm)
small mammals and reptiles when the opportunity arises. In its native Mexico habitat Tropical deciduous forests
this species lives in burrows in earth banks, where it is safe to molt and lay eggs, distribution Mexico
diet Mostly insects
and from which it ambushes its prey. Habitat destruction is now a threat, and, as
the species makes a popular pet, it is routinely bred in captivity.
legs are covered in
EYES specialized hairs sensitive
Like most spiders, the tarantula has to air movements and touch
eight simple eyes arranged at the
front of the head. Even so, it has poor
vision, and relies more on touch
than sight to sense its surroundings
and the presence of prey.

padded foot

JOINTS
Like all arthropods,
tarantulas have jointed
legs. Each leg consists
of seven tubular sections
of exoskeleton that connect
via flexible joints. Muscles
run through these joints
to move the sections.

FOOT
At the tip of each foot are two
claws used for grip as the tarantula
moves. As in other hunting spiders,
there are also pads of tiny hairs for
extra gripping on smooth surfaces.

SPINNERETS
Glands in the abdomen
produce silk in liquid form.
Using its hind legs, the
VENOMOUS FANGS tarantula pulls the silk from
Tarantula fangs hinge forward when attacking prey— tubes called spinnerets. The
unlike most other spiders, whose fangs are angled silk solidifies into threads
toward each other. Both fangs inject venom from used to make egg sacs and
muscular sacs within the head to paralyze the victim. line the spider’s burrow.
 dark abdomen contains
most of the animal’s
vital internal organs

orange-red
patella, or knee

HAIRY WEAPONS
Like many tropical American
tarantulas, this spider defends
itself by rubbing its hind legs
against its abdomen to release
the barbed, stinging hairs that
grow there. These urticating
hairs are tiny and light, and
designed to float in a cloud
around a predator’s face,
getting in its eyes, nose, and
palps are used for mouth. They cause intense
feeling, groping prey, irritation when lodged in skin.
and transferring
sperm to the female

UNDERSIDE
The head and thorax are
fused into one body part,
which carries the legs and
mouthparts. The abdomen
has openings for breathing
and reproduction, and two
pairs of spinnerets at the rear.
270 SEA SPIDERS
These fragile-looking marine animals distant cousins of the arachnids. Most sea spiders PHYLUM ARTHROPODA
live among seaweeds in shallow seas are small—less than 3/8 in (1 cm) in body length.
INVERTEBRATES • SEA SPIDERS

CLASS PYCNOGONIDA

and coral reefs in tropical waters. The They have three or four pairs of legs and the head ORDER 1

largest species live in the deep ocean. and thorax are fused together. Instead of clawlike FAMILIES 13
SPECIES 1,348
mouthparts they have a stabbing proboscis, which
Sea spiders of the class Pycnogonida are not true they use like a hypodermic needle to suck fluids
spiders, and are so remarkably different from other from their invertebrate prey. Their spindly body
arthropods that some scientists think they belong shape means they do not need gills; they rely
to an ancient lineage that is not closely related instead on oxygen seeping directly into all their
to any groups alive today. Others think they are cells through their body surface.

3/4 in YELLOW-KNEED
2 cm GIANT SEA SPIDER SEA SPIDER
Colossendeis megalonyx Unknown sp.
f: Colossendeidae f: Callipallenidae
One of the largest of all sea
spiders, this is a deep-sea Some species of sea spiders—such as this species
animal of sub-Antarctic of Australian reefs—are strikingly colored, often as
waters, with a leg span camouflage against their colorful surroundings.
2–4 in
of 28 in (70 cm). 5–10 cm

5⁄16 in SPINY SEA SPIDER

8 mm
Endeis spinosa
f: Endeidae
This sea spider—found around
European coasts but perhaps
ranging elsewhere—has an
especially spindly body, with a 5⁄16 in
long, cylindrical proboscis. 8 mm

3⁄16 in slender body one of eight multi-

5 mm FAT SEA SPIDER GRACEFUL SEA SPIDER colored, jointed legs
Pycnogonum litorale Nymphon brevirostre
f: Pycnogonidae f: Nymphonidae
Unlike most other sea spiders, One of the most common sea
this European species has a spiders of the N.E. Atlantic,
thick body and comparatively this species occurs in the
short, curved, clawed legs. It intertidal zone and
feeds on anemones. in offshore shallows.

HORSESHOE CRABS
The Merostomata are a small class of horseshoe crabs are most closely related to PHYLUM
CLASS
ARTHROPODA
MEROSTOMATA
sea-dwelling relatives of spiders and arachnids. The leaflike gills on the underside ORDER 1
scorpions. Of prehistoric origin, they of their abdomens are the forerunners of similar FAMILY 1

are widely regarded as “living fossils.” internal structures—the so-called book lungs— SPECIES 4
that arachnids use to breathe on land. The
Horseshoe crabs were far more diverse in pedipalps are used as a fifth pair of legs, one
prehistory, when animals like them may well pair more than in spiders. Horseshoe crabs grub spiny abdomen
have been the first chelicerates on Earth. Their around in muddy ocean waters to catch prey.
clawlike mouthparts and lack of antennae They come ashore in large numbers to breed,
indicate that, despite their hard carapace, laying their spawn in sand.

long, spine-
16–231/2 in
like tail
40–60 cm

JAPANESE MANGROVE
HORSESHOE CRAB HORSESHOE CRAB
Tachypleus tridentatus Carcinoscorpius rotundicauda
f: Limulidae f: Limulidae
fused head and
This horseshoe crab spawns in sandy thorax covered Associated with muddy-sand habitats
shores of E. Asia. Habitat destruction by carapace in S.E. Asia, this species feeds on insect
and pollution have depleted its numbers larvae and other invertebrates, 16–231/2 in
in some parts of its range. as well as small fishes. 40–60 cm
C RU S TAC E A N S 271
Most crustaceans are aquatic, breathe often used for brooding young. Gills tie most PHYLUM ARTHROPODA
with gills, and have limbs for crawling crustaceans to water, although woodlice and

IN VERTEBR ATES • CRUSTACEA NS

SUBPHYLUM CRUSTACEA

or swimming. A few are sedentary as some crabs have modified breathing organs ORDERS 56

adults, parasitic, or land living. that enable them to live in damp places on land. FAMILIES About 1,000

Being permanently under water, where their SPECIES About 70,000

The basic body plan of a crustacean consists of weight is supported, means that crustaceans can
a head, thorax, and abdomen, though in many develop a thick, heavy exoskeleton. Many species D E B AT E
groups the head and thorax are fused together. have an exoskeleton hardened with minerals, C R U S TA C E A N A N C E S T R Y
In crabs, lobsters, and prawns a frontal flap grows which are reabsorbed between molts when the
backward to cover the entire head and thorax outer coating of the body is shed. With their Crustaceans have many unique
with a shell-like carapace. Crustaceans are the body buoyed up by water, many crustaceans have features, including double antennae
and two-pronged limbs, that suggest
only arthropods that have two pairs of antennae become bigger than their land-based relatives.
they are exclusive descendants of
and primitively two-pronged limbs. Limbs on The world’s largest arthropod is the deep-sea a single common ancestor. However,
the thorax are usually used for locomotion, but Japanese spider crab (leg span 13 ft/4 m). At the recent DNA analysis shows that
in some species they are modified as pincers for other extreme, crustaceans make up the bulk insects are also descended from
feeding or defense. Many crustaceans also have of animal life in plankton, either as tiny larvae the crustacean group.
well-developed abdominal limbs, which are or as adult shrimps and krill.

WAT E R F L E A S BA RNAC L ES GIANT COPEPOD

Macrocyclops albidus
A N D R E L AT I V E S AND COPEPODS f: Cyclopidae
Copepods are tiny predators
The class Branchiopoda are primarily freshwater Like other marine crustaceans, the class of plankton; this widespread
crustaceans, which make up part of the plankton that Maxillopoda start life as tiny planktonic larvae. species even preys on
flourish in short-lived pools, surviving long periods Barnacle larvae cement themselves, head down, mosquito larvae, offering
1/32–1/8 in a potential means of
of drought as eggs. Their thorax carries leafy limbs for on rocks. Most copepods stay free-swimming, 1–2.5 mm controlling these insects.
breathing and filter feeding. Water fleas are typically although some become parasites.
encased in a transparent carapace. 2–4 in DEEP SEA GOOSE BARNACLE
3⁄16–1/2 in 5–10 cm Neolepas sp.
0.5–1.5 cm
f: Eolepadidae
This goose barnacle ally lives around
volcanic vents on the ocean floor,
3/8–1/2 in where it filter-feeds on other
1–1.5 cm organisms including bacteria.
BRINE SHRIMP 2–4 in/5–10 cm
Artemia salina 1⁄16–3⁄16 in
f: Artemiidae 2–5mm COMMON ACORN GIANT ACORN BARNACLE COMMON GOOSE
This soft-bodied, stalk-eyed animal BARNACLE Balanus nubilus BARNACLE
is found worldwide in salt pools and LARGE WATER FLEA Semibalanus balanoides f: Balanidae Lepas anatifera
swims upside down. Its hard-shelled Daphnia magna f: Archaeobalanidae The world’s largest barnacle, this f: Lepadidae
eggs can resist years of drought. f: Daphniidae Sensitive to drying out, this species occurs attached to rocks Goose barnacles are attached—
This N. American water flea, intertidal species is most below the intertidal level along mostly to oceanic flotsam—by
like its relatives, can brood eggs abundant at the base of the the Pacific N. American coast. a flexible stalk. This species
inside its carapace. These hatch barnacle zone on exposed occurs in temperate waters of
without fertilization, rapidly North Atlantic rocky coastlines. the N.E. Atlantic.
populating ponds as a result. 3/4–11/4 in
3/4 in 31/4–36 in
1⁄16 in 2–3 cm 8–90 cm
1.8 cm
1.5mm

COMMON
FISH LOUSE
SEA LOUSE ASIAN ACORN BARNACLE Argulus sp.
Caligus sp. Tetraclita squamosa f: Argulidae
f: Caligidae f: Tetraclitidae This is a flattened,
MARINE WATER FLEA A representative of a group of This acorn barnacle lives in fast-swimming crustacean
Evadne nordmanni copepods that are parasitic on the intertidal zone on Indo- with an oval carapace. It
f: Podonidae marine fishes, this crustacean Pacific shores. Recent research uses suckers to attach to 3⁄16–3/8 in
Most water fleas—named for attacks salmon and related species. has led to five subspecies fishes and suck their blood. 0.5–1 cm
their jerky swimming—occur in being recognized.
stagnant freshwater pools. This,
however, is a saltwater species 1/8–3/16 in
of oceanic plankton. 3–5 mm

VERNAL POOL TADPOLE SHRIMP 2 in

Lepidurus packardi 5 cm
f: Triopsidae GLACIAL COPEPOD
Tadpole shrimp are ancient, bottom-living Calanus glacialis
crustaceans of temporary freshwater pools. f: Calanidae
Relatives of this Californian species have changed two tails Within Arctic Ocean plankton, this copepod
little in 220 million years of evolution. is an important part of the food chain.
 SEED SHRIMP 3/4–11/4 in
2–3 cm
1⁄64 –1⁄16 in
0.5–2 mm
272 Seed shrimp belong to the class Ostracoda.
The entire body of a seed shrimp is enclosed GIANT SWIMMING CRAWLING SEED
SEED SHRIMP SHRIMP
within a two-valve hinged carapace, with just Gigantocypris sp. Cypris sp.
f: Cypridinidae f: Cyprididae
IN VERTEBR ATES • CRUSTACEA NS

the limbs poking through. If threatened, the

animal can shut itself inside. These small Most seed shrimp are tiny This widespread freshwater
crustaceans crawl through vegetation in crustaceans with two-valved crustacean belongs to a
carapaces; this is a large, deep-sea group of small, hard-shelled
marine and freshwater habitats; some use species with big eyes for hunting seed shrimp that crawl
their antennae to swim. bioluminescent prey. through detritus.

C R A BS A N D R E L AT I V E S ANTARCTIC KRILL
Euphausia superba
The Malacostraca class is the most diverse crustacean group. f: Euphausiidae
Swarms of these plankton-feeding
The basic features are head, thorax, and an abdomen with 11/2–21/4 in crustaceans are critical components of
multiple limbs. Two large orders within this class include 4–6 cm the Southern Ocean food chains that
the decapods, which have a carapace that curves around the support whales, seals, and seabirds.
fused head-thorax to contain a gill cavity, and the isopods
(woodlice and relatives), which lack a carapace and are the
largest group of land-living crustaceans.

1/2–7/8 in
1.5–2.2 cm
RELICT OPOSSUM SHRIMP
Mysis relicta
f: Mysidae 3/8–3/4 in
Translucent, feathery-legged opossum shrimp 1–1.8 cm
3/8–3/4 in carry their larvae in a brood pouch. Most live in
1–2 cm
COMMON coastal waters, but this species is found in fresh
FRESHWATER SHRIMP waters in the northern hemisphere.
Gammarus pulex
f: Gammaridae abdomen
This N. European amphipod,
abundant in freshwater streams,
belongs to a family of detritus-eating
freshwater shrimp. Related species
COMMON inhabit brackish waters.
SANDHOPPER
Orchestia gammarellus
f: Talitridae SPINY SKELETON
SHRIMP
One of many amphipods Caprella acanthifera
(sideways-flattened f: Caprellidae
crustaceans), this is a Like other skeleton shrimp,
European intertidal this is a slender, slow-moving,
sandhopper—so called predatory amphipod with few
because it can jump by legs. It clings to seaweeds in
flipping its abdomen. European rockpools.
1/2 in
13 mm

COMMON WATER
3/8–1/2 in SLATER
1–1.5 cm Asellus aquaticus
f: Asellidae
A common European member
of a family of freshwater 71/2–14 in
woodlice relatives, this species 19–36 cm
crawls among detritus in GIANT DEEPSEA
stagnant water. ISOPOD
Bathynomus giganteus
f: Cirolanidae
tail fin used A giant marine relative of
for swimming woodlice, this species crawls
3/4–11/4 in along ocean floors, scavenging
2–3 cm 4–43/4 in
10–12 cm on dead animals and
segmented occasionally taking live prey.
exoskeleton

BLACK-HEADED 3/8–3/4 in COMMON PILL

WOODLOUSE 1–1.8 cm WOODLOUSE
COMMON Porcellio spinicornis Armadillidium vulgare
SEA SLATER f: Porcellionidae f: Armadillidiidae
Ligia oceanica Often living alongside Pill woodlice are
f: Ligiidae humans, especially in characterized by their
A large coastal woodlouse, lime-rich habitats, this ability to roll into a ball
this European species lives distinctly marked woodlouse when disturbed. This
in rock crevices above the is native to Europe but has species is widespread
intertidal zone. It feeds invaded N. America. across Eurasia and
on detritus. introduced elsewhere.
 2–3 in 8–12 in
5–7.5 cm 20–30 cm

SINGAPORE
8–14 in BAMBOO
20–36 cm TIGER SHRIMP

IN VERTEBR ATES • CRUSTACEA NS

PRAWN Atyopsis moluccensis
Penaeus monodon f: Atyidae
f: Penaeidae From a family of mostly STRIPE-LEG SPINY
This widely farmed, giant freshwater river shrimp, LOBSTER
Indo-Pacific prawn belongs to a this is a filter-feeder from Panulirus femoristriga
group that does not brood its S.E. Asia that eats particles f: Palinuridae
fertilized eggs, but releases them caught on its fanlike Unlike true lobsters, spiny
directly into the sea. front legs. lobsters, such as this Indo-Pacific
species, lack claws and have a heavy,
spiny exoskeleton and long,
PEACOCK MANTIS 23/4–41/4 in stout antennae.
SHRIMP 7–11 cm
Odontodactylus scyllarus
f: Odontodactylidae 11/2–2 in
Mantis shrimp of warm marine 4–5 cm
shallows are intelligent and formidable
predators. This Indo-Pacific species uses stalked eyes COMMON AESOP SHRIMP
“clubs” on its front limbs to smash
PRAWN Pandalus montagui
the shells of crabs and snails.
Palaemon serratus f: Pandalidae
f: Palaemonidae Males of this cold-water,
11/8–7 in A large, commercially important North Atlantic shrimp
3–18 cm member of a family of shrimp with a sometimes change into
jagged-edged carapace, this species inhabits females after 13–16 months.
northeast Atlantic coasts, frequenting Other individuals remain
carapace the intertidal zone. male or female throughout
their lives.

31/4–43/4 in
SPOTTED CLEANER SHRIMP 8–12 cm
Periclimenes yucatanicus
1 in f: Palaemonidae
2.5cm
This Caribbean crustacean
lives in sea anemones and—
like many other reef-
dwelling shrimp—cleans
visiting fishes of their dead
skin and parasites.

WHITE-CLAWED CRAYFISH
Austropotamobius pallipes
f: Astacidae
Crayfish are small, mostly nocturnal,
2 in freshwater relatives of true lobsters.
5 cm This is a European stream species and
clublike
limbs deliver COMMON MARBLE the only crayfish native to Britain.
powerful blows SHRIMP
Saron marmoratus
f: Hippolytidae
11/4–2 in A member of the humpback
3–5 cm shrimp family, this Indo-Pacific long, stiff
reef species is a nocturnal antennae
predator of smaller animals.
It turns red at night.
8–91/2 in
20–24 cm
COMMON SHRIMP
Crangon crangon
f: Crangonidae
This shallow-water crustacean
is commercially fished on some
European coasts. It habitually long, spiny claws
hides in sand with only its eyes
and antennae showing.

2–8 in
5–20 cm
NORWAY LOBSTER
enlarged Nephrops norvegicus
SCULPTURED SLIPPER LOBSTER antennae SERRATED SLIPPER f: Nephropidae
Parribacus antarcticus LOBSTER This North Atlantic crustacean
f: Scyllaridae Ibacus brevipes tunnels in mud, often
Relatives of spiny lobsters, clawless slipper f: Scyllaridae cohabiting with goby fishes. It is
lobsters have flattened, platelike antennae. Like other slipper lobsters, this Indo-Pacific a small-bodied true lobster,
This nocturnal species lives on tropical 2–43/4 in crustacean can swim rapidly away from commercially marketed as
reefs with sandy bottoms. 5–12 cm predators by flapping its broad tail fan. langoustine or scampi.
 CRABS 3/8 in
274 A N D R E L AT I V E S 1 cm
ROBBER CRAB
Birgus latro
f: Coenobitidae
The largest land-living arthropod,
IN VERTEBR ATES • CRUSTACEA NS

this hermit crab is a relative of

12–16 in
30–40 cm squat lobsters and inhabits
Indo-Pacific island forests,
eating coconuts using
its massive pincers.
PINK SQUAT LOBSTER
Lauriea siagiani
f: Galatheidae
Many tropical squat lobsters are
associated with specific reef
organisms. This tiny, hairy
Indonesian species lives on
Xestospongia vase sponges.

BLUE-STRIPED
SQUAT LOBSTER
Galathea strigosa
f: Galatheidae
13/4 in 23/4–31/2 in Slender-clawed squat lobsters
2 cm unequal-sized claws 7–9 cm
such as this European species are
used for signaling decapods (ten-legged), but the last
anemone
attached leg pair is reduced, making them
to shell appear eight-legged.

ANEMONE PORCELAIN CRAB

Neopetrolisthes maculatus 
f: Porcellanidae
Porcelain crabs are tiny, eight-legged
decapods more closely related to squat
lobsters than true crabs.  This
Indo-Pacific species lives in giant
Stichodactyla anemones.
5⁄16–1/2 in
8–12 mm

antenna

INDO-PACIFIC
PEA CRAB
Pinnotheres sp.
f: Pinnotheridae
Tiny pea crabs complete their life
cycle on or in the body of other
marine invertebrates. This Philippine
species lives on cup corals.

11/2 in
4 cm

5–8 in
13–20 cm
21/4–4 in
6–10 cm
RED REEF HERMIT CRAB WHITE-SPOTTED ANEMONE HERMIT CRAB
Paguristes cadenati HERMIT CRAB Dardanus pedunculatus
f: Diogenidae Dardanus megistos f: Diogenidae
Hermit crabs inhabit discarded snail f: Diogenidae This hermit crab of Indo-Pacific reefs
shells that accommodate their soft This crustacean of the E. Atlantic and always carries a Calliactis anemone
coiled abdomen. This species lives on Indo-Pacific coasts is a “left-handed” on its shell, which shares its food and
Indo-Pacific and E. Atlantic reefs. hermit crab, with an enlarged left claw. provides camouflaged protection.
 2–21/2 in
5–6.5 cm

2–23/4 in
VELVET SWIMMING BLUE 5–7 cm
CRAB SWIMMING
Necora puber CRAB
f: Polybiidae Portunus pelagicus
Swimming crabs have f: Portunidae 2–4 in
paddlelike rear legs. This This Indo-Pacific swimming crab 5–10 cm
aggressive, red-eyed species favors sandy or muddy coastlines,
is common at low shore with juveniles entering the intertidal shell-like
level on rocky coasts in zone. Like its relatives, it preys on carapace
the N.E. Atlantic. other invertebrates. 11/2–21/4 in
4–6 cm

SPOTTED 13/4–31/2 in WARTY BOX CRAB

CORAL CRAB 4.5–9 cm Calappa hepatica EDIBLE CRAB
Carpilius maculatus f: Calappidae Cancer pagurus
f: Carpiliidae A tortoiselike crab, f: Cancridae
One of several species this Indo-Pacific, The most commercially
of brightly colored, sand-burrowing species important European
coral-living crabs with many habitually shields its face crab, this offshore species
fossil relatives, this large with its claws, earning it has a distinctive wide “piecrust” carapace
species is found in the Indian the alternative name and can live for more than 20 years.
and Pacific oceans. of shame-face crab.

11/2–2 in SPONGE CRAB PAINTED PEBBLE

4–5 cm Dromia personata CRAB
f: Dromiidae Leucosia anatum
This Atlantic crab often f: Leucosiidae
carries sponge pieces to hide Belonging to a family
itself from predators. Its of small, mostly diamond-
reduced hind legs are like 3/4–11/4 in shaped crabs with long
those of the related, 2–3 cm claws, this colorful
primitive hermit crabs. species occurs in the 12–16 in
Indian Ocean. 30–40 cm

JAPANESE SPIDER
CRAB
Macrocheira kaempferi
f: Inachidae
3/8–11/8 in With legs spanning 13 ft (4 m), this is the
1–3 cm world’s biggest arthropod. It occurs in
N.W. Pacific waters and reputedly
lives for up to 100 years.

31/4–4 in
CHRISTMAS 8–10 cm
ISLAND RED CRAB PANAMIC EUROPEAN
Gecarcoidea natalis ARROW CRAB FRESHWATER
f: Gecarcinidae Stenorhynchus debilis CRAB
This land-living crab lives f: Inachoididae Potamon potamios 11/2 –2in
in burrows only in the forests of Christmas Arrow crabs are small, f: Potamidae 4 –5cm
Island. Huge numbers march in an annual stalk-eyed, spiky-headed A member of a large
migration to the sea for spawning. spider crabs. This family of Eurasian crabs from alkaline
E. Pacific species is a fresh waters, this southern European
eyes on reef-dwelling scavenger. species spends much of its time on land.
stalks
ORANGE
3/8–3/4 in FIDDLER CRAB
1–2 cm Gelasimus vocans
f: Ocypodidae
Fiddler crabs are burrowing
beach crabs with an
enlarged signaling claw
in males. This species
lives on muddy
W. Pacific shores.

2–21/4 in
5–6 cm
AFRICAN RAINBOW CRAB 4–6 in
Ocypode (Cardisoma) armatum 10–15 cm
f: Ocypodidae
This shore-dwelling W. African
species feeds on fruits, vegetation, furry CHINESE
claws MITTEN CRAB
and animals. Only females go into Eriocheir sinensis
the sea to release eggs. f: Varunidae
Native to E. Asia, this furry-clawed, burrowing
river crab was introduced to N. America and
Europe, where it has become a pest.
276 AFRICAN
R A I N B OW C R A B BUCCAL CAVITY
Setae on the outer mouthparts scrub
O cy pode (C ard isoma) food from particles of sand and pass
IN VERTEBR ATES • CRUSTACEA NS

it inward to the mandibles

ar matum
African rainbow crabs dig deep individual burrows
above the shoreline on sandy beaches, dunes, mangrove
swamps, and estuaries. They mate in or near the burrows and
females then carry the fertilized eggs under their abdomen
for 2–3 weeks before releasing them into shallow water.
Millions of floating larvae may be produced by each female
per spawn, but only a few survive to make their way back
to continue their life on land.
EYE
The eyes are on long stalks and can
be rotated in their sockets, providing
all-around vision for locating food and
avoiding predators.

SHELL
To increase in size the crab
LEG JOINT must shed its hard shell,
Leg joints move in one a process called molting.
plane, like a human knee, Water is absorbed into the
but each one moves in a body until the shell opens
different plane, making up and can be discarded,
the crab very agile. revealing a new, larger
shell beneath.

UNDERSIDE
The abdomen in males, as shown here, is
narrow. In females it is broad and wide for
carrying the egg mass.

one of eight
walking legs

setae (stiff hairs) are

sensitive to touch and
help to detect food

sharp leg tip provides

excellent dexterity
size 4–6 in (10–15 cm)
habitat Sandy and muddy shorelines 277
distribution Atlantic coast of W. Africa
diet Opportunistic feeders, mainly on
decaying plants and animals

IN VERTEBR ATES • CRUSTACEA NS

moveable dactylus
allows the claw to
operate as a pincer

BEACHCOMBER
African rainbow crabs are most
active at night when they emerge
to forage, racing back to their
burrows if disturbed. They eat
plant and animal detritus, dead
fishes, and insects, leaving numerous
little balls of sand which they have
searched through for food.

static claw
against which the
dactylus moves
unequal-sized
claw-bearing leg
used for feeding and
buccal cavity social interactions
contains six pairs
of specialized eye on thick,
mouthparts elongated stalk

six-jointed
walking leg
278 INSECTS
Insects first appeared on land more of variants based on three major body regions— PHYLUM ARTHROPODA
than 400 million years ago and today the head, thorax, and abdomen. The head, made
INVERTEBRATES • INSECTS

CLASS INSECTA

they account for more species than up of six fused segments, houses the brain and ORDERS 29

any other class on the planet. carries the major sensory organs: compound FAMILIES About 1,000

eyes, secondary light-receptive organs called SPECIES About 1.1 million

Insects have evolved diverse lifestyles and although ocelli, and the antennae. The mouthparts are
most are terrestrial, there are also numerous modified according to diet, allowing the sucking D E B AT E
freshwater species but almost none are marine. of liquids or the chewing of solid foods. HOW MANY SPECIES?
They have a number of key features, such as a The thorax is made up of three segments, each
small size, an efficient nervous system, high bearing a pair of legs. The posterior two thoracic The actual number of insect species
reproductive rates, and—in many cases—the segments usually each bear a pair of wings. The likely to exist far exceeds the
1.1 million so far described, and
power of flight, which have led to their success. legs, which are each made up of a number of
many more are discovered each year.
Insects include (among others) beetles, flies, segments, can be greatly modified to serve a However, research based on sampling
butterflies, moths, ants, bees, and true bugs. Yet variety of functions, from walking and running to in species-rich rainforests suggests
for all their diversity, insects are remarkably jumping, digging, or swimming. The abdomen, that there could be as many as
similar. Evolution has modified the basic insect which is usually made up of 11 segments, contains 10–12 million species of insects.
anatomy many times over to produce a multitude the digestive and reproductive organs.

SI LV E R F I S H long front legs held forward

These primitive, wingless insects in the order

Zygentoma have a lengthy body that is covered ⅜–1⁄2 in
front segment of thorax
with scales. The head has a pair of long 1–1.5 cm
antennae and small eyes. The abdominal
segments have small
appendages (styles).
½ in
1.2 cm

FIREBRAT
SILVERFISH Thermobia domestica
Lepisma saccharina f: Lepismatidae
f: Lepismatidae This insect is found
This common domestic worldwide and lives under large, triangular
species can sometimes stones and in leaf litter. front wings
be a nuisance in kitchens, Indoors, it favors warm
where it feeds on tiny conditions and can be pale abdomen
scraps of dropped food. a pest in bakeries.
¾–1 in
1.5–2.5 cm

M AY F L I E S
Ephemeroptera are soft-bodied insects with slender legs and
two pairs of wings. The head has a pair of short antennae
and large, compound eyes. The end of the abdomen has two
or three long tail filaments. The life cycle is dominated by
the aquatic nymphal stages—the non-feeding adults live for MAYFLY
Ephemera danica
only a few hours or days. f: Ephemeridae
Breeding in rivers
and lakes with silty
3⁄16–7⁄32 in 1⁄4–½ in 3⁄8–3⁄5 in
bottoms, this large
5–7 mm 6–12 mm 10–15 mm
mayfly is widespread
in Europe. Adults have
long antennae and
three tail filaments.

POND OLIVE BLUE-WINGED OLIVE SIPHLONURUS LACUSTRIS

Cloeon dipterum Ephemerella ignita f: Siphlonuridae
f: Baetidae f: Ephemerellidae Very common in upland lakes
A widespread European species, Adults of this N. European species have in N. Europe, this summer
this mayfly breeds in a range of three tail filaments. The males’ rounded eyes mayfly has two long tails and
habitats from ponds and ditches have two parts, the larger upper portion greenish-gray wings. The
to troughs and water butts. for spotting females. hindwings are small.
DA M SELFLIES AND AZURE DAMSELFLY
Coenagrion puella
DR AG ON F LI ES f: Coenagrionidae 279
The blue- and black-marked males
These insects in the order Odonata have a distinctive of this N.W. European species rest
long body with a mobile head and large eyes, giving wings held backward frequently on floating vegetation.
good all-around vision. The adults have two pairs when at rest Females also have black markings
but are greenish elsewhere.
of similarly sized wings and are fast-flying hunters;
the nymphs capture prey under water using their
specialized mouthparts. Dragonflies are robust with a 11/2 in
rounded head, whereas damselflies are more slender 3.5 cm
with a broader head and widely separated eyes.

21/4–31/4 in 21/4–31/4 in 11/2 in

6–8 cm 5.8 –7.8 cm 3.6 cm

TWIN-SPOTTED PRINCE BASKETTAIL EMERALD DAMSELFLY

SPIKETAIL Epitheca princeps Lestes sponsa
Cordulegaster maculata f: Corduliidae f: Lestidae each leg
f: Cordulegastridae A widespread N. American species, Common in a wide band across Europe has two claws
This dragonfly is found in the the prince baskettail can be seen and Asia, this damselfly is found near still
E. US and S.E. Canada, where patrolling ponds, lakes, creeks, and or slow-moving, well-vegetated water.
it favors clean streams in rivers from dawn until dusk.
wooded habitats.

11/2–2 in
4.2–4.8 cm

2 in 3–31/2 in 21/2–3 in
5.3 cm 7.5–8.8 cm 6.5–7.6 cm
PLAINS CLUBTAIL COMET DARNER SWIFT RIVER CRUISER
Gomphus externus Anax longipes Macromia illinoiensis
f: Gomphidae f: Aeshnidae f: Macromiidae
Flying on warm, sunny days Occurring from Brazil to This N. American species patrols
and breeding in slow-moving, Massachusetts, the comet gravelly or rocky streams
BANDED muddy streams and rivers, darner can be found over lakes and rivers, but can also be seen
DEMOISELLE this dragonfly is widespread and large ponds and has a flying away from water along
Calopteryx splendens in the US. steady, regular flight pattern. roads or tracks.
f: Calopterygidae
Males of this large damselfly from N.W. Europe have
a metallic blue-green body and blue wing bands.
Females are metallic green and lack wing markings.

wing reddish
at base

three long tail filaments

red or dark-
orange abdomen

2¾–31/4 in 1–12∕5 in 11⁄2–1¾ in 2–3 in

7–8 cm 2.4–3.4 cm 4–4.6 cm 5–7.5 cm

GRAY PETALTAIL WHITE-LEGGED DAMSELFLY BROAD-BODIED CHASER FLAME SKIMMER

Tachopteryx thoreyi Platycnemis pennipes Libellula depressa Libellula saturata
f: Petaluridae f: Platycnemididae f: Libellulidae f: Libellulidae
A large dragonfly found in damp This damselfly from C. Europe breeds This C. European and C. Asian species Favoring warm ponds, streams,
broad-leaved forests of the east coast in slow-moving, weedy canals and breeds in ditches and ponds. The abdomen and even hot springs, this
of N. America, this species breeds rivers. The tibia of the hind leg is of the male is blue above, whereas that of species is common in
in boggy areas and seeps. expanded and appears slightly feathery. the female is yellowish brown. the S.W. US.
 STONEFLIES STICK AND E A RW I G S
280 Members of the order Plecoptera are soft, LEAF INSECTS These slender, slightly flattened, scavenging
slender-bodied insects with a pair of thin These slow-moving, herbivorous insects of the order insects belong to the order Dermaptera.
tail filaments and two pairs of wings. The Pseudophasmatidae have a stick- or leaflike body that may be They have short front wings with large,
INVERTEBRATES • INSECTS

nymphal stages are aquatic. smooth or spiny. Many are well camouflaged to avoid predators. fan-shaped hindwings folded beneath. The
flexible abdomen ends in a pair of
multipurpose forceps.
TWO-STRIPED 11∕2–21∕2 in
STICK INSECT 4.2–6.8 cm
Anisomorpha buprestoides
f: Pseudophasmatidae
3∕4–1 in Found in the S. US, this
2–2.8 cm species can eject an acidic,
defensive liquid from
glands in the thorax. 1∕2–3∕5 in
PERLA BIPUNCTATA 1.2–1.5 cm COMMON EARWIG
f: Perlidae JUNGLE NYMPH
STICK INSECT Forficula auricularia
The males of this species, which favors stony Heteropteryx dilatata f: Forficulidae
streams in upland regions, have much shorter f: Heteropterygidae This species can be found under bark and
wings and can be half the size of the females. This impressive species is found in in leaf litter. The female cares for her
Malaysia. Females are flightless eggs and feeds the young nymphs.
and green; males are smaller,
4–6 in winged, and brownish.
10–15.5 cm
JAVANESE LEAF INSECT
Phyllium bioculatum
f: Phylliidae
11⁄32–1/2 in 23∕4–33∕4 in Females of this S.E. Asian insect are large,
0.9–1.3 cm 7–9.4 cm winged, and very leaflike. The males are 1⁄2–11∕4 in
TAWNY EARWIG 1.6–3 cm
YELLOW SALLY smaller, more slender, and brown. Labidura riparia
Isoperla grammatica f: Labiduridae
f: Perlodidae The tawny earwig, the largest
Particularly common in limestone areas, this large, European species, is especially
species prefers clean, gravel-bottomed streams fan-shaped common along sandy river banks
and stony lakes. Males are shorter than females. hindwing and in coastal areas.

PR AY I NG M A N T I D S large, compound eyes on a

triangular head
These predatory insects of the order Mantodea have flattened, leaflike
abdomen
a triangular, highly mobile head with large eyes. The
enlarged, spined front legs are specially modified for extended prothorax
snatching prey. A pair of toughened front wings protects
the larger membranous hindwings folded beneath.

leaflike front wing

21/4–31/2 in
6–9 cm

spiny, enlarged
front femur

head crest
2–23/4 in
5–7 cm

COMMON
PRAYING MANTIS
Mantis religiosa 11∕4–23∕4 in
f: Mantidae 3–7 cm
The strike of a praying mantis
lasts a fraction of a second, and
impales the prey on the sharp CONEHEAD ORCHID MANTIS
spines of the front legs. MANTID Hymenopus coronatus
Empusa pennata f: Hymenopodidae
f: Empusidae With its flower-mimicking color and
A slender mantid with a distinctive head legs resembling petals, this S.E. Asian
crest, this S. European species eats small mantis is able to hide among foliage
flies and can be green or brown. and ambush small insects.
CRICKETS AND ½–1 in
1.4–2.4 cm
2–23⁄4 in
5–7 cm
1∕2–1 in
1.5–2.6 cm
GRASSHOPPERS 281
Orthoptera are mainly herbivorous.
They have two pairs of wings, although
some can be short-winged or wingless.
The hind legs are often large and used
for jumping. They sing by rubbing COMMON FIELD GRASSHOPPER WELLINGTON TREE WETA AFRICAN CAVE CRICKET
their front wings together or their Chorthippus brunneus Hemideina crassidens Phaeophilacris bredoides
hind legs on a wing edge. f: Acrididae f: Anostostomatidae f: Rhaphidophoridae
This grasshopper is typically found on Native to New Zealand, this nocturnal This C. African species is an omnivorous
short, dry, grazed grassland in Europe, insect lives in rotten wood and tree stumps. scavenger. It has very long antennae, an
N. Africa, and temperate Asia, where it is It eats plant material as well as small insects. adaptation to life in dark microhabitats.
most active on sunny days.
2–3¼ in 1∕2–3∕4 in
DESERT LOCUST 5–8 cm OAK BUSH 1.3–1.8 cm
Schistocerca gregaria CRICKET
f: Acrididae Meconema thalassinum
Crowding of nymphs after rain f: Tettigoniidae
stimulates this African insect to Found on a range of
transform from a solitary into a broadleaved trees, this
11∕2 in gregarious form. It swarms in European cricket feeds
4 cm the billions, devastating crops. on small insects after dark.
The female has a long,
LEAF-ROLLING MOLE CRICKET curved ovipositor. ovipositor for
CRICKET Gryllotalpa gryllotalpa laying eggs
Hyalogryllacris subdebilis f: Gryllotalpidae
f: Gryllacrididae This species from Europe
Found in Australia, this uses its strong front legs
species has relatively long warty surface
to burrow, just like a
wings and very long miniature mole. It is found
antennae, which may be 11∕2–13∕4 in in meadows and banks near
up to three times the 3.5–4.6 cm rivers, where the soil is
length of the body. damp and sandy.

HOUSE CRICKET
Acheta domesticus
f: Gryllidae
A nocturnal species, the house
cricket makes an attractive chirping
song. Originally from S.W. Asia and 1⁄2–3/4 in
N. Africa, it has spread into Europe. 1.4–2 cm
21∕4–31∕4 in
5∕8–7∕8 in 6–8 cm
FOAMING
1.7–2.3 cm GRASSHOPPER
SOUTHERN FIELD CRICKET bright red Dictyophorus spumans
Gryllus bimaculatus markings f: Pyrgomorphidae
f: Gryllidae Bright colors advertise that this
This cricket is widespread in S. Europe, South African grasshopper is toxic
parts of Africa, and Asia and lives on the to predators. It can also produce a
ground under wood or debris. noxious foam from its thoracic glands.

COC K ROAC H E S TERMITES

Members of the order Blattodea are 2–3¼ in These nest-building social insects of the order Isoptera live
5–8 cm
scavenging insects, with a flattened oval body. in colonies with different castes: reproductives (the kings
The downward-pointing head is often largely MADAGASCAN HISSING COCKROACH and queens), workers, and soldiers. Workers are generally
concealed by a shieldlike pronotum and there Gromphadorhina portentosa pale and wingless; reproductives have wings that are shed
are usually two pairs of wings. The tip of the f: Blaberidae after a nuptial fight; and soldiers have a large head and jaws.
This large, wingless cockroach is raised worldwide
abdomen has a pair of sensory organs, or cerci. as a pet. The male has prominent bumps SUBTERRANEAN TERMITE
on the thorax for male-to-male combat. Coptotermes formosanus
short terminal cerci f: Rhinotermitidae
Native to S. China, Taiwan, and
Japan, this invasive species has now
nymph spread to other parts of the world,
where it is a serious pest.
1∕4–7⁄32 in
11∕6–13∕4 in 6–7 mm
2.7–4.4 cm
AMERICAN
11⁄32–7∕10 in COCKROACH
0.8–1.3 cm Periplaneta americana PACIFIC DAMPWOOD
DUSKY COCKROACH f: Blattidae TERMITE
Ectobius lapponicus This species, which Zootermopsis angusticollis
f: Ectobiidae was originally from 1∕4–1∕2 in f: Archotermopsidae
Small and fast-running, this European Africa, is now found 0.8–1.5 cm This termite, found along the
species can be found among leaf litter worldwide. It lives Pacific coastal states of
and occasionally in foliage. It has been on ships and in N. America, nests in and feeds on
light brown pronotum introduced into the US. food warehouses. decayed, fungus-infected wood.
282
abdomen is lined with
I N V E RT E B R AT E S • I N S E C TS

sharp-tipped spines

large, strong,
defensive spines
line insides of
hind leg

claws for
gripping and
defense
wing pads

black-tipped spines on
side of thorax and head

ADOLESCENT
The small, nonoverlapping
wing pads show that this
female is still a nymph—
not yet sexually mature.
fewer At the next molt, when it
strong sheds its skin, the nymph will
spines hind leg become an adult, acquiring
short, stubby wings and a
functional ovipositor. After
UNDERSIDE mating, the abdomen will
The female’s underside is swell as the eggs develop.
dark green and, although
tapering it has fewer spines than
abdomen the top surface, it is well
protected by the spiny legs.
JUNGLE NYMPH 283
STICK INSECT
Het eropt ery x d ilatata

I N V E RT E B R AT E S • I N S E C TS
This species is also known as the Malayan jungle nymph.
Females are large, with bright green coloration on their upper
surface and darker green below. The adult male is much
smaller, more slender, and darker in color. Both have wings,
but the female is flightless. Nymphs and adults feed on the
foliage of a range of different plants, including durian, guava,
and mango. Mature females with eggs can be very aggressive.
If disturbed, they make a loud hissing sound using their short
wings, and splay their strong, spiny hind legs in a defensive
posture. If attacked, they will kick out. This lively, nocturnal
species has become a popular pet worldwide.
size Up to 6 in (15.5 cm)
habitat Tropical forest
MOUTHPARTS distribution Malaysia
At rest the jaws are diet Foliage of various plants
concealed behind two
pairs of food-handling
appendages called
palps. These are
covered with sensory
organs, which enable
the insect to taste the
surface of the leaves
on which it feeds. COMPOUND EYE
Compound eyes—typical
of insects—do not need
to be as acute as those
of many predatory
arthropods, but they
do need to be able to
detect movement and
potential enemies.
SEGMENTED BODY
The tough, spiny plates that
make up the body segments
are joined by soft membranes
that allow flexibility.

OVIPOSITOR
The female—which can lay
up to 150 eggs in her
lifetime—uses her ovipositor
to lay her large eggs one at a
time, concealed in leaf litter
or moist soil.

long, segmented FOOT

antennae sense the The foot is made up of a number
immediate environment, of short tarsal segments and
including air movements a long, spiny terminal segment,
which bears a pair of sharp,
curved claws.
 T RU E BUGS 1/32–1⁄16 in GLASSHOUSE
WHITEFLY 1⁄8–3⁄16 in
AMERICAN
LUPIN APHID
284 Abundant and widespread in terrestrial and
1–2 mm
Trialeurodes vaporariorum 3–5 mm Macrosiphum albifrons
f: Aleyrodidae f: Aphididae
aquatic habitats, members of the order This small, mothlike Aphids such as this US
Hemiptera range from minute, wingless insects bug is found in temperate species can quickly infest
INVERTEBRATES • INSECTS

to giant water bugs capable of catching fishes and regions worldwide and can plants because females can
be a serious pest of produce many offspring
frogs. The mouthparts are used for piercing and greenhouse crops. without fertilization.
sucking up liquids such as plant sap, dissolved
prey tissues, or blood. Many species are plant distinct, dark marginal area
pests and some transmit disease.

INDIAN CICADA
5⁄16–3/8 in 11⁄32–3∕8 in
Angamiana aetherea
8–10 mm 9–11 mm
f: Cicadidae
This cicada is found in
India. As with all cicadas,
males produce loud songs
both for courtship and 11/4–11/2 in
to signal aggression. 3.5– 4 cm

11/2 in
SPITTLE BUG FROGHOPPER 35–40 mm ASSAM
Aphrophora alni Cecopis vulnerata CICADA pale area at
f: Aphrophoridae f: Cercopidae Platypleura assamensis base of hindwing
Commonly occurring on The nymphs of this f: Cicadidae
a wide range of trees and conspicuously colored Known from N. India, this species 1/4–5⁄16 in
shrubs across Europe, this European species live is also found in Bhutan and parts 6–8 mm
froghopper can vary in communally underground of China where it prefers HORSE
temperate, deciduous forests. CHESTNUT SCALE
appearance from pale to in a protective, frothy mass Pulvinaria regalis
dark brown. and feed on plant root sap. f: Coccidae
31/4 in Although commonly
8 cm found on the bark of
horse chestnut trees, this
European scale insect
will also attack a range of
other deciduous species.

3/8–1/2 in
large eyespot 1–1.2 cm
THORN BUG
Umbonia crassicornis
1/2–3⁄4 in 1/4–5⁄16 in PEANUT-HEADED BUG f: Membracidae
1.3–1.8 cm 6–8 mm Fulgora laternaria Found in C. and
f: Fulgoridae S. America, almost
LEAFHOPPER CICADELLA VIRIDIS
Ledra aurita f: Cicadellidae This bug is found in C. and the whole body of this
f: Cicadellidae S. America and the West Indies. bug is concealed
Found feeding on grasses and The bulbous head was once
Mottled coloration helps sedges in wet, boggy, or marshy under the enlarged,
thought to glow. thornlike pronotum.
this flat-bodied leafhopper areas in Europe and Asia, this
from N. Europe to blend leafhopper can also be found
with the lichen-covered near garden ponds. elongated head
bark of oak trees
in its habitat. WART-HEADED BUG
Phrictus quinquepartitus
f: Fulgoridae
11/4 in Also known as the dragon- 1⁄16 –1/8 in
3.2 cm headed bug, this species can 2–3 mm ASH PLANT LOUSE
be found in Costa Rica, Psyllopsis fraxini
distinctive marking Panama, Colombia, and f: Psyllidae
across forewings parts of Brazil. Found commonly on ash trees,
the nymphs of this species cause
the edges of leaves to form red,
swollen galls when they feed.

1/8–5⁄32 in
3–4 mm

THISTLE LACE BUG

Tingis cardui
f: Tingidae
This bug occurs in most of Europe and
brightly colored feeds on spear, musk, and marsh thistles.
hindwings The body is covered in powdery wax.
 ⅜–1/2 in ⅜–1/2 in 5⁄32 in
1–1.2 cm 1–1.6 cm 4 mm
285
long antennae

INVERTEBRATES • INSECTS
3⁄16 in
5 mm

EUROPEAN HAWTHORN COMMON BIRCH BARK BUG

TORTOISE BUG SHIELD BUG FLOWER BUG Aradus betulae 3/4 in
Eurygaster maura Acanthosoma haemorrhoidale Anthocoris nemorum f: Aradidae 1.8 cm
f: Scutelleridae f: Acanthosomatidae f: Anthocoridae The flattened body of
This bug feeds on a wide This attractive European species This predatory bug can be this European bug allows
range of grasses but can feeds on the buds and berries of found on a wide range of it to live under the bark
sometimes be a minor hawthorn and occasionally other plants. Despite its small size of birch trees, where
pest of cereal crops. deciduous trees such as oak. it can pierce human skin. it feeds on fungi.

leaflike
hind legs
5⁄32–3⁄16 in
single sharp claw 4–5mm
strong front leg
BED BUG LEAF-FOOTED BUG
Cimex lectularius Anisocelis affinis
f: Cimicidae f: Coreidae
This widespread species Found in Mexico and
feeds on the blood of C. America, the leg
humans and other expansions of this
warm-blooded mammals. herbivorous bug may
These wingless, flattened provide camouflage and
bugs are active after dark. protect it from enemies.

⅜–1/2 in 5⁄16–3⁄5 in 5⁄16–1/2 in

1–1.2 cm 8–15 mm 8–11 mm

COMMON POND WATER BOATMAN TOAD BUG

SKATER Corixa punctata Nerthra grandicollis
Gerris lacustris f: Corixidae f: Gelastocoridae
f: Gerridae Using its powerful, oarlike The warty surface and drab
Widespread and immediately rear legs for swimming, coloring of this African bug
recognizable as it darts around this common European allow it to hide in mud and
on the surface of water, the pond species feeds on algae and debris, where it ambushes
skater locates prey by the ripples detritus in ponds. insect prey.
it sends out.

⅜–1/2 in ⅜–7⁄10 in
1–1.5 cm 1–1.3 cm
SAUCER BUG
Ilyocoris cimicoides
f: Naucoridae
The saucer bug traps air from the
surface under its folded wings
and hunts prey in the shallow
margins of lakes and slow rivers. 1/4 in
31/4–4 in This species is from Europe. 6 mm
8–10 cm

hair-fringed GIANT WATER BUG WATER MEASURER COMMON GREEN CAPSID

hind leg Lethocerus sp. Hydrometra stagnorum Lygocoris pabulinus
f: Belostomatidae f: Hydrometridae f: Miridae
Using its strong front legs and This slow-moving European bug This widespread bug can be
toxic saIiva, this tropical bug can lives around the margins of ponds, a serious pest to a range of
pair of appendages acts overpower larger vertebrate lakes, and rivers and feeds on plants, including fruit crops such
as breathing siphon prey such as frogs and fishes. small insects and crustaceans. as raspberries, pears, and apples.
 T RU E BUGS
286
5⁄16–3/8 in strong front legs
8–10 mm for holding prey
FIRE BUG
INVERTEBRATES • INSECTS

Pyrrhocoris apterus
1⁄4–5⁄16 in
6–8 mm
f: Pyrrhocoridae
This unmistakable black and red bug
is a gregarious, flightless species
widespread in C. and S. Europe.
It feeds on seeds. ¾–11/2 in
spine at side 2–4 cm
of thorax

1/2 in 3⁄8–1/2 in
1.2–1.4 cm 1.1–1.4 cm

false “eye” spot

on front wing

COMMON GREEN FOREST BUG

SCARLET
SHIELD BUG Pentatoma rufipes
SHIELD BUG Palomena prasina f: Pentatomidae
Eurydema dominulus f: Pentatomidae Found on a range of deciduous
f: Pentatomidae This bug is very common and trees, this European species, reddish
widespread across Europe. It recognizable by its blunt- orange
This European species, which leg band
can be red or orange, feeds on feeds on a wide range of plants shouldered appearance, feeds
plants of the cress family and and can be a minor pest. on sap and small insects.
can be a pest of brassicas.

WHITE-SPOTTED
11/4–11/2 in ¾–1 in 1⁄2–¾ in ASSASSIN BUG
3–3.5cm 1.8–2.2 cm 1.3–1.7 cm Platymeris biguttata
f: Reduviidae
Like all assassin bugs, this African
long, hair-fringed species has toxic saliva, but unlike
hind legs used for many it can spit its saliva, causing
swimming temporary blindness.

WATER STICK INSECT WATER SCORPION COMMON BACKSWIMMER

Ranatra linearis Nepa cinera Notonecta glauca
f: Nepidae f: Nepidae f: Notonectidae
Using its specialized, long This aquatic bug crawls Found in ponds, lakes, canals, and
front legs to catch prey, around in the margins of ditches in Europe, this species is
including small fishes, this shallow pools to catch small capable of taking vertebrate prey
slender bug favors deeper, prey. It breathes through a such as tadpoles and small fishes.
well-vegetated pools. long respiratory tail. The underside faces up as it swims.

PA R A SI T IC L IC E
3⁄16 in
5 mm

These wingless insects of the order Phthiraptera are ectoparasites,

living on the bodies of birds and mammals. Their mouthparts are
modified for chewing skin fragments or sucking blood, and their
legs for gripping tightly to hair or feathers.

1⁄10–1/8 in HUMAN 1⁄10–1/8 in CHICKEN 1/32–1⁄16 in

2.5–3 mm BODY LOUSE 2.5–3 mm BODY LOUSE 1–2 mm
Pediculus humanus humanus Menacanthus stramineus
HUMAN HEAD LOUSE f: Pediculidae f: Menoponidae GOAT LOUSE
Pediculus humanus capitis This subspecies may have Found worldwide as an Damalinia caprae
f: Pediculidae evolved from the head louse ectoparasite of chickens, f: Trichodectidae
This louse glues its eggs, after the invention of clothing, infestations of this pale, Found worldwide on
called “nits,” to head hair. to which it glues its eggs. flattened chewing louse can goats, this biting louse
Infestation outbreaks are It transmits disease. cause feather loss and infection. can also survive for
common among school a few days on sheep
children. A related species but is not able to
attacks chimpanzees. breed on them.
BARKLICE THRIPS ALDERFLIES AND
AND BOOKLICE Members of the order Thysanoptera DOBSONFLIES 287
Common on vegetation and in litter, insects of are tiny insects, typically with two The Megaloptera have two pairs of wings, which
the order Psocoptera are small, squat, and pairs of narrow, hair-fringed wings. are held rooflike over the body when they are at

INVERTEBRATES • INSECTS
soft-bodied. The head has threadlike antennae They have large compound eyes and rest. The aquatic larvae, which are predatory
and bulging eyes. They eat microflora and distinctive mouthparts for piercing and have abdominal gills, pupate on land
some species are pests of stored products. and sucking. in soil, moss, or rotting wood.
FLOWER THRIP 1/32–1/24 in EASTERN DOBSONFLY
Frankliniella sp. 1–1.5 mm Corydalus cornutus
5∕32–1/4 in f: Thripidae f: Corydalidae
4–6 mm Flower thrips are found all Found in N. America, the male
over the world and can be pests on crops such of this species has very long
as peanuts, cotton, sweet potatoes, and coffee. mandibles used for combat and
for gripping the female.
PSOCOCERASTIS
GIBBOSA
f: Psocidae 1⁄64–1/24 in SNAKEFLIES
0.6–1.5 mm
Native to Europe and parts Snakeflies, in the order Raphidioptera, are 4–51⁄2 in
of Asia, this relatively large woodland insects with a long prothorax, a 10–14 cm
bark louse can be found on
a wide range of deciduous broad head, and two pairs of wings. Adults
and coniferous trees. and larvae eat aphids and other soft prey.

RAPHIDIA NOTATA ALDERFLY

LIPOSCELIS LIPARIUS f: Raphidiidae Sialis lutaria 1/2–1 in
f: Liposcelididae Found in deciduous or f: Sialidae 1.4–2.6 cm
This very widespread species coniferous woodlands in The female of this widespread
favors dark, damp microhabitats Europe, this snakefly is species lays a mass of up to
and can be a pest in libraries and 1/2–3/4 in usually associated with oak 2,000 eggs on twigs or leaves
granaries if the humidity is too high. 1.6–1.8 cm trees, where it eats aphids. near water.

L AC E W I NGS
A N D R E L AT I V E S
Insects of the order Neuroptera have conspicuous
eyes and biting mouthparts. The pairs of net-veined
wings are held rooflike over the body at rest. The 1/2 in 11/2 in
larvae have sickle-shaped mouthparts, which form 1.4 cm 4 cm
sharp, sucking tubes.
11/4 in MANTISPA STYRIACA SPOON-WINGED
3 cm f: Mantispidae LACEWING
Like a miniature praying mantid, Nemoptera sinuata
this species, found in S. and f: Nemopteridae
C.Europe, lives in lightly wooded Common to areas of S.E.
areas, where it hunts small flies. Europe, this delicate species
feeds on nectar and pollen
at flowers in woodland and
OWLFLY open grassland.
Libelloides macaronius
f: Ascalaphidae 3/8–1/2 in
1–1.2 cm
Capturing insect prey in midair, this
species will only fly on warm sunny GREEN
days. It can be found in C. and S. Europe LACEWING
and parts of Asia. Chrysopa perla
f: Chrysopidae
spotted wings This widespread European species has
a characteristic bluish-green tinge and
GIANT LACEWING black markings and is commonly
Osmylus fulvicephalus found in deciduous woodland.
f: Osmylidae
This European species can
be found among shady
woodland vegetation by
streams, where it eats
small insects
and pollen.

ANTLION 2–21/4 in
Palpares libelluloides 5–5.5 cm
f: Myrmeleontidae
This large, day-flying, Mediterranean species with males have organs for
1/2 in distinctively mottled wings can be found in rough clasping females
1.3 cm grassland, warm scrubby habitats, and sand dunes.
 BEETLES
288 Members of the order Coleoptera—the largest
insect order—range from minute to very large
species. A distinguishing feature are the
INVERTEBRATES • INSECTS

toughened front wings, called elytra, which elongated

meet down the body’s midline and protect the head
larger membranous hindwings. Beetles occupy
every aquatic and terrestrial habitat, where they VIOLET
are scavengers, herbivores, or predators. GROUND BEETLE
Carabus violaceus
f: Carabidae
1∕8–3∕16 in A nocturnal hunter, this beetle 1–12∕5 in
3–5 mm is common in many habitats, 2.8–3.4 cm
including gardens. It is native to
Europe and parts of Asia.

11∕2 in
4 cm
5⁄16–3⁄8 in
8–10 mm

WOODWORM BEETLE JEWEL BEETLE COMMON RED

Anobium punctatum Chrysochroa chinensis SOLDIER BEETLE
f: Anobiidae f: Buprestidae Rhagonycha fulva
Adapted to breeding in wood This metallic species is native f: Cantharidae VIOLIN BEETLE
in buildings and furniture, to India and S.E. Asia, where A common sight on flowers in Mormolyce phyllodes
this species is now widespread its larvae burrow into the the summer, this European f: Carabidae
and can be a serious pest. wood of deciduous trees. beetle can be found in meadows The shape of this S.E. Asian
and woodland margins. beetle allows it to squeeze
under bracket fungi and tree
bark, where it feeds on 21∕4–4 in
7∕32–3∕8 in 6–10 cm
7–10 mm insect larvae and snails.
1–11∕2 in
2.5–3.8 cm
wide, flat
elytron
elytra with 3∕16–5∕16 in
metallic sheen 5–8 mm

ANT BEETLE
Thanasimus formicarius 5∕16–3∕8 in 11∕4 in
f: Cleridae 8–10 mm 3 cm
Associated with coniferous
trees in Europe and N. Asia,
the larvae and adults of LARDER BEETLE GREAT DIVING BEETLE CLICK BEETLE WHIRLIGIG
this beetle prey on bark Dermestes lardarius Dytiscus marginalis Chalcolepidius limbatus BEETLE
beetle larvae. f: Dermestidae f: Dytiscidae f: Elateridae Gyrinus marinus
Found in Europe and parts of This large beetle lives This beetle is found in f: Gyrinidae
Asia, this beetle feeds on in weedy ponds and lakes woodland and grassland in the A common European species,
animal remains but also lives in Europe and N. Asia. warmer parts of S. America. Its this beetle is a surface-
in buildings, where it eats It eats insects, frogs, larvae are predators in rotten dweller on ponds and lakes.
stored produce. newts, and small fishes. wood and soil. It uses its paddle-shaped legs
to skim over the water.

5∕16–3∕8 in 1 in 1∕4–3∕8 in 1∕2–3∕4 in curved horns

8–11mm 2.5 cm 6–10 mm 1.5–2 cm

male has wings

SCREECH BEETLE COMMON GLOWWORM FOUR-SPOTTED HISTER BEETLE MINOTAUR BEETLE

Hygrobia hermanni Lampyris noctiluca Hister quadrimaculatus Typhaeus typhoeus
f: Hygrobiidae f: Lampyridae f: Histeridae f: Geotrupidae
This European beetle feeds on small This beetle is found across Europe and Asia, Widespread across Europe, this beetle This species is found in sandy areas
invertebrate prey in slow-moving where it favors rough grassland. The is found in dung and sometimes carrion, in W. Europe and buries sheep and
rivers and muddy ponds. It makes a wingless female—known as a glowworm— where it feeds on small insects rabbit droppings, on which the
high-pitched grating noise if handled. emits a greenish light to attract a mate. and their larvae. larvae feed, in burrows.
 yellow
antennal tip
1∕8–5⁄32 in
3–4 mm
3⁄16–5⁄16 in
5–8 mm
289
1 in 21∕4–61∕2 in
2.6 cm 6–17 cm

INVERTEBRATES • INSECTS
red parallel-
sided elytra

POWDER POST BEETLE SEXTON BEETLE HERCULES BEETLE

Lyctus opaculus Nicrophorus investigator Dynastes hercules 21∕4–4 in
f: Bostrichidae f: Silphidae f: Scarabaeidae 5.5–10 cm
This N. American This species is found all over the The largest species of rhinoceros
beetle breeds in dry northern hemisphere in woodland beetle (genus Dynastes), this beetle
wood and reduces it to and grassland. It buries the carcasses feeds on rotting fruits in
a fine powder. of small animals and the female lays an rainforests in C. and S. America.
egg on it, providing food for the larva. Its larvae breed in decaying wood. GOLIATH BEETLE
massive
Goliathus cacicus
mandibles
f: Scarabaeidae
NET-WINGED BEETLE This species, the world’s heaviest insect,
Platycis minutus lives in equatorial Africa, where the
f: Lycidae adults feed on ripe fruits or tree sap.
This small beetle is associated
with decaying wood in mature antennal club
and ancient woodland
across Eurasia.

3 in
7.5 cm
3∕8–3⁄4 in
1–2.1 cm

STAG BEETLE PHANAEUS DEMON

Lucanus cervus f: Scarabaeidae
f: Lucanidae Native to C. America,
This impressive beetle lives in this very variable beetle
woodland in S. and C. Europe. breeds in the dung of large
Its larvae develop over 4–6 herbivorous animals in
years, mainly in decaying 5∕32–1⁄4 in grassland and pasture.
oak stumps. Males use their 4–6 mm
enlarged, antlerlike mandibles
in contests over females. SAP BEETLE
Glischrochilus hortensis
f: Nitidulidae
Often found feeding at
fermenting sap flows or ripe golden
DEVIL’S COACH fruits, this W. European member iridescence
HORSE of the pollen beetle family is
Staphylinus olens associated with decaying trees
f: Staphylinidae such as birch.
A European species found
among woodland and garden
leaf litter, this beetle raises
its abdomen in a threatening
display if disturbed.
3∕4–11∕10 in
2–2.8 cm

EMUS HIRTUS
f: Staphylinidae
Native to S. and C. Europe, this hairy
rove beetle feeds on other insects
attracted to cow and horse midline
manure or carrion. between
elytra

3∕4–11∕4 in
2–3.2 cm hind leg claw
GOLD BEETLE
Chrysina aurigans
f: Scarabaeidae
Found in Costa Rica and Panama,
long, segmented this beetle lives in deciduous forests,
antennae usually in rotting logs. The adults are 3∕4–13∕8 in
nocturnal and attracted to light. 2–3.5 cm
 BEETLES
290
INVERTEBRATES • INSECTS

3∕8–½ in 1–11∕2 in 2–21∕4 in

1–1.2 cm 2.4–3.4 cm 5–6 cm
1∕8 in ANTLIKE BEETLE THICK-LEGGED BLACK OIL BEETLE
3 mm Anthicus floralis FLOWER BEETLE Meloe proscarabaeus
f: Anthicidae Oedemera nobilis f: Meloidae
Found across Europe and parts of Asia, f: Oedemeridae This European beetle is found in
this small beetle lives in and around Often seen feeding on pollen at warm meadows, heaths, and
decaying vegetation, manure, and flowers in meadows, this beetle coastal areas. The larvae develop
compost heaps. is native to S.W. Europe. Its larvae inside the nests of wild bees.
develop inside plant stems.

1∕2–3∕4 in ½–3⁄5 in
1.8–2.2 cm 1.4–1.6 cm

RED-SPOTTED
LONGHORN BEETLE
Batocera rufomaculata
f: Cerambycidae
Also known as the mango
borer or fig borer, this
tropical species can be found
throughout India and
GIBBIFER CARDINAL BEETLE S.E. Asia.
CALIFORNICUS Pyrochroa coccinea
f: Erotylidae f: Pyrochroidae
This species is found in This distinctive and somewhat
moist woodland in the flattened European beetle is
S.W. US. The larvae feed often seen crawling on old PHOSPHORUS
on fungi growing on trees tree stumps, inside which its VIRESCENS
and decaying wood. larvae develop. f: Cerambycidae
Native to W. Africa, the
larvae of this beetle burrow
into some economically
important species such as
large cephalic horn the cola tree.
long, feathery 11∕4–11∕2 in
antennae 2.8–3.6 cm

11/4–1½ in 11∕2–21∕4 in
3–3.5 cm 4.2–5.5 cm

WALLACE’S
LONGHORN BEETLE
Cyriopalus wallacei
f: Cerambycidae
This impressive beetle
can be found in the
rainforests of S.E. Asia,
where its larvae bore into
certain trees.
spines on 1½–2 in
leg joint 3.2–5 cm

DICRONORHINA
DERBYANA
f: Cetoniidae
TAURHINA POLYCHROUS This beetle is found in
f: Cetoniidae southern and eastern parts of
Native to E. Africa, especially Tanzania, this Africa, where the larvae feed
beetle exists in a number of color forms. The blue on decaying plant matter
form is considered to be a subspecies. and animal dung.
 291
11∕4–11∕2 in 1∕4–5∕16 in 7∕32–11∕32 in 3∕16–5∕16 in

INVERTEBRATES • INSECTS
3–3.5 cm 6–8 mm 7–9 mm 5–8 mm

SCARLET LILY BEETLE EYED LADYBUG SEVEN-SPOT LADYBUG

Lilioceris lilii Anatis ocellata Coccinella septempunctata
f: Chrysomelidae f: Coccinellidae f: Coccinellidae
Originally from Europe and This ladybug is native to Europe and This species is very common in a
Asia, this striking pest has is associated with coniferous trees, wide range of habitats across
spread to other parts of the especially spruce and pine, where Europe, and is now established
world where lilies are grown. it feeds on aphids. in N. America.

1∕8–3⁄16 in 5∕32–3⁄16 in 1∕8–5∕32 in

3–5 mm 4–5 mm 3–4 mm

JEWELED FROG BEETLE

Sagra buqueti
f: Chrysomelidae
This beetle develops inside certain BROAD BEAN WEEVIL LARCH LADYBUG TWENTY-TWO SPOT
large-stemmed vines in S.E. Asia, Bruchus rufimanus Aphidecta obliterata LADYBUG
particularly Thailand. Like many f: Chrysomelidae f: Coccinellidae Psyllobora vigintiduopunctata
attractive tropical species, they This species is a serious pest of This European species lives on f: Coccinellidae
are sold to collectors. broad bean crops. The adults conifers such as larch, fir, and pine, This European species lives on
feed on the pollen and their where it feeds on scale insects and low-growing plants in meadows
larvae tunnel into the seeds. conifer aphids. and, unlike most ladybugs, feeds
on fungi such as mildews.

3∕4–11∕4 in BAPTISIA WEEVIL

2–3 cm Trichapion rostrum elongated neck used for
f: Brentidae rolling leaves
This small weevil feeds on the seeds of white
and cream wild indigo (Baptisia) plants on the
prairies of N. America. 1 in
2.6 cm
1∕10–5∕32 in
2.5–4 mm GIRAFFE-NECKED WEEVIL
Trachelophorus giraffa
f: Attelabidae
This beetle lives in the rainforests of
Madagascar. It rolls the leaves of the
Dichaetanthera tree with its neck, and the
female lays her egg inside the roll.

3∕8–½ in 7∕10–3∕4 in
1–1.3 cm 1.8–2 cm

3∕4–1 in
2–2.4 cm
CRATOSOMUS
RODDAMI
f: Curculionidae
This species lives in C. and
S. America. Its larvae are
stem and bark borers of CHURCHYARD BEETLE
certain wild fruit trees. Blaps mucronata
f: Tenebrionidae
ONYMACRIS This flightless, ground-living species
CANDIDIPENNIS is usually active after dark and can
claw for f: Tenebrionidae be found in dark and damp places,
gripping Long legs and white wing cases help this where it eats decaying matter.
diurnal beetle survive in the arid deserts
along the southwestern coast of Africa.
BLUE BANDED WEEVIL
Eupholus linnei BLACK VINE WEEVIL
f: Curculionidae 5∕16–½ in Otiorhynchus sulcatus
Found on certain islands in 8–12 mm f: Curculionidae
E. Indonesia, this spectacular Widespread in Europe, N. America, and
weevil, like many others in the Australasia, this weevil is a serious pest of a
genus Eupholus, feeds on yams. wide range of garden and cultivated plants.
 SCORPIONFLIES FLEAS
292 These predatory insects of the order Mecoptera Members of the order Siphonaptera, fleas are
3/4–1 in
generally have an elongated and cylindrical body. 1.8–3 cm laterally flattened, wingless, blood-sucking
The large, narrow wings may be clear or ectoparasites of mammals and some birds.
INVERTEBRATES • INSECTS

marked with spots or bands of dark color. Their head has short, piercing mouthparts
Some species are short-winged or completely and pairs of simple lateral eyes. The hind legs
wingless. The head, which has large eyes and are highly specialized for jumping.
threadlike antennae, is characteristically
extended downward in the form of a beak, COMMON
SCORPIONFLY CAT FLEA
at the end of which are biting mouthparts. Panorpa communis Ctenocephalides felis
f: Panorpidae f: Pulicidae
SNOW SCORPIONFLY Native to W. Europe, This species is very
1/8–3⁄16 in this insect lives in shady common worldwide.
Boreus hyemalis 3–5 m
f: Boreidae hedgerows and woodland Also found on dogs, it
The adult of this small, margins and can often be can feed on the blood of
seen resting on nettles. 1⁄32–1⁄16 in several other animals,
wingless European insect 1–2 m
can only be seen in the fall too—including humans.
and winter. It breeds
among moss.

T RU E F L I E S proboscis
for stabbing 5⁄32–3⁄8 in
True flies belong to the order Diptera, and are recognized and sucking 4–10 m
by a single pair of membranous front wings. The
hindwings are reduced, forming a pair of balancing MOSQUITO
Culex sp.
organs called halteres. Most flies are beneficial to humans f: Culicidae
as pollinators, predators, and recyclers, but wild and There are more than 1,000
domestic animals and many people are affected by species in this genus worldwide.
fly-borne diseases. Some flies are also crop pests. While some are disease vectors,
many are essential in food chains.

1/8 in
3 mm
7⁄32–1/2 in
7–13 m 3/8–1/2 in
1–1.4 cm

AGROMYZA RONDENSIS
f: Agromyzidae haltere
This small fly can be found
all over Europe. The larvae
make leaf mines in a wide
variety of grass species.
BEE FLY GIANT BLUE lateral ST. MARK’S FLY
Bombylius major ROBBER FLY hair tuft Bibio marci
f: Bombyliidae Blepharotes splendidissimus f: Bibionidae
This species is widespread f: Asilidae This common, springtime
in temperate regions of the This large fly is found in 1–11⁄4 in European fly can sometimes
northern hemisphere. Adults E. Australia. It catches quite large 2.5–3 c be seen in great numbers,
are nectar-feeders and their prey on the wing and flies with flying clumsily and mating on
larvae are parasites in bee nests. a distinctive buzzing sound. grass and low vegetation.

3/8–1⁄2 in 1/32–1⁄16 in
1–1.2 cm 1–2 mm

3⁄16 –3/8 in 5⁄16–3/8 in

PLATYURA 5–10 mm 8–11 mm plumelike
MARGINATA antennae
f: Keroplatidae
Widespread throughout DASINEURA SISYMBRII CABBAGE ROOT FLY BUZZER MIDGE
W. Europe, this fly lives f: Cecidomyiidae Delia radicum Chironomus plumosus
in wooded areas. The This tiny fly lays eggs on the f: Anthomyiidae f: Chironomidae
larvae live inside rotting flower heads of field cress. This small European fly can be a serious This fly is found all across the northern
wood, where they eat A pale, spongy gall develops, pest of wild and cultivated brassicas hemisphere. The larvae are called bloodworms
other small insects. inside which the larvae feed. such as cabbage, turnip, and rape. and live in muddy-bottomed ponds.
 HOUSE FLY prominent 3⁄4–1 in
Musca domestica red eyes 2–3 cm
f: Muscidae 293
This species is found worldwide
and is the most common fly
around dwellings, where it long legs can be

INVERTEBRATES • INSECTS
transmits a number of disease- shed if caught
5⁄32–1⁄4 in causing organisms to foodstuffs.
4–6 mm

GRASS FLY
Meromyza pratorum MARSH
f: Chloropidae CRANE FLY
This fly occurs across the northern Tipula oleracea
hemisphere, especially in sandy f: Tipulidae
coastal regions. The larvae burrow in Originally from Europe, this
stems of marram grass and reed grass. crane fly has been introduced
to N. America and now some
upland parts of S. America.
1⁄4–5⁄16 in
It is often found by water.
6–8 mm red-orange
wing base

1/4–7⁄32 in 5⁄32–1/4 in
6–7 mm 4–6 mm

POECILOBOTHRUS LESSER HOUSE FLY

NOBILITATUS Fannia canicularis
f: Dolichopodidae f: Fanniidae 5⁄32 in
This European species can be Able to breed in almost any 4 mm
found in damp habitats near decaying, semiliquid matter, this
water. Males display in sunny species is particularly associated
patches by waving their wings. with human habitation.

1/8–3⁄16 in 1/2–2/3 in
3–5 mm 1.4–1.8 cm

SIMULIUM
ORNATUM
f: Simuliidae
This small fly lives in Europe
and Asia but has been
introduced elsewhere. The
adults feed on animal blood
and can transmit bovine
MOTH FLY FLESH FLY onchocerciasis, a disease
Clogmia albipunctata Sarcophaga carnaria commonly known as
f: Psychodidae f: Sarcophagidae river blindness.
This very widespread Widely distributed
fly species resembles through Europe and Asia,
a tiny moth. The larvae this fly feeds on nectar
breed in dark, wet places and liquids from rotting
such as drains, tree matter. The female lays
holes, and sewage. small larvae on carrion. males with longer
eyestalks win fights
for territory

FERRUGINOUS ROTHSCHILD’S ACHIAS

BEE-GRABBER Achias rothschildi
Sicus ferrugineus f: Platystomatidae
f: Conopidae Males of this Papua New Guinean
5⁄16–1/2 in This European fly lays eggs in the fly have very elongate eyestalks,
8–14 mm
abdomen of certain bumble bees. which are used in territorial and 1/2–2/3 in
The larvae develop as internal mating displays. 1.5–1.8 cm
parasites, killing the host.

pale face with large,

compound eyes

1⁄32–1⁄16 in 3/8–1/2 in
1–2 mm 1–1.2 cm

FARMYARD MIDGE BLUEBOTTLE

Culicoides nubeculosus Calliphora vicina
f: Ceratopogonidae f: Calliphoridae
This widespread European fly breeds in Found in Europe and N. America, this
mud contaminated by dung or sewage. species is especially common in urban areas
Adults suck the blood of horses and cattle. and breeds in dead pigeons and rodents.
 T RU E F L I E S
294

1⁄8 –3⁄16 in
YELLOW DUNG FLY 3–5 mm 1∕16–1∕8 in
Scathophaga stercoraria ½–3∕5 in
1.3–1.5 cm 2–3 mm
f: Scathophagidae
This very common fly is found BANDED BROWN HORSE FLY SEPSIS SP. COMMON FRUIT FLY
in many parts of the northern hemisphere, Tabanus bromius f: Sepsidae Drosophila melanogaster
where it breeds in cattle and horse dung. The f: Tabanidae f: Drosophilidae
larvae feed on the dung, while the adults prey These flies are common and
Widespread in Europe and the widespread in a wide variety A common laboratory
on other insects attracted to the dung. Middle East, this fly mainly attacks animal, this widespread
of habitats. The larvae
horses but will feed on some other develop in animal dung species has a distinctive dark
animals, including humans. and rotting matter. patch on the abdomen and
breeds in rotting fruits.
1∕4–3∕8 in
6–11 mm fleshy, toothed
proboscis for
attacking prey 3⁄8–½ in
1–1.2 cm

MARSH
SNIPE FLY
Rhagio tringarius
f: Rhagionidae
This predatory fly can
be found on low 1∕4– 3∕5 in
vegetation growing on 8–14 mm
damp scrubby or boggy SYRPHUS RIBESII
ground throughout f: Syrphidae
much of Europe. The larvae of this nectar-
feeding fly are major predators
of aphids in the northern
hemisphere. Its coloration
mimics that of wasps and
bees to deter predators.

11∕32–½ in 3∕8–½ in 3∕5–3∕4 in

0.9–1.2 cm 1–1.2 cm 1.5–2 cm

MARMALADE LEUCOZONA DRONE FLY

HOVERFLY LEUCORUM Eristalis tenax
Episyrphus balteatus f: Syrphidae f: Syrphidae
f: Syrphidae This northern hemisphere This European fly, which
Very common in a variety species can be found visiting has been introduced to
of habitats including gardens, this flowers in damp woodlands in N. America, is a good mimic
European fly feeds on pollen and the spring and early summer. of the honey bee. The larvae
nectar. The larvae eat aphids. Its larvae prey on aphids. breed in stagnant water.

5∕16 in 11∕2–23∕4 in
8 mm 3.5–7 cm

5∕32–3⁄16 in
4–5 mm

yellow, bristly
body

FOREST FLY PENICILLIDIA FULVIDA GAUROMYDAS HEROS

Hippobosca equina f: Nycteribiidae f: Mydidae
f: Hippoboscidae Widespread in sub-Saharan This S. American insect, the
Occurring mainly in wooded Africa, this bloodsucking, world’s largest fly, breeds inside
areas in Europe and parts of Asia, wingless, ectoparasitic fly the nests of leaf-cutter ants,
this bloodsucking fly attacks can be found on a wide where the young are thought to
horses, deer, and sometimes cattle. range of bat species. eat scarab beetle larvae.
 295
1–11⁄4 in 5∕32–1∕4 in
2.4–3 cm 4–6 mm
sensory bristles
for detecting

INVERTEBRATES • INSECTS
air movement
RODENT BOT FLY
Cuterebra fontinella
f: Oestridae SNAIL-KILLING FLY
This parasitic fly lives in N. America.
Pherbellia cinerella
The larvae develop inside the bodies of
f: Sciomyzidae
rodents such as white-footed mice. This species is found in damp
meadows and marshy areas
across Europe and parts of
1∕5–3⁄16 in Asia. The larvae prey on
4.5–5.5 mm snails of the family Helicidae.

striped
thorax
3∕8–½ in
1–1.2 cm
APPLE MAGGOT
Rhagoletis pomonella
f: Tephritidae
Native to N. America, this fly is an apple
pest but also attacks hawthorn, plums,
cherries, and other related fruit trees.

1∕4– 3∕5 in 11∕4–2 in

8–15 mm 3–5 cm smoky wings

SAVANNAH PANTOPHTHALMUS DANCE FLY

TSETSE FLY BELLARDII Empis tessellata
Glossina morsitans f: Pantophthalmidae f: Empididae
f: Glossinidae Found in the rainforests Native to Europe and Asia, this fly
Confined to parts of Africa, this fly is one of the of Costa Rica, Venezuela, and can be found in meadows and
species that transmits the trypanosome parasite Ecuador, the larvae of this hedgerows, where it feeds on
that causes sleeping sickness in humans. broad body large fly feed inside tree roots. nectar and other insects.

CADDISFLIES
Closely related to the Lepidoptera, these slender,
rather mothlike insects of the order Trichoptera
are covered with hairs, not scales. The head has
long, threadlike antennae and weakly developed 1∕8–1∕5 in 11∕8–11∕4 in
mouthparts. The two pairs of wings are held tent- SALT AND PEPPER MICROCADDIS 3–4.5 mm GREAT RED SEDGE 2.8–3.2 cm
like over the body at rest. Caddisfly larvae are aquatic Agraylea multipunctata Phryganea grandis
and often construct species-specific portable f: Hydroptilidae f: Phryganeidae
This small caddisfly is widespread and common This European caddisfly breeds in weedy
shelters out of small stones or plant fragments. in N. America. It has narrow wings and lakes and slow rivers. The larvae make a case
breeds in algae-rich ponds and lakes. from cut leaf sections arranged in a spiral.

1∕2–3⁄5 in 1∕4–½ in DARK-SPOTTED SEDGE

1.2–1.5 cm 8–13 mm Philopotamus montanus
f: Philopotamidae
This European caddisfly has shortish
antennae and breeds in fast-flowing rocky
streams. The larvae make tubelike nets on
the undersides of rocks.

1∕2–3⁄5 in
1.2–1.7 cm

MOTTLED SEDGE
MARBLED SEDGE Glyphotaelius pellucidus
Hydropsyche contubernalis f: Limnephilidae
f: Hydropsychidae This European caddisfly breeds in
This caddisfly flies in the evening and lakes and small ponds. The larvae
breeds in rivers and streams. The aquatic make a case out of fragments of
larvae weave a net to catch food. withered tree leaves.
 MOTHS AND 4–61/2 in 4–61⁄2 in
296 BUTTERFLIES 10–16 cm 10–16 cm

Members of the order Lepidoptera are

covered with minute scales. They have large
INVERTEBRATES • INSECTS

compound eyes and a feeding tube called

a proboscis. Larvae, known as caterpillars,
pupate, metamorphosing into their adult form.
Most moths are nocturnal and rest with their
wings open; the majority of butterflies are POLYPHEMUS MOTH
diurnal and rest with their wings closed. AMERICAN Antheraea polyphemus
MOON MOTH f: Saturniidae
Actias luna Common and widespread
HERCULES MOTH 8–101/2 in f: Saturniidae in the US and S. Canada,
Coscinocera hercules 20–27 cm This lime-green N. American moth this moth has large
f: Saturniidae has distinctive long hindwing tails. eyespots on the wings, to
Found in New Guinea and Australia, this The caterpillars feed on a range startle predators.
moth is one of the largest in the world. hindwing tail of deciduous trees.
Only the male has the long hindwing tails.

21/4–31/2 in 13⁄4–3 in 2–31⁄4 in 2–31/4 in

6–9 cm 4.5–7.5 cm 5–8.5 cm 5–8 cm

GIANT LEOPARD MOTH OAK EGGAR LAPPET MOTH PINE-TREE LAPPET

Hypercompe scribonia Lasiocampa quercus Gastropacha quercifolia Dendrolimus pini
f: Erebidae f: Lasiocampidae f: Lasiocampidae f: Lasiocampidae
The distribution of this striking moth Ranging from Europe to N. Africa, Found in Europe and Asia, the specific This moth is widespread in
extends from S.E. Canada as far south as the caterpillars of this moth feed name of this large moth refers to its coniferous forests across Europe
Mexico. The caterpillars feed on the foliage of bramble, oak, resting appearance, which resembles and Asia, where the caterpillars
on a variety of plants. heather, and other plants. a cluster of dead oak leaves. feed on pine, spruce, and fir.

white chevrons on forewing CASE-BEARING

3/8–1/2 in CLOTHES MOTH
1–1.6 cm Tinea pellionella
f: Tineidae
Found in W. Europe and parts
11/2–13/4 in of N. America, this species
3.5–4.5 cm can cause serious damage to
wool clothing and carpets.
AUSTRALIAN MAGPIE MOTH
Nyctemera amicus
f: Erebidae feathery
antennae
Widespread in Australia and also found
in New Zealand, the caterpillars of this
day-flying moth feed on plants such
as groundsel and ragwort.
13/4–23/4 in
4.5–7 cm GARDEN TIGER
Arctia caja
f: Erebidae
This unmistakable moth is
found across the northern
hemisphere. The hairy
caterpillars feed on
a range of low-growing
plants and shrubs.
23/4–31/4 in 1–11/4 in
7–8 cm 2.5–3 cm

ILIA UNDERWING VAPOURER MOTH

Catocala ilia Orgyia antiqua
f: Erebidae f: Erebidae
This widespread species from N. Now found across the northern abdomen
America has distinctive, red-banded hemisphere, the female of this
hindwings. The caterpillars feed European species has tiny
on the foliage of oak trees. wings and is unable to fly.
 91/2–12 in
24–30 cm

GIANT AGRIPPA
Thysania agrippina
f: Noctuidae
Found in C. and parts of S. America,
this species has one of the largest
wingspans of any moth in the world.

SNOUT MOTH
Vitessa suradeva
f: Pyralidae
This moth occurs in India, parts
of S.E. Asia, and New Guinea. The
caterpillars feed in a web on young
leaves of poisonous shrubs.
11/2–2 in
4–5 cm
1–11/4 in
2.4–2.8 cm

21/4–4 in 21/2–33/4 in
6–10 cm 6.5–9.5 cm
DIVA MOTH
Divana diva
f: Castniidae
ACACIA CARPENTER Found in tropical forests in
MOTH S. America, this day-flying
SMALL MAGPIE SILVER-SPOTTED Endoxyla encalypti species is well camouflaged
Anania hortulata GHOST MOTH f: Cossidae at rest despite its brightly
f: Pyralidae Sthenopis argenteomaculatus The robust, white caterpillars colored hindwings.
Found in hedgerows and waste f: Hepialidae of this large and distinctive
ground, the caterpillars of this very This species occurs in S. Canada and Australian moth burrow into
common European species feed parts of the US. The caterpillars mainly the woody tissue of certain 31/2–43/4 in
on rolled nettle leaves. feed inside the roots of alder trees. species of the acacia tree. 9–12 cm

large eyespot to
scare predators 2–21/2 in 11/4–11/2 in 11/2–2 in
5–6.5 cm 3–4 cm 4–5 cm

LARGE EMERALD ARGENT AND SABLE CLARA’S SATIN MOTH

Geometra papilionaria Rheumaptera hastata Thalaina clara
f: Geometridae f: Geometridae f: Geometridae
This moth occurs across Europe The common name for this day-flying moth The caterpillars of this moth,
and the temperate parts of Asia. from the northern hemisphere comes from the which occurs in E. and
The caterpillars feed mainly on heraldic terms for silvery-white and black. S.E. Australia and N. Tasmania,
the foliage of birch trees. feed on acacia foliage.

23/4–3 in 11/2–21/4 in
7–7.5 cm SILK MOTH 4–6 cm
Bombyx mori
f: Bombycidae
Originating in China, the
wavy pattern silk moth has been bred in
of black and captivity, on a diet of white
orange lines mulberry leaves, for
thousands of years. Its
cocoon is used to make silk.
4–61/2 in
10–16 cm
1–11/4 in
2.5–3 cm
WHITE
PLUME-MOTH
Pterophorus pentadactyla
WALLICH’S OWL MOTH COPPERY f: Pterophoridae
Brahmaea wallichii DYSPHANIA Common across Europe
f: Brahmaeidae Dysphania cuprina on dry grassland, waste
This large moth occurs from f: Geometridae ground, and gardens,
N. India to China and Japan. The Widespread across S.E. Asia, this brightly the caterpillars of this
caterpillars eat the foliage of colored moth flies during the day and is thought distinctive species feed long legs
ash, privet, and lilac. to be unpalatable to birds. on hedge bindweed.
 MOTHS AND BUTTERFLIES
298
INVERTEBRATES • INSECTS

2–21/4 in 13/4–21/2 in 11/2–2 in 21/4–21/2 in

5–6 cm 4.5–6.2 cm 3.5–5 cm 5.5–6.5 cm

REGENT SKIPPER GUAVA SKIPPER HORNET MOTH BUFF-TIP

Euschemon rafflesia Phocides polybius Sesia apiformis Phalera bucephala
f: Hesperiidae f: Hesperiidae f: Sesiidae f: Notodontidae
This brightly colored species is This species can be found from To deter predators, the adults of this This moth is found in Europe
native to tropical and subtropical S. Texas to as far south as Argentina. moth look very similar to hornets but are and eastward to Siberia. At rest
forests in E. Australia, where The caterpillars feed inside harmless. The caterpillars bore into the its folded wings wrap around its
it can be seen feeding at flowers. rolled-up guava leaves. trunk and roots of poplars and willows in body for camouflage, closely
Europe and the Middle East. resembling a broken-off twig.

COMMON
MORPHO 33/4–6 in
Morpho peleides 9.5–15 cm
f: Nymphalidae
This butterfly is widespread
21/4–23/4 in 31/2–43/4 in in tropical forests in
6–7 cm 9–12 cm
C. and S. America.
The adults feed on
ELEPHANT HAWK MOTH VERDANT SPHINX the juices of
Deilephila elpenor Euchloron megaera rotting fruits.
f: Sphingidae f: Sphingidae
This pretty, pinkish hawk moth is This distinctive moth is widespread
widespread in the temperate parts throughout sub-Saharan Africa. The
of Europe and Asia. The caterpillars caterpillars feed on the foliage of metallic blue color is
feed on bedstraw and willowherb. creepers of the grape family. attractive to mates

31/4–4 in
8.5–10 cm

11/4–11/2 in 13/4–2 in 2–21/4 in

3–4 cm 4.5–5 cm 5–6 cm

SIX-SPOT BURNET LITTLE WOOD SATYR MONARCH BUTTERFLY

Zygaena filipendulae Euptychia cymela Danaus plexippus
f: Zygaenidae f: Nymphalidae f: Nymphalidae
Brightly colored and distasteful This woodland butterfly is found A well-known migrant, this butterfly has
to birds, this day-flying moth can from S. Canada to N. Mexico. The spread from the Americas to many other
be found in meadows and woodland caterpillars feed on grasses in parts of the world. The caterpillars feed
clearings across Europe. clearings close to water. on the milkweed plant.
QUEEN CRACKER WHITE ADMIRAL
Hamadryas arethusa Ladoga camilla
f: Nymphalidae f: Nymphalidae
long The common name for this This butterfly can be found
antennae butterfly, found in forests across temperate Europe and hindwing tail
from Mexico to Bolivia, Asia to Japan. The caterpillars
21/4–23/4 in refers to the clicking noise feed on the foliage of the
6–7 cm it makes when it flies. honeysuckle.

23/4–31/2 in 31/2–43/4 in
7–9 cm 9–12 cm

PURPLE EMPEROR INDIAN LEAF BUTTERFLY

Apatura iris Kallima inachus
f: Nymphalidae f: Nymphalidae
Found in dense oak woodland throughout With undersides that resemble a
Europe and Asia as far east as Japan, the brown leaf, this butterfly is perfectly
male of this butterfly is iridescent purple camouflaged at rest with its wings
whereas the female is dull brown. folded. It occurs from India to S. China.
 SMALL WHITE pointed front 11/2–21/2 in
Pieris rapae wing
f: Pieridae
4–6.5 cm
299
This butterfly can now
be found worldwide. The
caterpillars feed on wild and
cultivated cabbages and
mustards, and can be pests.

11/2–2 in
3.5–5 cm

CALIFORNIA DOG-FACE
Zerene eurydice
f: Pieridae
3–33/4 in This butterfly is restricted to parts
7.5–9.5 cm of California and, sometimes,
western Arizona. It breeds on
a shrub called Napa False Indigo.

11/2–2 in
4–5 cm
ORANGE TIP
Anthocharis cardamines
f: Pieridae
This butterfly can be found
in meadows across temperate
Europe and Asia as far as Japan.
The caterpillars feed on lady’s
smock and garlic mustard.
23/4–31/4 in
7–8.5 cm
ORANGE-BARRED
SULPHUR
Phoebis philea
f: Pieridae
The distribution of this
butterfly extends from S. Brazil
to C. America, S. US, and
sporadically farther north. The
caterpillars feed on sennas.

2–23/4 in
5–7 cm
MADAGASCAN SUNSET MOTH
Chrysiridia rhipheus
f: Uraniidae
This colorful, day-flying moth,
with iridescent wing scales, is endemic
to Madagascar. It feeds on certain
poisonous shrubs of the spurge family.
TIGER
iridescent red PIERID
wing marking dark spots on Dismorphia amphione
hindwing f: Pieridae
This colorful butterfly, which
21/4–31/4 in mimics unpalatable butterfly species
6–8 cm for protection from predators, is
widespread and common from
Mexico southward to S. America.
4–6 in
10–15 cm
21/4–3 in 2–23/4 in
underside 5.5–7.5 cm 5–7 cm

SMALL POSTMAN
Heliconius erato
f: Nymphalidae
This butterfly is common along
forest edges and open ground
from C. America to S. Brazil. The
caterpillars eat passionflower foliage.
BLACK-VEINED CLEOPATRA
WHITE Gonepteryx cleopatra
OWL BUTTERFLY Aporia crataegi f: Pieridae
Caligo idomeneus f: Pieridae This butterfly is common in
f: Nymphalidae Occurring across Europe, N. Africa, countries around the Mediterranean,
The underside of this large species, native to and Asia as far as Japan, this particularly in lightly wooded coastal
S. America, has prominent owllike eyespots distinctive butterfly breeds on areas and scrubland. The caterpillars
to deter predators when the butterfly is at rest. hawthorn and blackthorn. feed on species of buckthorn.
 MOTHS AND BUTTERFLIES
300 10–12 in
25–31 cm
24∕5–31/4 in 21∕4–4 in
7–8.5 cm 6–10 cm
INVERTEBRATES • INSECTS

QUEEN
ALEXANDRA’S
BIRDWING
Ornithoptera alexandrae
f: Papilionidae
Found only in
S.E. Papua New Guinea
east of the Owen
Stanley Ranges, this
BIG GREASY BUTTERFLY ZEBRA endangered species,
Cressida cressida SWALLOWTAIL the largest butterfly
f: Papilionidae Photographium marcellus in the world,
This species is found in grassland f: Papilionidae is now protected.
and drier forests in Australia and Found in damp woodlands in eastern N. America,
Papua New Guinea, where its this butterfly has distinctive black and white
pipevine foodplants grow. markings. The caterpillars feed on pawpaw.

6–7 in
15–18 cm

4¾–7½ in 3–31∕2 in 11∕2–21∕4 in

12–19 cm 7.5–9 cm 4–5.5 cm

COMMON GREEN BIRDWING

RAJAH BROOKE’S BIRDWING Ornithoptera priamus SWALLOWTAIL
Troides brookiana f: Papilionidae Papilio machaon
f: Papilionidae This large butterfly occurs from Papua New Guinea f: Papilionidae GREEN
This butterfly lives in the tropical forests of Borneo and the Solomon Islands to tropical N. Australia. This species is found in wet meadow, DRAGONTAIL
and Malaysia, where the adults sip fruit juices and The caterpillars eat pipevines. fenland, and other habitats across Lamproptera meges
nectar. The caterpillars feed on pipevines. the northern hemisphere. f: Papilionidae
The caterpillars feed on This unmistakable butterfly,
various umbellifers. which hovers while it feeds at
flowers, can be found from
red spot India to China and through
S. and S.E. Asia.

short, robust
antenna

13/4–2 in
4.5–5 cm

21∕4–3½ in
6–9 cm
APOLLO
Parnassius apollo
f: Papilionidae
This butterfly can be found in flower-rich meadows
in mountainous regions of Europe and Asia.
The caterpillars feed on stonecrop.

zig-zag pattern
on hindwing
23∕4–31∕4 in
7–8 cm

31∕4–3½ in
BLUE 8–9 cm
TRIANGLE
Graphium sarpedon
f: Papilionidae SCARCE SPANISH FESTOON
Common and widespread SWALLOWTAIL Zerynthia rumina
from India to China, Iphiclides podalirius f: Papilionidae
Papua New Guinea, and f: Papilionidae This species lives in scrub, meadows, and
Australia, this butterfly Despite its name, this species is quite widespread rocky hillsides in S.E. France, Spain,
feeds on nectar and in Europe and across temperate Asia to China. Portugal, and parts of N. Africa. The
drinks from puddles. The caterpillars feed on blackthorn. caterpillars feed on species of birthwort.
 31/4–5½ in SMALL COPPER 1–11∕4 in
8–14 cm Lycaena phlaeas 2.5–3 cm
f: Lycaenidae
Common in Europe,
N. Africa, and Asia as far as
Japan, this species can also
be found in N. America.
The caterpillars feed on
sorrels and docks.
11∕4–11∕2 in
3–4 cm

TIGER BROWN HAIRSTREAK BLUE THAROPS

SWALLOWTAIL Thecla betulae Menander menander
Papilio glaucus f: Lycaenidae f: Lycaenidae
f: Papilionidae This species can be found in This fast-flying butterfly occurs
This species is widespread in hedgerows, scrubland, and in tropical forests from Panama
N. America. The young woodland across Europe and to northern S. America. Little
caterpillars, which resemble temperate Asia. The caterpillars is known about its life cycle
bird droppings, feed on a 11∕2–13∕4 in feed at night on blackthorn. or caterpillars.
variety of trees and shrubs. 3.5–4.5 cm

1–11∕2 in 3∕4–1 in 11∕4–11∕2 in

2.5–3.5 cm 2–2.5 cm 3–4 cm

ADONIS BLUE SONORAN BLUE

Lysandra bellargus Philotes sonorensis DUKE OF BURGUNDY FRITILLARY
f: Lycaenidae f: Lycaenidae Hamearis lucina
This European species can be This rare butterfly is restricted to f: Lycaenidae
found in chalky grassland, rocky washes and desert cliffs in This species ranges across C. Europe as far
where the caterpillars feed on California. The caterpillars feed as the Urals and favors flower meadows,
horseshoe vetch. The male is sky on succulents known as stonecrop where its food plants, cowslip
blue while the female is brown. or “live-forever” plants. and primrose, grow.

S AW F L I E S , WA S P S , STEM SAWFLY
Cephus nigrinus
7∕32–11∕32 in
7–9 mm
BEES, AND ANTS f: Cephidae
This slender, all-black sawfly is
Hymenoptera typically have two pairs of wings, joined in widespread in W. Europe. The larvae
flight by tiny hooks. Apart from sawflies, they have a “wasp- bore downward inside the stems of
waist” and females have an ovipositor that may be modified wild and some cultivated grasses.
to sting. Many wasps are predatory or parasitic. Bees are
vital pollinators and ants play a role in many ecosystems. ¾–9∕10 in
1.8–2.2 cm

7∕32–11∕32 in
7–9 mm CIMBICID SAWFLY
Trichiosoma lucorum
f: Cimbicidae
This stout-bodied
European sawfly is found
fine tufts on in woodland, hedgerows,
wing margin and scrubby areas. slender
¾–11∕2 in antennae
The larvae feed on 2–4 cm
TENTHREDO ARCUATA birch and willow.
f: Tenthredinidae
Widespread in meadows, this distinctive
black- and yellow-marked European sawfly lays large
its eggs on clovers. head
PERGID SAWFLY
Unknown sp.
f: Pergidae
These herbivorous insects are found in ¾ in
Australia and S. America. Many species 2 cm
attack eucalyptus trees, and young
larvae feed gregariously. 7∕32–11∕32 in
7–9 mm
HORNTAIL
Urocerus gigas
f: Siricidae
This impressive species
ROSE SAWFLY can be found across the
Arge ochropus northern hemisphere.
f: Argidae RED-HEADED SAWFLY The female drills
Recognizable by the Acantholyda erythrocephala deep into softwood
black along the wing f: Pamphiliidae trees to lay
edge, this European Originally found in Europe and Asia, her eggs.
sawfly visits flat-topped this sawfly has spread to North
7∕32–3∕8 in flowers to feed. The America. The larvae feed communally
7–10 mm larvae feed on wild roses. under silken webs among pine needles. ovipositor
 S AW F L I E S , WA S P S , 1∕8–3∕8 in
BRACONID WASP
302 BEES, AND ANTS 3–10 mm
Unknown sp.
f: Braconidae
1∕32–1∕8 in Found worldwide,
1–3 mm braconid wasps large
compound eye
INVERTEBRATES • INSECTS

generally live as
parasites on caterpillars
and the larvae of beetles
and flies. The females
of some species have
a long ovipositor.

FIG WASP
Unknown sp.
f: Agaonidae
Fig wasps are found in tropical and subtropical bright metallic
regions. Many are essential pollinators, breeding coloration
inside specific species of fig.

3∕16–3∕10 in
5–6.5 mm

OAK APPLE
GALL WASP
Biorhiza pallida
f: Cynipidae
This wasp emerges from root
3∕4–9∕10 in
galls on oak trees in Europe 2–2.2 cm
and Asia. It lays eggs
in buds, inducing the growth
of oak apple galls.

GIANT 3∕4–11∕2 in 7∕10–3∕4 in

WOOD WASP 2–4 cm 1.8–2 cm
Rhyssa persuasoria
f: Ichneumonidae
Found in pine forests
across the northern LISSONOTA SP. SPLENDID
hemisphere, these very f: Ichneumonidae EMERALD WASP
large wasps drill into The genus Lissonota contains Stilbum splendidum
trees to lay their egg on many very similar wasps. This f: Chrysididae
the larvae of horntails. species drills into a trunk to This large wasp from N. Australia
lay its eggs in a wood-boring lives as a parasite on the larvae
moth larva. of solitary mud-nesting wasps.
1∕8–5∕32 in They are known to visit flowers to feed.
3–4 mm
1∕32–1⁄8 in
1–3 mm

3∕8–½ in
TORYMUS SP. 1–1.6 cm
f: Torymidae hard, dimpled
This wasp uses her long ovipositor to drill body surface gives
through gall tissue and lay eggs on the PTEROMALID CHALCID EUROPEAN ½–3∕5 in protection from stings
WASP WASP BEEWOLF 1.3–1.5 cm
developing larvae inside.
Mesopolobus typographi Chalcis sispes Philanthus triangulum
f: Pteromalidae f: Chalcididae f: Crabronidae
This European and Asian This wasp is found in Found in S. and C.
wasp is a hyperparasite; it is Europe and parts of Asia, Europe and N. Africa,
a parasite on another wasp where it lives as a parasite this wasp makes a nest in
21∕2–31∕4 in that is itself a parasite on on the aquatic larvae of sandy areas stocked with
6–8 cm the larvae of bark beetles. large soldier flies. honey bees as food for
its larvae.

smoky
orange wing 13∕4–21∕4 in 11∕32–3∕5 in
4.5–5.5 cm 9–14 mm

TARANTULA HAWK MAMMOTH WASP

Pepsis heros Megascolia procer VELVET ANT
f: Pompilidae f: Scoliidae Mutilla europaea
This large S. American Native to Borneo, Java, f: Mutillidae
wasp hunts for tarantulas, and Sumatra, this wasp This European species lives in sandy
which it paralyzes paralyzes the larvae of areas and rough grassland. The
and buries as food scarab beetles and lays its female is wingless, and the larvae
for its larvae. eggs on them. eat the larvae of bumble bees.
 5∕16– 1∕2 in HORNET ½–11∕4 in 1⁄2–3∕4 in
7.5–12 mm Vespa crabro 1.5–3 cm 1.4–2.2 cm
f: Vespidae 303
This large social wasp
lives in Europe and
Asia and has been
introduced elsewhere.
TIPHIID WASP It favors woodland, BUFF-TAILED
Methocha articulata where it makes its nest BUMBLE BEE
f: Tiphiidae in hollow trees. Bombus terrestris
The wingless females of this European species f: Apidae
live in sandy areas, where they are a parasite on This social bee, which is native to
the larvae of tiger beetles inside their burrows. wasp-waist C. and S. Europe and N. Africa,
has been introduced elsewhere. It
½–3∕4 in is an important crop pollinator.
COMMON 1.2–1.9 cm
WASP
Vespula vulgaris
f: Vespidae
Native to the northern
hemisphere, this wasp
feeds its young on
insects and makes a
paper nest from chewed very hairy
wood fibers. body

½ in
TAWNY 1.2–1.6 cm
MINING BEE
Andrena fulva
f: Andrenidae
Found in C. Europe, this
early spring bee makes a
nest underground in grassy ½ in
HONEY 1.2–1.4 cm
areas, raising a little mound
of soil at the entrance. BEE ORCHID
Apis mellifera BEE
2∕5–½ in 11∕32–3⁄5 in f: Apidae Euglossa asarophora
1.1–1.3 cm 9–15 mm The western honey bee, an f: Apidae
important crop pollinator, is Like all orchid bees, this species lives
now found worldwide. In the in colonies in the rainforests of
wild it nests in hollow trees, but S. America. Males collect oils and
is widely kept in domestic hives. resins from orchids to attract a mate.
GREAT
CARPENTER BEE 3∕16–11∕32 in
Xylocopa latipes 5–9 mm
f: Anthophoridae
Found all over S.E. Asia,
PLASTERER BEE this large bee excavates
Colletes sp. nest cavities in tree
3∕5 in
f: Colletidae branches or wooden 1.6 cm
These solitary, ground-nesting timbers and posts.
bees are common across the 11∕4–11∕2 in
3.3–3.6 cm WOOD ANT
northern hemisphere. They line Formica rufa
their nests with a waterproof f: Formicidae
abdominal secretion. Common throughout Europe,
this ant is an important predator
compound of insects in forests. It squirts LEAF-CUTTER
eye formic acid from the rear of its ANT
abdomen as a defense. Atta sp.
f: Formicidae
Leaf-cutter ants live in
C. and S. America.
They can make vast
underground nests,
where they cultivate a
special fungus on chewed
leaf fragments as food.

soldier

1∕8–½ in
3–12 mm
band of
pale hair ½–3∕4 in
3∕8–1∕2 in 1.3–2 cm
1.1–1.7 cm

SWEAT BEE ARMY ANT

WOOL CARDER BEE Halictus quadricinctus Eciton burchellii
Anthidium manicatum f: Halictidae f: Formicidae
f: Megachilidae This solitary bee is found across This S. American ant
This European bee makes nests in existing S. Europe and the Mediterranean. forms large colonies of up
holes in wood or masonry, which it lines with It makes large clusters of cells to 2 million individuals
hairs that it collects from certain plants. underground in which to rear its young. and attacks large prey.
304 R I B B ON WO R M S B RYO Z O A N S
Marine worms of the phylum Nemertea are voracious These minute animals form corallike colonies,
predators, seizing their prey with a proboscis and filter-feeding with microscopic tentacles and,
INVERTEBRATES • RIBBON WORMS

swallowing it whole or sucking its fluids. in turn, providing food for grazing invertebrates.
Ribbon worms are soft slimy PHYLUM NEMERTEA
Many bryozoans resemble PHYLUM BRYOZOA
animals, cylindrical in shape CLASSES 4 coral, but they are more CLASSES 3
or somewhat flattened. Many ORDERS 4 advanced animals than ORDERS 7
simply wriggle across the FAMILIES 48 cnidarians. A colony— FAMILIES About 160

seabed, but some can also SPECIES 1,350 genetically a single individual— SPECIES 6,409

swim. One species, Lineus is made up of thousands of

longissimus, the bootlace worm, has been recorded growing to 100 ft tiny bodies called zooids, each consisting of a retractable fan of
(30 m) in length. Like others, it has a fragile body that is easily broken. tentacles surrounding the mouth. Microscopic, beating hairs on the
The proboscis of a ribbon worm is not part of its gut, but emerges tentacles waft particles of food down to the mouth, which leads into
from a sac on its head just above the mouth. In some species the a U-shaped gut. Waste exits by the anus in the body wall. Although a
proboscis has sharp spikes for gripping or piercing prey, and even magnifying glass is needed to see individual zooids, colonies assume a
venom for immobilizing it. These well-armed species prey on other variety of forms, depending upon species. Some are encrustations
invertebrates, such as crustaceans, polychaete worms, and mollusks. on rock or seaweed, others grow into bristly thickets or fleshy lobes.
Other ribbon worms feed on dead material. Many are heavily calcified like stony corals; others are soft.

FALSE CORAL
3–4 in BRYOZOAN
8–10 cm Myriapora truncata
f: Myriaporidae
10–12 in A distinctive bryozoan of the
PACIFIC STRIPED 25–30 cm RED RIBBON WORM Mediterranean, the colonies of
RIBBON WORM Nipponnemertes pulchra this species consist of thick
Baseodiscus hemprichii f: Cratenemertidae cylinders that branch repeatedly.
f: Valenciniidae Found in cold, ocean waters
This colorful ribbon worm around the world, this ribbon
lives on reefs of the Indo-Pacific. 1⅛–11/4 in
worm varies in color from 3–4 cm
It hunts for invertebrate prey on head orange to pink or red.
the bottom of the sea.
smooth
slimy body
COMMON
ATLANTIC RIBBON
WORM
Tubulanus annulatus
f: Tubulanidae
A large ribbon worm, this small ring of
species lives in muddy tentacles around
sediment in intertidal or central mouth
offshore waters of the
4–8 in
N.E. Atlantic and 10–20 cm
Mediterranean.
HORNWRACK
Flustra foliacea
4¾–30 in
12–75 cm
f: Flustridae
Typical of its family, this bryozoan forms erect,
leafy colonies. This stiff, bushy offshore species
is common on rocky N. European coasts.

2–8 in
5–20 cm
6–8 in
15–20 cm

PINK LACE BRYOZOAN LACY CRUST BRYOZOAN

Iodictyum phoeniceum Membranipora membranacea
f: Phidoloporidae f: Membraniporidae
Lace bryozoans—sometimes inaccurately The bryozoans known as sea mats grow
called “lace corals”—form tall, rigid in encrusting lacy sheets. This species
colonies. This colorful species lives on forms fast-growing colonies on kelp
coastlines around S. and E. Australia. fronds of the N.E. Atlantic.
LA MPSHELLS MOLLUSKS 305
Lampshells look very like bivalve mollusks, Mollusks are a large, highly diverse group, ranging from blind
but feed instead with tentacles. They filter-feeding bivalves cemented to rock, through voraciously

INVERTEBRATES • MOLLUSKS
belong to a different, very ancient phylum. grazing snails and slugs, to lively, intelligent octopus and squid. 
The soft body PHYLUM BRACHIOPODA
A typical mollusk has a soft body carried PHYLUM MOLLUSKA
of a lampshell is CLASSES 3 on a large muscular foot, and a head that CLASSES 9
enclosed within a ORDERS 5 senses the world using eyes and tentacles. ORDERS 53
two-valved shell FAMILIES About 30 The viscera (internal organs) are contained FAMILIES 609

that has one valve SPECIES 414 in a visceral hump, which is covered by a SPECIES 71,719

larger than the fleshy mantle. This mantle overhangs the

other. The shell is either attached to the seabed with edge of the hump, creating a groove called
a rubbery stalk or is directly cemented to rock. The the mantle cavity, which is used for
shell valves are positioned top-and-bottom around the breathing. In most species, this mantle
animal—not around the sides, as in bivalve mollusks. also secretes substances used to make the
Like mollusks, lampshells have a fleshy mantle attached shell. Most mollusks feed with a tongue
to the inner surface of the shell, which encloses a called a radula. Coated in teeth made of
mantle cavity. In the mantle cavity there is a ring of chitin, the radula moves forward and
tentacles with microscopic beating hairs that drive backward through the mouth to rasp at
particles to a central mouth—just like the feeding food. Bivalves lack a radula. They feed
structure in bryozoans. instead by siphoning water through their
The fossil record shows that lampshells were much shell. Most species trap particles of food
more common and demonstrated much greater in the mucus on their gills.
diversity in the warm, shallow seas of the Paleozoic Era.
They began to decline dramatically during the time of w ith an d w ithout sh ells
the dinosaurs—perhaps because bivalve mollusks were A shell is not only a refuge from predators:
more successful. it is also protection from drying out. Some
snails even have a trapdoor called an
These limpets have grazed the rock around
operculum to seal the opening of the them bare, but they cannot reach the green
EUROPEAN LAMPSHELL
shell. The shell is hardened with minerals algae growing on top of their own shell.
¾–1¼ in
2–3 cm Terebratulina retusa obtained from the diet and surrounding
f: Cancellothyrididae water. It is coated in tough protein; and smooth on the inside, to allow the body to
Found from the N.E. Atlantic to
the Mediterranean, this lampshell slide in and out—some groups have a lining of mother-of-pearl. Many shell-less
has pear-shaped valves and attaches mollusks defend themselves with chemicals that make them distasteful, or even
to vertical rocks using a short stalk.
poisonous, and advertise this with striking colors.
Most cephalopods—tentacled mollusks, such as octopus and squid—lack an
obvious shell. Members of this group are entirely predatory and have a horny beak
1¼–2¼ in
3–5.5 cm for chewing on flesh. Their muscular foot is modified into their characteristic
tentacles, which are used for grasping and swimming.

g e t t i n g ox yg e n
Most mollusks are aquatic and so they have gills for breathing, usually projecting
into the mantle cavity and irrigated by a current of water. In most land snails and
slugs the mantle cavity is filled with air and functions as a lung. Because many
freshwater snails probably evolved from this group, they have a lung, too, and must
frequently surface to breathe.

PACIFIC LAMPSHELL
hinged, two- Terebratalia transversa
valved shell f: Terebrataliidae A PL ACOPHOR A NS
An abundant lampshell of the northern
Pacific, this short-stalked species has a More wormlike than mollusklike, the
⅜–1/2 in
variable smooth or ribbed shell. Aplacophora are small cylindrical burrowers ⅛ –3¼ in
3 mm–8 cm COMMON
1–1.5 cm that feed on detritus or other invertebrates GLISTENWORM
INARTICULATED in the sediment of deep oceans. They lack a Chaetoderma sp.
LAMP SHELL
shell, but their clearly molluskan features f: Chaetodermatidae
Novocrania anomala This wormlike mollusk
f: Craniidae include a rasping, tonguelike radula. The is covered by a tough
A brachiopod of North Atlantic mantle cavity is reduced to an opening cuticle. It burrows
waters, this species cements its through muddy sediment
shell to rock, and superficially at the rear of the body, into which the in the depths of the
resembles a limpet. animal discharges waste from the gut. North Atlantic.
306 B I VA L V E S
Highly specialized aquatic mollusks, called a byssus. Others use a well-developed PHYLUM MOLLUSKA
bivalves are instantly recognizable, muscular foot for burrowing into muddy sediment.
I N V E RT E BR AT E S • BI VA LV E S

CLASS BIVALVIA

with a hinged shell that opens up to A few bivalves, such as scallops, swim freely, using ORDERS 19

access food and oxygen-rich water. a method of jet propulsion. FAMILIES 105

Water is pumped in and out of the shell through SPECIES 9,733

Bivalves are identifed from their shell, which tubes, called siphons. The water supplies oxygen
consist of two plates, called valves, joined together and food, which are collected as the water passes
by a hinge. To protect against predators and the over modified gills. Food particles caught in the
dangers of drying out, the shell clamps tightly shut sticky mucus around the gills are wafted toward
by contracting powerful muscles, sealing most or the mouth by microscopic hairs.
all of the body inside. Seashore species are regularly
exposed by receding tides. b i va lv e s a n d p e o p l e
Within the class there is a huge variation in size, Mussels, clams, and oysters are important sources
ranging from the tiny fingernail clam at about of food, and some oysters create fine-quality pearls
¼ in (6 mm) to the giant clam, which can grow to by coating a foreign object with layers of nacre.
A queen scallop claps the two valves of
4½ ft (1.4 m) across. Some bivalves attach to rocks Bivalves are also useful indicators of water quality, its shell together to accelerate away from the
and hard surfaces by a bundle of tough threads, as many cannot survive high levels of pollution. unwelcome attention of a predatory starfish.

OYST E R S A N D 4–4¾ in
COCK’S COMB
OYSTER
SCALLOPS 10–12 cm Lopha cristagalli
f: Ostreidae
The Ostreoida feed on tiny food particles Close relatives of scallops,
filtered from the seawater. Many oyster many species of oyster
are prized as food and for
species live permanently submerged in their pearls. This is an
coastal waters, fixed to the rock by the Indo-Pacific species.
gland that produces byssus thread in
other bivalves. Scallops, by contrast,
can swim freely by clapping their
valves open and shut. spiny left
valve
4¾–6 in
12–15 cm
3¼–4 in
8–10 cm
GREAT
SCALLOP EDIBLE OYSTER
Pecten maximus CAT’S TONGUE OYSTER Ostrea edulis
f: Pectinidae Spondylus linguafelis f: Ostreidae
As well as swimming freely, f: Spondylidae Formerly abundant in Europe, this
scallops have jet-propelled One of a family of thorny oysters commercially important species is
escape responses. This with a colorful mantle, the cat’s now overfished in some areas. The
commercial species inhabits 4–4¾ in tongue oyster of the Pacific is species is inedible during summer
fine sands of European coasts. 10–12 cm named for its spiny exterior. months when breeding.

A RC C L A M S MARINE FA N M U SSE L S
A N D R E L AT I V E S MUSSELS A N D R E L AT I V E S
The shell of these bivalves is closed by The Mytiloida have a distinctively shaped, Members of the Pterioida include long-
two strong muscles, and hinge along elongate, asymmetrical shell that hinged, winged oysters and T-shaped
a straight line bearing a continuous attaches to rocks by byssus threads. hammer-oysters as well as fan mussels.
series of teeth. Like their relatives, Only one of their two shell-closing The group also contains commercially
the Arcoida have a reduced foot and muscles is well developed. important marine pearl oysters.
large gills that help trap food particles.
3¼–4 in 10–16 in
2–2¾ in 2–2¼ in 8–10 cm 25–40 cm
5–7 cm 5–6 cm

NOAH’S ARK EUROPEAN BITTERSWEET

Arca noae Glycymeris glycymeris
f: Arcidae f: Glycymerididae COMMON MUSSEL FLAG FAN MUSSEL
Arc shells are square-edged, Dog shells are rounded relatives Mytilus edulis Atrina vexillum
thick-shelled bivalves. Noah’s of arc shells. This N.E. Atlantic f: Mytilidae f: Pinnidae
Ark is a byssus-attached species, fished in Europe, has The most commercially important mussel in Europe, Fan mussels, such as this species from western
intertidal species on rocky a sweet flavor but is tough this long-lived species occurs in dense beds. It can European coastlines, have a triangular shell
coasts of the E. Atlantic. when overcooked. tolerate the low salinity of estuarine waters. attached by a byssus in soft sediment.
F R E S H WAT E R 4–6 in
COCKLES AND
MUSSELS 10–15 cm R E L AT I V E S ¾–1½ in
307
2–4 cm
The Unionoida are the only exclusively The largest order of bivalves, the Veneroida include a CRADLE DONAX
freshwater order of bivalves. The tiny large range of marine animals. Most have short siphons Donax cuneatus

I N V E RT E BR AT E S • BI VA LV E S
larvae use their valves to clamp onto that are often fused together. Some, notably the cockles, f: Donacidae
fishes, forming cysts on the gill fins are agile, capable of burrowing or even leaping with One of a family of triangular
wedge shells that are fast
and feeding on blood or mucus before their foot. Others are attached to rocks by a byssus. burrowers in surf, this is a
dropping off as juvenile mussels. tropical Indo-Pacific species.
11/4–1½ in ZEBRA MUSSEL
3–4 cm
3½–4 in Dreissena polymorpha 2–21/4 in
9–10 cm f: Dreissenidae 5–6 cm
This freshwater mussel is
anchored by byssus threads, but
can detach and crawl on a slender 6–8 in
FRESHWATER foot. Native to E. European 15–20 cm
PEARL MUSSEL waters, it has spread elsewhere.
SWAN MUSSEL Pinctada margaritifera SWORD RAZOR
Anodonta sp. f: Margaritiferidae CLAM
f: Unionidae Well-known for its good- COMMON EDIBLE 1½–2 in Ensis siliqua OBLONG
Bitterling fishes lay eggs in living quality pearls, this mussel lives
COCKLE 4–5 cm f: Pharidae TRAPEZIUM
Cerastoderma edule Razor shells, such as this Trapezium oblongum
freshwater mussels, such as this buried in sand or gravel f: Cardiidae f: Trapezidae
Eurasian species. The mollusk beneath fast-flowing rivers N.E. Atlantic species, live
becomes a nursery for fish fry. in Eurasia and N. America. Sand-burrowing cockles in burrows, feeding and An Indo-Pacific bivalve,
have a shell with radiating respiring through their this animal lives attached
ribs. This N.E. Atlantic emergent siphon, but drop to rocks by byssus threads,
species, often found in quickly using their muscular usually in crevices or
T RU E C L A M S A N D vast numbers, is fished
in N. Europe.
foot when disturbed. beneath coral rubble.
B O R I N G B I VA LV E S 31/4–3½ in
8–9 cm
Long-siphoned bivalves in the order Myoida typically 2–2¾ in
1–11/4 in
burrow in mud or bore through wood or rock. In 2.5–3 cm
5–7 cm
piddocks the front of the shell acts like a file to bore
burrows in soft rock. Shipworms use their shell valves
to drill through wood. 4¾–6 in
12–15 cm SUNRISE TELLIN WEST AFRICAN TELLIN
4¾–6 in Tellina radiata Peronaea madagascariensis
12–15 cm f: Tellinidae f: Tellinidae
The shell of this tellin from Tellins, such as this
the Caribbean has a variable rose-colored tropical species,
banding pattern and is often feed on particles in sediment,
found on beaches. drawn in through their long,
extendable siphon.
concentric rings
COMMON PIDDOCK SOFT SHELL CLAM
Pholas dactylus Mya arenaria ROYAL COMB VENUS STRIPED TELLIN
f: Pholadidae f: Myidae Hysteroconcha dione Tellinella virgata
This common piddock of the An edible clam with a thin f: Veneridae f: Tellinidae
N.E. Atlantic is phosphorescent shell, this N. Atlantic species Named after the comblike spines This Indo-Pacific species has
and, like other species, lives in is especially abundant in on its shell, this Venus clam occurs a decoratively patterned shell,
burrows bored in wood or clay. muddy estuaries, where it on tropical American coastlines. as do many tellins.
burrows in soft sediment.
2–31/4 in 11/4–1½ in
5–8 cm 3–4 cm
SHIPWORM
Teredo navalis 21/4–31/4 in
f: Teredinidae 6–8 cm
½–¾ in This highly modified, widespread bivalve uses
1.5–2 cm its ridged shell valves as a drill to bore deep,
chalk-lined burrows in wood, damaging ships.

WAT E R I N G P O T S RED CALLISTA

Callista erycina
RINGED DOSINIA
Dosinia anus
A N D R E L AT I V E S f: Veneridae f: Veneridae
Many Venus clams are prized This Venus clam is found
The order Pholadomyoida includes clams by collectors, including this on the coastlines of New
and tropical watering pots. Scarcely Indo-Pacific species, also Zealand, and is one of
recognizable as bivalves, watering pots known as the sunset clam. many species harvested for
live encased in chalky tubes, and draw human consumption.
algae within
detritus and water into the tube through mantle exposed
a perforated plate at the anterior end. to sunlight

6–6½ in GIANT CLAM large, scale-

15–17 cm Tridacna gigas like flutes 12–16 in
f: Cardiidae 30–40 cm
PHILIPPINE WATERING POT The world’s largest 31/4–4½ ft
FLUTED GIANT CLAM
Verpa philippinensis bivalve, this long-lived 1–1.4 m Tridacna squamosa
f: Penicillidae and now endangered f: Cardiidae
This bizarre Indo-Pacific watering pot belongs to a coral-dwelling animal Like other Indo-Pacific giant clams, this species
family named for their wide, perforated ends, fringed lives on sandy beds in opens its shell during the day so that algae in its
with tubules. It lives partially buried in sediment. the Indo-Pacific. colored mantle can photosynthesize to make food.
308 G A S T RO P O D S
Gastropods are by far the largest class sea slugs. Gastropods originated in the oceans PHYLUM MOLLUSKA
of mollusks. The word gastropod and they still have their greatest diversity there;
INVERTEBRATES • GASTROPODS

CLASS GASTROPODA

means “stomach-foot,” because the other species live in fresh water or on land. ORDERS 21
FAMILIES 409
animal appears to crawl on its belly. SPECIES About 67,000
tor s ion
Most snails and slugs glide along a stream of slime The juvenile gastropod undergoes a process
on a single muscular foot, rasping at their food called torsion: the entire body within the shell D E B AT E
with a radula, a tonguelike sandpaper. Most twists around 180 degrees, so that the respiratory A COMMON TWIST
gastropods use their radula to rasp at vegetation, mantle cavity comes to lie above the animal’s
algae, or the fine film of microbes that coat head. This allows the vulnerable head to withdraw Sea snails were originally grouped
underwater rocks, but some are predatory. They into the safety of the shell. In marine snails, such together because their body twists
as periwinkles and limpets, this body form persists during development so that their gills
usually have a distinct head with well-developed
face forward. But the ancestor of all
sensory tentacles. Snails have a coiled or helical into adulthood. Because of this, marine snails are snails probably had this feature, and
shell into which they can withdraw; slugs lost described as prosobranchs, meaning “forward now marine snails are split into groups
their shell during the course of evolution. The gills.” In sea slugs, however, the body twists based on other characteristics. Their
class also includes a number of animals that back again, and they are called opisthobranchs, exact relationships remain debatable.
deviate from this basic plan, such as swimming or “hind gills.”

T RU E L I M PE T S 11/4–2 in
NERITES AND
Primitive algae-grazing limpets, the 3–5 cm R E L AT I V E S
Patellogastropoda have a slightly coiled, A small but diverse group, Cycloneritimorpha
conical shell. Their powerful muscles are well known in the fossil record. They include
clamp down firmly on intertidal rocks, marine, freshwater, and land-living forms, some
protecting them from predators, with a coiled shell, and a few like limpets. There ¾–2 in
desiccation, and waves. are also species with a trapdoor to their shell. 2–5 cm
½–¾ in
ZIG-ZAG NERITE 1.2–2 cm BLEEDING
COMMON LIMPET Neritina communis TOOTH NERITE
Patella vulgata f: Neritidae Nerita peloronta
f: Patellidae An inhabitant of Indo-Pacific f: Neritidae
This species grazes algae from rocks on mangroves, this is a highly Named for the blood-red mark on
shores of the N.E. Atlantic at high tide, variable species, with white, its shell opening, this Caribbean
then returns to a depression in the black, red, or yellow shells even intertidal mollusk can survive out
rock that matches the shape of its shell. in the same population. of water for prolonged periods.

TOP SH E L L S A N D R E L AT I V E S ¾–1 in
2–2.5 cm
The Vetigastropoda are marine gastropods that graze algae and microbes with a
brushlike radula. The shells range from those of keyhole limpets, with the barest
suggestion of a spire, to the coiled globes and pyramids of top shells, which can seal
themselves inside using an operculum, or trapdoor.
31/4–4¾ in
8-12 cm GIANT TOP SHELL LISTER’S KEYHOLE LIMPET CHECKERED TOP SHELL
Rochia nilotica Diodora listeri Phorcus turbinatus
f: Tegulidae f: Fissurellidae f: Trochidae
Many top shells have a Keyhole limpets are close relatives of Top shells have a “spinning-top”
thick mother-of-pearl lining. abalones and top shells. Like others, shell that is closed with a
This large Indo-Pacific this W. Atlantic species ejects circular operculum. This is a
species is used to oxygen-depleted water Mediterranean species.
make jewelry. through its “keyhole.”

shell shaped like

spinning top 8–12 in
20–30 cm
2–2¾ in ½–1¾ in
banded 5–7 cm 1.5–4.5 cm RED ABALONE
pattern Haliotis rufescens
f: Haliotidae
SILVER MOUTH Abalones have an ear-
TURBAN SHELL shaped shell lined
Turbo argyrostomus with thick mother-of-
f: Turbinidae pearl and punctuated
Turban shells are close with holes for exhaling
relatives of top shells, but water. This N.E. Pacific
have a calcified operculum. kelp-grazing species
This is an Indo-Pacific species. is the largest.
TOW E R SH E L L S PERIWINKLES, WHELKS,
A N D R E L AT I V E S A N D R E L AT I V E S 309
These tall-spired snails typically live in muddy or The largest and most diverse order of marine snails, the Caenogastropoda
sandy sediment. They feed on particles in the water are divided into several groups. Epitoniids, including wentletraps (bottom-

INVERTEBRATES • GASTROPODS
that they circulate through their mantle cavity. The living) and violet snails (floating), are specialist predators of cnidarians.
Cerithioidea are found in marine, freshwater, and Littorinids graze on algae, and include periwinkles, cowries, and conches.
estuarine habitats. They are slow-moving and often Whelks and their relatives are predators that project a long siphon out
group together in very large numbers. through a groove in the shell.
¾–1½in
2–4 cm
GREAT SCREW SHELL PRECIOUS
1–21/4 in 23/8–6 ½ in 1–2¾ in WENTLETRAP
2.5–5.5 cm Turritella terebra 6–17 cm 2.5–7 cm Epitonium scalare
f: Turritellidae f: Epitoniidae
A filter feeder of muddy sediments, this Wentletraps (from the
Indo-Pacific snail belongs to a group variously German for spiral staircase)
known as tower shells or auger shells. prey on anemones and corals
and have cutting jaws. This
ROUGH CERITH WEST INDIAN is an Indo-Pacific species.
Rhinoclavis aspera WORM SHELL LARGE VIOLET SNAIL
f: Cerithiidae Vermicularia spirata Janthina janthina
Ceriths are snails often f: Turritellidae 1–6½ in f: Epitoniidae
abundant in shallow-water Free-floating males of this 2.5–16 cm Violet snails float in tropical waters,
tropical marine Caribbean unwound screw shell preying on floating cnidarians. They
sediments. Like others, attach to solid material, often secrete mucus to create rafts of
this Indo-Pacific species embedding themselves in sponges. bubbles that keep them buoyed up.
lays strings of eggs They then develop into larger
attached to solid material. sedentary females. TIGER COWRIE
Cypraea tigris
f: Cypraeidae
The fleshy lobes of a cowrie’s mantle
older spikes SUNBURST CARRIER wrap right around the smooth shell
often broken Stellaria solaris 4–6 in when it crawls. This Indo-Pacific
f: Xenophoridae 10–15 cm species preys on other invertebrates.
Carrier snails cement
objects such as pebbles or ATLANTIC SLIPPER ¾–2 in
shells of other animals to LIMPET 2–5 cm
their shell for camouflage. Crepidula fornicata
This species inhabits f: Calyptraeidae
Indo-Pacific waters. Unrelated to true limpets,
½-21/4 in FOOL’S CAP filter-feeding slipper
1.5–6 cm Capulus ungaricus limpets form mating
f: Capulidae towers, with smaller males
2¾-5 in
This North Atlantic on top changing sex to
6–13 cm gastropod resembles replace dead females beneath.
unrelated limpets,
attaching itself to stones or
even the shell of other
mollusks, such as scallops.
COMMON
PELICAN’S FOOT
Aporrhais pespelecani
f: Aporrhaididae
Cousins of conches, pelican’s foot snails
are mud-dwelling detritus-eaters with
extensions to their shell that look like
webbed feet. This species inhabits the
Mediterranean and the North Sea.
12–16½ in
30–42 cm

attached detritus

¾–11/4 in 6–12 in
2–3 cm 15–31 cm

COMMON PINK CONCH

PERIWINKLE Aliger gigas
Littorina littorea f: Strombidae
f: Littorinidae Conches are mostly tropical,
Periwinkles, such as this European medium to large marine grazing
species, form a family of intertidal snails. Strombus conches, such as
snails with a globular spired shell this giant W. Atlantic species,
closed by an operculum. have a flare-lipped shell.
 PERIWINKLES, WHELKS, 3∕16–11/4 in 31/4–41/4 in
310 A N D R E L AT I V E S 5 mm–3 cm 8–11 cm
INVERTEBRATES • GASTROPODS

GIANT FROG SHELL

Tutufa bubo
f: Bursidae
4–121/2 in Tropical marine frog shells 4–20 in
10–32 cm are warty. This Indo-Pacific 10–50 cm
species eats bristleworms
using a proboscis, first
anesthetizing them TRUMPET TRITON POLI’S NECKLACE SHELL COMMON NORTHERN
with saliva. Charonia tritonis Euspira nitida WHELK
f: Charoniidae f: Naticidae Buccinum undatum
Tropical intertidal tritons and This is a European member of f: Buccinidae
related frog and helmet shells are a family of sand-burrowing A large predator of other mollusks
predators of other invertebrates. predators of bivalves. Necklace and tube worms, this North Atlantic
warty shell This Indo-Pacific species shells are named for their snail also consumes carrion, and is a
eats the coral-feeding necklacelike ribbons of spawn. popular seafood.
crown-of-thorns starfish.

3/4–11/4 in 11/4–11/2 in
2–3 cm 3–4 cm

canal for
siphon

21/4–41/4 in 31/2–6 in
6–11 cm 9–15 cm

PRICKLY PACIFIC DOG WHELK TEXTILE CONE

DRUPE Nucella lapillus Conus textile
Drupa ricinus f: Muricidae f: Conidae
f: Muricidae This North Atlantic snail Cone shells use their radula
A member of the dog belongs to a whelk family that to harpoon their prey and
whelk family, this snail catch prey by using the radula inject venom. Some, such as
lives on coral reefs of the to drill holes in the shells of this Indo-Pacific species, can
Indo-Pacific, where it preys barnacles and mollusks, helped be harmful to humans.
on bristle worms. by secretions of enzymes.
3–41/2 in 1⅛–5 in
7.5–11.5 cm 3–13 cm

BANDED TULIP IMPERIAL HARP TENT OLIVE

Cinctura lilium Harpa costata Oliva porphyria
f: Fasciolariidae f: Harpidae 23/4–8 in
7–20 cm
f: Olividae
This is a Caribbean snail Harp snails are sand-living SUBULATE AUGER The largest of the olive
related to Buccinum whelks. predators of crabs, trapping Terebra subulata shells, this species from the
Other sand-dwelling species them with their broad foot and f: Terebridae Pacific coasts of Mexico and
with a longer siphonal canal digesting them with saliva. This Typical of many augers, this Indo-Pacific species has a patterned shell. S. America has a colorful,
are called spindle shells. is an Indian Ocean species. Augers burrow through surface layers of sand to prey on worms. glossy shell.

F R E S H WAT E R
GILLED SNAILS
COMMON
The Architaenioglossa are the only order of gilled snails containing no marine RIVER SNAIL
species. Most live in fresh water, but some live on land. Apple snails have gills in Viviparus viviparus
f: Viviparidae
the mantle cavity that can function as a lung, allowing them to survive periods A European freshwater
of drought. All species have an operculum, or trap door, for closing their shell. snail with gills, this is a
relative of the apple snails
4–6 in CHANNELED and, like them, has a trapdoor
10–15 cm APPLE SNAIL (operculum) to close its shell.
Pomacea canaliculata
f: Ampullariidae
A typical apple snail, this
tropical American species 11/4–11/2 in
has been introduced 3–4 cm
elsewhere and has become
sensory an invasive pest.
tentacles
muscular
foot
SEA HARES SWIMMING SEA SLUGS
Sensory water-tasting head stalks, a common The Gymnosomata are also called sea angels, because winglike flaps
311
feature of sea slugs, are so big in the large sea hares they have evolved flaps from their muscular foot and use for swimming
that they resemble ears. The Anaspidea have a them to fly through water. The related sea butterflies or

INVERTEBRATES • GASTROPODS
small internal shell and, like sea angels, can swim Thecosomata also have flaps, but retain a fragile shell.
with their foot flaps.
foot flap
2¾–8 in
7–20 cm

SPOTTED SEA HARE COMMON SEA ANGEL

Aplysia punctata Clione limacina 1½–2 in
f: Aplysiidae f: Clionidae 4–5 cm
Like other seaweed-eating sea hares, this Sea angels swim freely with soft, transparent “wings.”
European species gathers in large breeding groups. As This cold-water species preys on a related gastropod
a defense, it releases ink when disturbed by predators. called the Arctic sea butterfly (Limacina helicina).

T RU E SE A SLUGS
The Nudibranchia are the largest group of sea slugs. 2–31/4 in
Nudibranch means “naked gills” and refers to the fact 5–8 cm
that these animals’ gills are exposed on their back,
rather than enclosed in a mantle cavity. Many species
have vibrant color patterns to advertise their toxicity.

4–5 in 1½–2 in
10–13 cm
BLACK-MARGINED OPALESCENT SEA SLUG 4–5 cm
SEA SLUG Hermissenda crassicornis
Doriprismatica atromarginata  f: Myrrhinidae
f: Chromodorididae This intertidal northern Pacific species belongs to a
This common Indo-Pacific sea slug lives in group of sea slugs with body outgrowths that store
shallow waters, feeding on sponges. It varies the still-potent stinging organs of their jellyfish prey.
¾–2 in in color from off-white to pale yellow.
2–5 cm

VARICOSE SEA SLUG ANNA’S SEA SLUG

Phyllidia varicosa Chromodoris annae external
f: Phyllidiidae f: Chromodorididae gills
This Indo-Pacific sea slug is a Like other Chromodoris sea slugs,
common inhabitant of rocks, this variable species from the
rubble, and sand in coastal waters, western Pacific Ocean specializes
where it preys on sponges. in feeding on sponges.
rhinophores
(scent detectors)
2¾–31/4 in head end
7–8 cm

ELEGANT SEA SLUG

Okenia elegans
f: Goniodorididae
Typical of its family, this sea slug feeds
on sea squirts. It occurs in waters around
Europe, including the Mediterranean.
gills obtain
oxygen
4–4¾ in
10–12 cm

mantle

VARIABLE NEON SEA SLUG SPANISH DANCER

Nembrotha kubaryana Hexabranchus sanguineus
f: Polyceridae 12–16 in f: Hexabranchidae
This Indo-Pacific sea slug preys on sea squirts and 30–40 cm A giant Indo-Pacific sea slug, this species gets its name
incorporates their defensive chemicals into its from its free-swimming movements; its red mantle is said
mucus. Many of its relatives prey on bryozoans. to resemble the ruffled skirt of a flamenco dancer.
 F R E S H WAT E R A I R- shell opening coiled shell
312 BR E AT H I NG SNA I L S to left

Snails of the superorder Hygrophila have their mantle cavity

developed as a lung; unlike most marine gastropods, they must
return to the water surface to breathe. Mainly herbivorous,
they are common in alkaline or neutral, weed-choked, stagnant ⅜–⅝ in
or slow-moving fresh water. 1–1.6 cm

GREAT POND SNAIL EUROPEAN

Lymnaea stagnalis BLADDER SNAIL
1–2 in f: Lymnaeidae Physella acuta
2.5–5 cm Widespread throughout f: Physidae
temperate parts of the northern Most snails have a shell that opens to the 1¼–11/2 in
hemisphere, this is a common right (when the opening faces the observer), 3–3.5 cm
pond snail in still or slow- but members of the freshwater physid
moving fresh water. family have a left-opening shell.

3∕16–5⁄16 in
5–8 mm COMMON RIVER LIMPET GREAT RAMSHORN SNAIL
Ancylus fluviatilis Planorbarius corneus
f: Planorbidae f: Planorbidae
Not a true limpet, but a limpetlike Ramshorn snails have a flat-coiled
member of the ramshorn snail shell, instead of the spire shell typical
family, this species is common in of snails. Most—like this Eurasian
fast-flowing European waters. species—occur in still, fresh water.

L A N D A I R- BR E AT H I NG 6–9 in
15–22 cm
1¼–11/2 in
3–3.5 cm
SNAILS AND SLUGS
The Stylommatophora are land snails and slugs that breathe air
using a lung in their mantle cavity. They have eyes at the tips GIANT AFRICAN SNAIL CUBAN LAND SNAIL
Lissachatina fulica Polymita picta
of their tentacles. Many are hermaphrodite and have both male f: Achatinidae f: Cepolidae 4–8 in
and female reproductive organs. Their courtship behavior This giant land snail from E. Africa This snail is found only in the mountain 10–20 cm
involves the exchange of spearlike darts before mating. has been introduced to many other forests of Cuba. There are many varieties
warm parts of the world, with different-colored shells.
becoming an invasive pest.
1–1¾ in ¼ in
2.5–4.5 cm 3¼–4¾ in 7–8 mm
8–12 cm

shell

BROWN GARDEN COMMON DOMED DISK SNAIL EUROPEAN

SNAIL SHELLED SLUG Discus patulus BLACK SLUG
Cornu aspersum Testacella haliotidea f: Discidae Arion ater
f: Helicidae f: Testacellidae Discid snails make up a f: Arionidae
This variably colored A typical member of family characterized by Black specimens of this
species with a wrinkled a family of slugs with a certain primitive features European slug predominate
shell is widespread in small external shell, this and a flat, coiled shell. in the north, but other
European woods, European species preys This is a N. American individuals are orange.
hedgerows, dunes, on earthworms. woodland species. Although it eats plants, it
and gardens.  also clears garden waste.
6–10 in
eye on tip 15–25 cm
of tentacle PACIFIC
BANANA SLUG
Ariolimax columbianus
f: Ariolimacidae
1¼–2 in Named for its yellow color,
3–5 cm the Pacific banana slug lives in
wet coniferous forests on the
western coast of N. America.

ROMAN SNAIL
Helix pomatia
f: Helicidae BROWN-LIPPED ¾–1¼ in 4–12 in
Widespread in C. Europe SNAIL 2–3 cm 10–30 cm
on calcium-rich soils, Cepaea nemoralis ASHY-GRAY
this is the largest snail f: Helicidae SLUG
of the region and is A close relative of the Roman Limax cinereoniger
bred locally for human snail, this W. European f: Limacidae
consumption. species has a highly variable Keel-back slugs have a small
shell color, and banding internal shell. This large species of
patterns for camouflage European woodland exists in a
in different habitats. number of color varieties.
CEPHALOPODS 313
Cephalopods are agile hunters in the gills and is then expelled through a short funnel. PHYLUM MOLLUSKA
mollusk phylum. Their sophisticated Cephalopods can move backward quickly by

INVERTEBRATES • CEPHALOPODS
CLASS CEPHALOPODA

nervous system enables them to hunt expelling water through the funnel, to propel ORDERS 9

down fast-moving prey. themselves forward. Squid, cuttlefish, and some FAMILIES About 50

octopus have fins on the sides of the mantle, used SPECIES 822

Cephalopods include the most intelligent of all for open-water swimming. Most octopus spend
invertebrates. Many species signal their mood their time on the sea floor.
using pigment-containing structures in their skin Only the open-ocean nautilus has a coiled shell.
called chromatophores. The class is organized In squid the shell is reduced to a penlike structure
by number of arms. Squid and cuttlefish have that gives some internal support, in cuttlefish to
eight arms used for swimming, plus two longer a similar, calcified structure called a cuttlebone.
retractable arms, called tentacles, with suckers Most species of octopus have lost their shell.
for gripping prey. Octopus lack tentacles, but have Cephalopods are fast-moving predators that use
suckers on all eight of their arms. The cephalopod their arms to grasp prey, and a parrotlike beak to
mantle encloses a cavity, which contains the gills. dispatch it. Squid catch free-swimming prey, while
A North Pacific giant octopus
To obtain oxygen, cephalopods take in water cuttlefish and octopus take slower, bottom-crawling discharges black-pigmented ink in
through the sides of the mantle. It passes over the crustaceans, such as crabs. self-defense as it jets away from a predator.

18–20 in 18–20 in
45–50 cm 45–50 cm

6–9½ in
15–24 cm
BROADCLUB
CUTTLEFISH
CHAMBERED NAUTILUS Sepia latimanus
Nautilus pompilius f: Sepiidae
f: Nautilidae AUSTRALIAN GIANT CUTTLEFISH This large cuttlefish is wide-ranging
This Indo-Pacific species is the Sepia apama throughout the Indo-Pacific region.
largest and best-known living member f: Sepiidae It is an abundant predator on coral reefs,
of a small group of cephalopods The largest known cuttlefish, this animal inhabits feeding on prawns and shrimp.
otherwise known only as fossils. waters off the southern coast of Australia, living
in seagrass beds and rocky reefs.
16–20 in
40–50 cm
8–20 in
20–50 cm

COMMON CUTTLEFISH 21/4–23/4 in

Sepia officinalis 6–7 cm WHIP-LASH SQUID
f: Sepiidae Mastigoteuthis sp.
Like many other cuttlefish, this f: Mastigoteuthidae
species migrates inshore to FLAMBOYANT CUTTLEFISH Reddish whip-lash squid such
spawn on muddy sediments. It Metasepia pfefferi as this species hover with broad fins in
occurs off the coasts of Europe
f: Sepiidae prey-snatching deep water, waiting for prey with its
Unusual among cuttlefish for its habit of tentacles
and South Africa. long tentacles extended.
using its tentacles to walk on ocean floors,
this Indo-Pacific cuttlefish has recently
been shown to be poisonous.
3/4–11/4 in 10–14 in 12–18 in 1½–13/4 in
2–3 cm 25–35 cm 30–45 cm 3.5–4.5 cm

BERRY’S BIGFIN REEF

BOBTAIL SQUID SQUID
Euprymna berryi Sepioteuthis lessoniana
f: Sepiolidae f: Loliginidae COMMON SQUID RAM’S HORN SQUID
Bobtail squid, such as this A pair of large fins makes this Loligo vulgaris Spirula spirula
Indo-Pacific species, are Indo-Pacific squid resemble a f: Loliginidae f: Spirulidae
small cuttlefish relatives. cuttlefish. It communicates This commercially important squid This small, deep-ocean cephalopod
They have a pen, not by flashing light from special is common in the N.E. Atlantic and contains a spiral gas-filled shell that
cuttlebone, and the rounded organs, which contain Mediterranean. Like related species, acts as a float; the animal rises
body has lobed fins. light-emitting bacteria. it has prominent side fins. toward the ocean surface at night.
314 COM MON OCTOPUS
O c topu s v u lgar is
Whether it is extricating a lobster from a lobster pot or hunting a scuttling crab,
INVERTEBRATES • CEPHALOPODS

the common octopus is one of the most intelligent of all the invertebrates. This
sea-living mollusk has excellent vision and eight grasping arms, or tentacles, which
are also used for crawling. The octopus can change color in an instant and squeeze
through very narrow crevices. And its horny beak can bite into prey, whose
scattered shell fragments frequently litter the sand around its lair. But in spite of
its versatility, the common octopus is short-lived. Upon hatching, an infant octopus
enters the oceanic plankton with half a million of its siblings, before descending
to the seabed two months later. If it survives being eaten, it takes
little more than a year to mature and spawn its own size Tentacle span 5–10 ft (1.5–3 m)
habitat Rocky coastal waters
offspring before dying. The common octopus is distribution Widespread in
widespread in tropical and warm temperate tropical and warm temperate waters
oceanic waters, and may consist of several diet Crustaceans, shelled mollusks
similar species.

fleshy mantle encloses a cavity

that contains vital organs,
including gills for breathing

INTELLIGENT
MOLLUSK
An octopus is an agile
predator with an advanced
nervous system. Two-thirds
of its neurons are located in
its arms, which operate with
a remarkable degree of
independence from its brain.
Experiments show that it also
has considerable problem-
solving intelligence, and
excellent long- and short-
term memory.
 UNDERSIDE
The octopus is a cephalopod, which literally means
“head-footed.” Its eight mobile arms radiate from 315
the underside of the head, where the mouth can
be seen at the center. BEAK

INVERTEBRATES • CEPHALOPODS
This octopus is a
predator of crustaceans,
among other prey. Its
jaws are a parrotlike
beak, which is strong
enough to penetrate
the tough carapace of
a crab or lobster.

SKIN
The skin contains special
cells called chromatophores.
FUNNEL, OPEN AND CLOSED These contain pigment
On one side of the mantle, just behind the which can change the
head, is a funnel, which the octopus uses octopus’s color—to blend in
in three ways: to expel water after gills have with its surroundings SUCKER
extracted oxygen; to rapidly discharge water or to signal its mood if Each arm has two series of suckers.
as jet propulsion for a quick getaway; and angry or afraid. These give the octopus an excellent
to squirt a confusing cloud of ink at the grip, allowing it to negotiate
enemy before fleeing. the sea bottom and coral reefs
with ease. The suckers also have
receptors that enable the octopus
to taste what it is touching.
prominent eye
with a horizontal,
slit-shaped pupil

tough, warty skin can

change color, texture, and
shape for camouflage

long, muscular arms for

moving over the seabed
and grasping objects

cuplike suckers
 CEPHALOPODS
316
INVERTEBRATES • CEPHALOPODS

4–6 in 8 in 11/4–21/4 in 61/2 ft

10–15 cm 20 cm 3–6 cm 2m
VAMPIRE
SQUID WINGED ARGONAUT
Vampyroteuthis infernalis DUMBO OCTOPUS Argonauta hians
f: Vampyroteuthidae Grimpoteuthis plena f: Argonautidae
This deep-sea cephalopod—with f: Opisthoteuthidae Argonauts—also known as paper
characteristics seemingly intermediate Named for its earlike fins used in nautiluses—are octopus relatives.
between squid and octopus—has fins swimming, this cephalopod lives at Females produce a paper-thin egg-case NORTH PACIFIC
projecting from its mantle and light- depths of 10–13,000 ft (3–4,000 m), that resembles a shell. This species is GIANT OCTOPUS
producing organs covering the body. preying on other invertebrates. distributed widely around the world. Enteroctopus dofleini
f: Enteroctopodidae
Perhaps the largest of all
octopuses, this animal is
juvenile 5–10 ft surprisingly short-lived.
1.5–3 m Females are diligent carers
of their huge broods.

39–59 in 20–28 in
PACIFIC LONG- CARIBBEAN REEF 1–1.5 m 50–70 cm
ARMED OCTOPUS OCTOPUS
Octopus sp. Octopus briareus COMMON OCTOPUS
f: Octopodidae f: Octopodidae Octopus vulgaris
One of several octopuses with An octopus of the W. Atlantic f: Octopodidae
especially long arms, adults of and Caribbean, this coral- Globally widespread in tropical and
this species frequent sandy dwelling species often traps its warm temperate waters, this octopus
lagoons, but translucent prey by spreading its webbed typically has a warty body and two
2 in juveniles live among arms like a net. rows of suckers on its arms.
5 cm oceanic plankton.

ATLANTIC OCTOPUS
Octopus sp.
f: Octopodidae
DNA analysis has revealed a
number of similar octopus
species, including this one,
which are related to the
yellow background common octopus, revealing
color hidden biodiversity.

39 in
1m

MIMIC
OCTOPUS
Thaumoctopus mimicus
f: Octopodidae
Many octopuses can change
color, but this Asian species
can change shape, too—
even disguising itself to
look like other marine
animals such as sponges,
corals, and jellyfish.

funnel

BLUE-RINGED OCTOPUS
Hapalochlaena lunulata
f: Octopodidae
This octopus preys on
crustaceans and fishes in the black and blue ring
W. Pacific, paralyzing them markings warn that
6–8 in with venomous saliva that is this is a highly
15–20 cm also potentially fatal to humans. venomous species
CHITONS 317
Flattened, rock-hugging chitons are although the shell itself contains cells that enable PHYLUM MOLLUSKA
among the most primitive of mollusks. them to react to light. The plates are fringed by

INVERTEBRATES • TUSK SHELLS

CLASS POLYPLACOPHORA

Most are grazers of algae and microbe the edge of the mantle, known as the girdle. The ORDERS 2

films, and occur in coastal waters. mantle forms a fleshy skirt around the chiton. FAMILIES 20

This overhangs grooves running down each side SPECIES 1,026

A chiton’s shell, popularly known as a “coat-of-mail” of the animal, which channel water, delivering
shell, is made up of eight interlocking plates that oxygen to gills that project into the grooves. The girdle
fringe
are sufficiently flexible to allow the animal to bend radula, or rasping tongue, is coated with minute
when gliding over uneven rocks—or even to roll up teeth reinforced with iron and silica, so chitons
when disturbed. Chitons have no eyes or tentacles, can graze on the toughest encrusting algae.

31/4 in/8 cm 11/2–2 in/4–5 cm 3/4–31/4 in/2–8 cm 1 in/2.5 cm

MARBLED CHITON GREEN CHITON WEST INDIAN DECKED CHITON

Chiton marmoratus Chiton glaucus FUZZY CHITON Ischnochiton comptus
f: Chitonidae f: Chitonidae Acanthopleura granulata f: Ischnochitonidae
Like other chitons, the shell A variable-colored chiton, f: Chitonidae Ischnochiton chitons have a
plates of this Caribbean this species occurs along the This spiky-fringed Caribbean spiny or scaly girdle
mollusk are composed coastlines of New Zealand and chiton can tolerate exposure fringe. This is a common
entirely of a chalky mineral Tasmania and, like most chitons, to sun, and can live high up intertidal species of
called aragonite. is active at night. in the intertidal zone. W. Pacific shores.

eight-plated
shell
12–13 in 2–31/4 in
30–33 cm 5–8 cm 11/2–23/4 in
4–7 cm

GUMBOOT
CHITON
Cryptochiton stelleri BRISTLED HAIRY 1/2–2 in
f: Acanthochitonidae CHITON CHITON 4–5 cm
In one chiton family Mopalia ciliata Chaetopleura papilio
the fleshy girdle f: Mopaliidae f: Chaetopleuridae LINED CHITON
overlaps the chain mail Named for its prominent A bristly-girdled species with Tonicella lineata
plates. In this North hairy girdle, this chiton of the brown-striped chainmail f: Ischnochitonidae
Pacific species, the Pacific coast of N. America is plates, the hairy chiton is This brightly colored chiton occurs on
largest chiton, it forms sometimes found beneath found beneath rocks on North Pacific coasts, where it may be camouflaged
a leathery skin. long-moored boats. South African coasts. when feeding on red encrusting algae.

TUSK SHELLS 11/4–11/2 in

3–4 cm

Curious, mud-shoveling tusk shells live PHYLUM MOLLUSKA

EUROPEAN TUSK
Antalis dentalis
in marine sediments and have a tubular CLASS SCAPHOPODA f: Dentaliidae
curved shell—open at both ends and ORDERS 2 A northeast Atlantic species, the
European tusk is widespread
FAMILIES 13
unlike the shell of any other mollusk. SPECIES 576
in sandy areas of coastal waters.
In places its empty shells are
Tusk shells are common, but are rarely seen because found in huge numbers.
they usually occur far offshore, with the wide base of the hollow tusk buried in mud. The 2–31/4 in
5–8 cm
eyeless head and foot reach deeper into the sediment, probing for food with their tentaclelike
structures called captacula, and tasting the mud with chemical detectors. When food BEAUTIFUL TUSK
is found—small invertebrates or detritus—the tentacles bring it to the mouth, where it is Pictodentalium formosum
f: Dentaliidae
ground up by the radula, a rasping tongue typical of mollusks. Tusk shells lack gills; the This colorful tropical
animal’s fleshy mantle encloses a water-filled, oxygen-extracting tube that runs the length tusk shell has been recorded
in marine sediments in Japan,
of the shell. When levels of oxygen run low, the tusk shell contracts its foot and squirts the the Philippines, Australasia,
stale water out through the top end of the shell. Fresh water enters by the same route. and New Caledonia.
318 ECHINODERMS
Echinoderms include an astonishing The water circulates through tubes that run through PHYLUM ECHINODERMATA
array of sea creatures, ranging from the animal and is forced into tiny, soft projections
INVERTEBRATES • ECHINODERMS

CLASSES 5

filter-feeding feather stars to grazing near the surface, called tube feet, which can move ORDERS 35

urchins and predatory starfish. back and forth and stick to surfaces. In feather FAMILIES 173

stars these tube feet point upward on the feathery SPECIES 7,447

This is the only large invertebrate phylum entirely arms, capturing particles of food, which are then
restricted to salt water. Echinoderms are sluggish, moved toward the central mouth. In other
slow-moving inhabitants of the sea floor and most echinoderms thousands of downward-pointing
have a body form built around a five-part radial tube feet are used in locomotion—together they
symmetry. Echinoderm means “spiny-skinned”— pull the animal over sediment and rock.
a reference to the hard, calcified structures, called
ossicles, that make up the internal skeleton of defense
these animals. In starfish these are spaced out The tough, prickly skin of most echinoderms
within the soft tissue so that the animal is offers good protection from potential predators,
reasonably flexible, but in urchins they are fused but these animals defend themselves in other
together to form a solid internal shell. In sea ways, too. Many urchins are also covered with
cucumbers the ossicles are so minute and sparsely formidable spines; some can inflict serious wounds
distributed—or even absent—that the entire in humans. Many species have tiny pincerlike
animal is soft-bodied. projections, sometimes venomous, that can
remove cluttering debris as well as deter potential
h y d r au l i c f e e t predators. Soft-bodied sea cucumbers rely on
Echinoderms are the only animals with a water noxious chemicals to protect them—and are
transport system. Sea water is drawn into the often brightly colored as a warning. As a last
The tiny tube feet on which echinoderms
center of the body through a perforated, sievelike resort, some sea cucumbers can even spew out move around the seabed are seen here on
plate, usually on the upper surface of the body. sticky tangles or eject their gut. a crown-of-thorns starfish, Acanthaster planci.

F E AT H E R STA R S
A feather star carries five basic filter-feeding arms, which in some species
branch into a dense cluster. A mouth and anus point upward from the center
of the star. Some members of the class Crinoidea crawl along on their
featherlike arms. Others, known as sea lilies, are attached by a stalk.

PRETTY FEATHER STAR

Cenometra emendatrix
f: Colobometridae
The arms of a feather star
are fringed with smaller
branches called pinnules
that catch particles of
food more effectively. In
this tropical Pacific
species they are white.
4–6 in
10–15 cm

feathery arms

coral
base

4–8 in 4–6 in RED

10–20 cm 10–15 cm FEATHER STAR
Himerometra robustipinna
GOLDEN FEATHER STAR YELLOW FEATHER STAR f: Himerometridae
Davidaster rubiginosus Anneissia bennetti This feather star inhabits
f: Comatulidae f: Comasteridae tropical Indo-Pacific coastal
Like other crinoids, this species Like many crinoids, this common waters, clinging to corals and
from the W. Atlantic is a W. Pacific species filter feeds most sponges. The pale clingfish
suspension feeder, catching voraciously at night and when lives among its arms, probably 4–6 in
plankton with its feathery arms. underwater currents are strongest. for protection from predators. 10–15 cm
U RC H I NS A N D
R E L AT I V E S 319
Sea urchins, which belong to the class Echinoidea,
4–4 ¼ in 1½–2 in 8 in
gave their name to the whole phylum. The hard 10–11 cm 4–5 cm 20 cm
ossicles of an urchin are plates that interlock to
form a shell called a test. The animal’s spines help INDO-PACIFIC SAND DOLLAR BANDED SEA URCHIN RED URCHIN
in locomotion and defense. The mouth generally Sculpsitechinus auritus Echinothrix calamaris Astropyga radiata
faces downward and the anus upward—with f: Astriclypeidae f: Diadematidae f: Diadematidae
This Indo-Pacific urchin of coastal The shorter spines of this Indo-Pacific This Indo-Pacific lagoon species
rows of tube feet running between the two. sands is a typical member of a group reef-dwelling urchin are painfully belongs to a family of tropical urchins
of burrowing sea dollars, named for venomous. Cardinalfishes (Apogonidae) with long hollow spines. It is often
their flattened appearance. often hide among them for protection. carried by a crab, Dorippe frascone.
3¼–4 in
8–10 cm
6½–7 in
16–18 cm
FIRE URCHIN 8–10 in
Asthenosoma varium 20–25 cm
f: Echinothuridae
This somewhat flat Indo-Pacific species
of sandy lagoons has a shell sufficiently
flexible for the urchin to enter
crevices. It can deliver a painful sting.

flexible test
MATHA’S URCHIN EDIBLE URCHIN
Echinometra mathaei Echinus esculentus
f: Echinometridae f: Echinidae
This Indo-Pacific species This large, globular urchin common
belongs to a group of urchins on N.E. Atlantic coasts has a
characterized by thick spines. variable color. It grazes on
It lives in crevices on coral reefs. invertebrates and algae,
especially kelp.

3¼–4 in 2¾–3¼ in
8–10 cm 7–8 cm
PURPLE URCHIN SEA POTATO
Strongylocentrotus purpuratus Echinocardium cordatum
f: Strongylocentrotidae f: Loveniidae
An inhabitant of the underwater Heart urchins, such as this globally
kelp forests along the Pacific widespread species, are detritus-feeding
coast of N. America, this burrowers that lack the obvious radial
urchin is often used in symmetry of other urchins. short, venomous
biomedical research. spines

SEA CUCUMBERS 2–3¼ in

5–8 cm
6–7 in
15–18 cm
Sea cucumbers are soft, tubular animals,
with a mouth surrounded by a ring of SEA APPLE CUCUMBER
food-collecting tentacles at one end, and Pseudocolochirus violaceus
an anus at the other. Some Holothuroidea YELLOW SEA CUCUMBER f: Cucumariidae
Colochirus robustus Sea apples are brightly colored,
shovel or burrow in sediments using their f: Cucumariidae highly poisonous, reef-dwelling
tentacles; others use rows of tube This echinoderm belongs to a family of thick- sea cucumbers. This species has
feet to crawl along the seabed. skinned sea cucumbers. Like this Indo-Pacific yellow or orange tube feet, but
species, many are covered in knobbly projections. variable body color.
sticky mass
ejected to 10–12 in
repel predators 25–30 cm warning
OCELLATED SEA coloration
CUCUMBER 15–23½ in
Bohadschia argus 38–60 cm
VERMIFORM SEA
f: Holothuriidae CUCUMBER
Distributed from the Synapta maculata
W. Indian Ocean around EDIBLE SEA CUCUMBER f: Synaptidae
Madagascar to the South Holothuria (Halodeima) edulis Typical of its family, this Indo-Pacific
Pacific, this large species is f: Holothuriidae species is a soft-bodied, wormlike
variable in color, but Large sea cucumbers are especially abundant 23½ in sea cucumber that burrows in soft
is always spotted. in tropical waters. This Indo-Pacific species is 60 cm sediment. It lacks tube feet.
harvested and dried for human consumption.

spikes provide grip 14–16 in

on ocean floor 35–40 cm

23½–30 in 12 in
GIANT CALIFORNIA 60–75 cm 30 cm
SEA CUCUMBER
Apostichopus californicus PRICKLY REDFISH DEEP SEA CUCUMBER
f: Stichopodidae Thelenota ananas Kolga hyalina
One of the largest sea cucumbers f: Stichopodidae f: Elpidiidae
of N. America’s Pacific coast, This echinoderm belongs to a family of sea Virtually worldwide in distribution at depths of up
this fleshy, spiked species is cucumbers covered in fleshy spikes. It occurs on to 5,000 ft (1,500 m), this is one of many little-
fished as a local delicacy. sandy seabeds around Indo-Pacific coral reefs. known sea cucumbers found on ocean floors.
 BR I T T L E STA R S 8–12 in
20–30 cm
SHORT-SPINED
BRITTLE STAR
320 These animals are named for the fact that their long, Ophioderma sp.
f: Ophiodermatidae
thin, sometimes branching, arms easily break off. A brittle star of W. Atlantic
There is one opening to the gut in the central disk—a coastal waters, this species
INVERTEBRATES • ECHINODERMS

downward-facing mouth. Some Ophiuroidea use their lives in submerged grass

beds, where it preys
arms to trap food particles; others are predatory. on other invertebrates,
such as shrimp.
long, flexible arms
COMMON BRITTLE STAR
Ophiothrix fragilis
f: Ophiotrichidae 8–10 in GORGON’S HEAD
This N.E. Atlantic brittle BLACK 20–25 cm BRITTLE STAR
star—often found in dense 8–12 in BRITTLE STAR Gorgonocephalus caputmedusae
populations—has spiny arms, 20–30 cm Ophiocomina nigra f: Gorgonocephalidae
some of which it holds up f: Ophiocomidae This species, named for its
to trap food particles. This large brittle star of coiled, snakelike branching
European waters can either arms, is common around
4¾–6 in filter feed on particles or European coasts. Larger
12–15 cm scavenge on detritus. It can animals are found in places
be common on rocky coasts of strong currents, where they
with strong currents. can collect more food.

STA R F I SH
Like most other echinoderms, starfish crawl on the seabed PURPLE
using their rows of tube feet. These run along grooves in the SUNSTAR
starfish’s arms. Most species of Asteroidea are five-armed, Solaster endeca
but a few have more than five arms, and some are virtually 14–16 in f: Solasteridae
35–40 cm Sunstars are large, spiny
spherical in shape. Many prey on other slow-moving starfish with many arms.
invertebrates; others scavenge or feed on detritus. Although Living on offshore mud 20–23½ in
their skin is studded with hard ossicles, in the cold waters of the 50–60 cm
starfish are sufficiently flexible to northern hemisphere, this
species has from 7 to 13 arms.
catch prey.
SEVEN ARM STARFISH
Luidia ciliaris
f: Luidiidae
8–9½ in MOSAIC STARFISH Unlike most other starfish, this large
20–24 cm Plectaster decanus Atlantic species has seven arms. It
f: Echinasteridae has long tube feet and is fast-moving
Like a number of other in its pursuit of other echinoderms.
members of its family,
this striking species of
S.W. Pacific rocky coasts
varies greatly in color.

tube feet on
underside of
broad base arms
8–10 in to arms
20–25 cm
WARTY STARFISH
Echinaster callosus
f: Echinasteridae
Belonging to a family of stiff-bodied RED CUSHION STAR
starfish with conical arms, this coarse, spiny Porania (Porania)
W. Pacific species is unusual for upper surface pulvillus
its pink and white warts. f: Poraniidae
This distinctive,
smooth-topped starfish
with short arms and 4–4¾ in
fringe of spines occurs 10–12 cm
on European rocky coasts,
4¾–5 in 4–10 in often on kelp holdfasts.
10–12 cm 10–25 cm

16–20 in 32–39 in
40–50 cm 80–100 cm

BLOOD HENRY STARFISH OCHER STARFISH COMMON STARFISH

Henricia oculata Pisaster ochraceus Asterias rubens GIANT SUNFLOWER STARFISH
f: Echinasteridae f: Asteriidae f: Asteriidae Pycnopodia helianthoides
This relative of the warty starfish lives This N. American Pacific relative of the This common N.E. Atlantic predator f: Asteriidae
in tidal pools and kelp forests of the common Atlantic starfish—like others of of other invertebrates is unusual among One of the world’s largest starfish, this many-armed
N.E. Atlantic. Its body secretes mucus its family—hunts invertebrates. It is an echinoderms in tolerating estuarine conditions. species preys on mollusks and other echinoderms. It
to trap food particles. important predator of mussels. It often occurs in huge local populations. lives among offshore seaweed on the N.E. Pacific coast.
 20–23½ in CROWN-OF-THORNS
50–60 cm STARFISH
Acanthaster planci 321
f: Acanthasteridae
This giant Indo-Pacific
starfish is a predator of coral 8–12 in
polyps, threatening some 20–30 cm
reef ecosystems. It has BLUE STARFISH
10–20 arms and sharp, Linckia laevigata
mildly venomous spines that f: Ophidiasteridae
can cause injury to humans.
Snake-armed starfish have a smooth,
spineless surface. This Indo-Pacific
species is usually blue, but a
minority are purple or orange.
It frequently lives in association
with detritus-eating shrimp.

8–10 in
20–25 cm
10–12 in
25–30 cm

PURPLE STARFISH KNOBBLY

Ophidiaster ophidianus STARFISH
f: Ophidiasteridae Pentaceraster cumingi
This snake-armed starfish f: Oreasteridae
occurs in warmer waters of This large, strikingly
the northeast Atlantic, on patterned cushion star occurs
coarse seabeds around the on sandy and rubble-strewn
Mediterranean and as far as sea bottoms of tropical
W. Africa. central and E. Pacific.

venomous spines
4–4¾ in
10–12 cm
2¼–2¾ in
6–7 cm

NECKLACE STARFISH CUMING’S PLATED

Fromia monilis STARFISH
f: Goniasteridae Neoferdina cumingi
There are 15 Fromia species, f: Goniasteridae
sometimes difficult to tell apart, This Pacific species lives on shallow
10–12 in 8–10½ in but all are brightly colored to reefs but others in the family occur at
25–30 cm 20–27 cm depths below 3,300ft (1,000m). All
deter predators. This common
shallow-water species is are characterized by rows of granular
rows of red widespread in the Indo-Pacific. plates on their upper surface.
tubercles 8–10 in
20–25 cm

RED GENERAL STARFISH GRANULATED STARFISH INDO-PACIFIC CUSHION STAR flattened body
Protoreaster lincki Choriaster granulatus Culcita novaeguineae
f: Oreasteridae f: Oreasteridae f: Oreasteridae
From the cushion star family, this A giant Indo-Pacific echinoderm This coral-feeder of the Indo-Pacific
Indian Ocean species grazes on algae of rubble slopes and coral reefs, belongs to a family of starfish that become
as a juvenile, but preys on other this cushion star grazes on algae shorter-armed as they get bigger, eventually
invertebrates when adult. and detritus in shallow waters. maturing into dumpy cushion shapes.
4–4¾ in
10–12 cm
6–8 in
groups of stiff
15–20 cm spines on short,
blunt arms 1½–2 in
4–5 cm SMALL CUSHION STAR
GOOSEFOOT STARFISH Asterina gibbosa
Anseropoda placenta f: Asterinidae
f: Asterinidae This short-armed variable- ICON STARFISH
A thin, flat starfish with colored starfish, from the Iconaster longimanus
ill-defined arms, the goosefoot same family as the goosefoot, f: Goniasteridae
starfish of the E. Atlantic scavenges dead material Many echinoderms develop from planktonic
is a predator of bottom- on the rocky coasts of larvae, but this Indo-Pacific starfish lacks a
dwelling crustaceans. the N.E. Atlantic. larval stage. It produces large, yolky eggs.
322
C H O R DAT E S
Only three to four percent of known animal species are
A NIM ALS • CHORDATES

chordates, yet they include the largest, fastest, and most

intelligent animals alive today. Most chordates have a skeleton
made of bone or cartilage, but their defining feature is the
rodlike notochord—the evolutionary forerunner of the spine.
Fossil records indicate that the earliest lifestyles
known true chordates were small, It is perhaps surprising that one of the
streamlined animals, just an inch long. simplest chordate groups, the sea squirts
Living at least 500 million years ago, (tunicates), do not have a skeleton or
they had no hard parts, except for a stiff backbone, and spend their adult lives
but flexible cartilaginous notochord. The fastened in one place. However, in their
notochord ran the length of their body, planktonic larval stage they do have a
creating a framework for their muscles notochord, indicating their shared
to pull against. Today’s chordates all have ancestry with vertebrate chordates such as
this feature, and a small number keep it us. In contrast to these simple chordates,
throughout life. However, in the vast vertebrate chordates are often fast-moving
majority of chordates—from fishes animals, with rapid reactions, a well-
and amphibians to reptiles, birds, and developed nervous system, and a sizeable TUNICATES
The subphylum Tunicata contains just over 3,000 species. Larval
mammals—the notochord is present brain. Birds, mammals, and a few fishes tunicates have a notochord, and resemble tadpoles. The adults
only in the early embryo. As the embryo use energy from food to keep their body are filter feeders, the most common type being sea squirts.
develops, the notochord disappears, at a constant optimum temperature. AMPHIBIANS
replaced by an internal skeleton made Chordates vary in the way they Numbering more than 8,200 species, amphibians include frogs
and toads, salamanders, newts, and caecilians. Most start their
of cartilage or bone. These animals are reproduce and raise their young. All lives as aquatic larvae, but as adults they take up life on land.
known as vertebrates, from the column mammals bear live young, except for the
of vertebrae that makes up their spine. duck-billed platypus and echidnas. Almost
Unlike shells or body cases, bony all other chordates lay eggs, though there
skeletons work on an amazing spectrum are examples of live-bearing species in all
of scales. The smallest vertebrate, a vertebrate groups except birds. The number
freshwater fish called Paedocypris of offspring produced is related to the
progenetica, is less than ⅜ in (1 cm) long amount of parental care. Some fishes
and weighs several billion times less than a produce millions of eggs, and play no part
blue whale, which is the largest chordate, in raising their young, whereas mammals
and the biggest animal that has ever lived. and birds have much smaller families.

CHORDATE LOBE-FINNED
FISHES
TREE CARTILAGINOUS REPTILES
TUNICATES FISHES AND BIRDS

CHORDATES VERTEBRATES
continued from
invertebrates
LANCELETS RAY-FINNED MAMMALS
FISHES
Tunicates and lancelets evolved
before the vertebrates, though which
is the oldest group is still debated. JAWLESS FISHES AMPHIBIANS
LANCELETS FISHES
Small marine chordates with slender bodies and a notochord Fishes are the most diverse and numerous of all the vertebrate chordates. They are classified into a number of different classes, within
throughout life, lancelets live half buried in the seabed. the subphylum Vertebrata. These classes of fishes are often grouped together as a superclass, Pisces, and their variety reflects their
There are 20 species in the subphylum Cephalochordata. evolutionary past. Almost 34,000 living species are known.

REPTILES BIRDS MAMMALS

With over 11,000 species, reptiles inhabit every continent Birds are the only living animals that have feathers. There are about Readily identified by their fur or hair (even whales have
on Earth, with the exception of Antarctica. Unlike birds 10,000 species of birds, though recent studies suggest there may be up some), mammals are the only chordates that raise their young
and mammals, they are cold-blooded, and covered in scales. to 18,000. All lay eggs, and many show highly developed parental care. on milk. There are more than 6,300 known mammal species.
324 FISHES
Fishes are the most diverse chordates and can be found in all types
ANIMALS • FISHES

of watery habitat—from tiny, freshwater pools to the deep ocean.

Almost without exception, they breathe using feathery gills to
absorb oxygen from the water, and almost all swim using fins.

PHYLUM CHORDATA Fishes are not a natural, single group but are, in fact,
CLASSES PETROMYZONTI seven classes of vertebrate chordates, of which the
MYXINI
ELASMOBRANCHII
familiar ray-finned (bony) fishes are by far the most
HOLOCEPHALI populous. The majority of fishes are cold-blooded
ACTINOPTERYGII
COELACANTHI
and so their body temperature matches that of the
DIPNEUSTI surrounding water. A few top predators, such as white
ORDERS 66–81 sharks, can maintain a flow of warm blood to the
FAMILIES 560–588
SPECIES About 33,900
brain, eyes, and main muscles, which allows them to
hunt actively even in very cold water. Most fishes are
protected by scales or bony plates embedded in the
skin. In fast swimmers these are lightweight, and
provide streamlining as well as protection from
abrasion and disease. Although a few fishes slither or
shuffle over the seabed, most swim using fins. With
the notable exception of rays, the tail fin usually
provides the main propulsive force. Paired pectoral
and pelvic fins (on the sides and underside respectively)
provide stability and maneuverability, helped by up to
three dorsal fins on the fish’s upper side and one or
two anal fins. Fishes avoid collisions—especially in
Overlapping scales cover the body large shoals—by using special sensory organs that
of a bicolor parrotfish, forming a flexible
layer that protects against injury. detect the vibrations made as they or other animals
move through the water. Most fishes have a row of
these organs running along both sides of the body,
called the lateral line. While they also use the senses
of hearing, touch, sight, taste, and smell found in other
chordates, only fishes have a lateral line system.

r e p ro d u c t i v e s t r at e g i e s
The seven classes of fishes have very different ways
of reproducing. Most ray-finned fishes have external
fertilization and shed masses of eggs and sperm
Predatory sharks lunge into a huge directly into the water to compensate for the huge
shoal of sardines, which swim ever closer numbers that are eaten before they can develop into
together for protection.
juvenile fishes; a few instead guard their eggs until
hatching. In contrast, cartilaginous fishes (sharks,
rays, and chimaeras) all have internal fertilization and
produce eggs or young at an advanced developmental
stage. This requires a high input of energy—only
a few young are produced at one time, but these
have a good chance of survival. The eggs of lampreys
(jawless fishes) hatch into larvae that live and feed
for many months before undergoing metamorphosis
into the adult form.
The huge mouth of a male gold-specs BRIGHT SPECTACLE
jawfish provides an unlikely nest for its The fabulous colors of the Banggai cardinalfish, seen here
eggs. He will not feed during incubation. sheltering among giant anemones, attract aquarium owners.
f i s h g ro u p s
The seven classes show fundamental
differences, especially in skeleton
and gills, but also many similarities
in body form required to function
in an aquatic environment.
j aw l e s s f i s h e s

326
c a rt i l ag i n o u s
fish es

327
r ay - f i n n e d f i s h e s

334
lobe-f i n n e d f ish e s

353
326 J AW L E S S F I S H E S
Unlike other vertebrates, jawless fishes as ammocoetes. These live in mud and feed on PHYLUM CHORDATA
have no biting jaws, although they do detritus. After about three years the larvae of
F I S H E S • J AW L E S S F I S H E S

CLASSES PETROMYZONTI

have teeth. Once abundant and diverse, the anadromous species change into adults and MYXINI

few living representatives remain. swim out to sea where they feed for several years. ORDERS 2

The freshwater species live and breed in rivers FAMILIES 4

SPECIES 128
There are two living groups of jawless fishes— and lakes. Hagfishes lay eggs on the seabed.
lampreys and hagfishes—and the relationship
between them is still debated. Instead of jaws, pa r a s i t e s a n d s c av e n g e r s D E B AT E
lampreys have a round, sucker mouth encircled by Lampreys are notorious for feeding parasitically on ARE LAMPREYS AND
rasping teeth, while hagfishes have a slitlike mouth larger fishes when they are adult. Clinging on with H A G F I S H E S R E L AT E D ?
with teeth inside on the tongue. Circular gill its sucker mouth, a lamprey grinds its way through
Traditionally lampreys and hagfishes
openings run along the sides, seven in lampreys the skin of its victim eating either, or both, flesh were classified together as
and a variable number (1–16) in hagfishes. For and blood. They can be a nuisance to fishermen, Cyclostomata, separate from jawed
body support and muscle attachment, both groups as they damage and kill fishes in nets and fish farms. vertebrates. Detailed morphological
have a simple, rodlike structure called a notochord. However, many lampreys, especially freshwater comparisons and early molecular
In addition, lampreys may have a few cartilage species, just eat small invertebrates. The sucker work allied lampreys with jawed
supports around the notochord. mouth is also useful when the fishes are swimming vertebrates, leaving hagfishes as a
upstream against the current—they use it to cling non-vertebrate chordate. Since about
d i f f e r i n g l i f e c yc l e s onto rocks so they can have a rest and to move 2010, advanced genetic techniques
While hagfishes live permanently in the deep pebbles when excavating a nest site in the river bed. (of micro-RNAs) have provided strong
evidence for grouping them together
ocean, lampreys inhabit temperate coastal waters Hagfishes live in mud burrows in the seabed, once again.
and fresh waters worldwide. All lampreys breed emerging at night to feed on both live and dead
in fresh water. Coastal species are anadromous— invertebrates. If the opportunity arises, they will feed
that is, like salmon, they swim up rivers into fresh on rotting whale or fish carcasses, from which they
4 ft/1.2 m
water to spawn, after which they die. The eggs rasp off bits of flesh. If attacked, hagfishes produce
hatch into wormlike, burrowing larvae, known copious, gill-clogging slime to deter the predator.

L A M PR EYS AN D H AGFISH ES
The orders Petromyzontiformes and Myxiniformes are the only two SEA LAMPREY
Petromyzon marinus
groups of living jawless fishes—often called agnathans. They share an f: Petromyzontidae
eellike shape, lack of jaws, and possession of a notochord, but also have Salmon in the North Atlantic sometimes
numerous anatomical differences. Lampreys have a long larval stage fall prey to this parasitic fish, which latches
spent in fresh water, while hagfishes lay eggs on the seabed that hatch onto its victim and scrapes out its flesh.
into miniature adults.
6 1/2 in
first 16 cm
dorsal fin

3 1/4 ft
1m

gill openings
BROOK LAMPREY
Lampetra planeri
f: Petromyzontidae
Larvae and adults of this common northern
European lamprey live in streams and rivers.
Once metamorphosis has taken place, the
adults stop feeding and devote all their 30 in
energy to spawning, after which they die. PACIFIC LAMPREY 76 cm
Entosphenus tridentatus
f: Petromyzontidae sucker mouth with
Some populations of this lamprey are land- rasping teeth
rows of backward- locked in lakes. Those in the ocean rasp flesh
pointing teeth and blood from fishes and sperm whales.

16 in temporary
40 cm knot in body

ATLANTIC HAGFISH 5 ft
Myxine glutinosa tail tapers to 1.5 m
f: Myxinidae thin filament
Hagfishes detect food by smell and
their sensory tentacles. They knot their
body to give leverage when feeding and to
slide off defensive slime. This species lives
in the N. Atlantic and Mediterranean.
C A RT I L AG I N O U S F I S H E S 327
These fishes have a skeleton of pliable live entirely in rivers. Most open-water cartilaginous PHYLUM CHORDATA
cartilage rather than the hard bone fishes are continually on the move because, unlike

FISHES • CARTILAGINOUS FISHES

CLASSES ELASMOBRANCHII

found in most other vertebrate groups. bony fishes, they do not have a gas-filled swim HOLOCEPHALI

Most are predators with acute senses. bladder to maintain neutral buoyancy and therefore ORDERS 14–17

may sink if they stop swimming. The whale shark FAMILIES 57

SPECIES 1,338
Sharks, rays, and chimaeras are all cartilaginous and other surface-feeding species have a large, oily
fishes, although chimaeras have distinct anatomical liver to prevent this. Predatory sharks are well
differences from the other two groups. In known for their amazing ability to smell blood
chimaeras, the upper jaw is fused to the braincase and hone in on wounded fishes and mammals.
and so cannot be moved independently. They also Cartilaginous fishes also have the ability to detect
have teeth that grow continuously. In contrast, the weak electrical fields that surround living
sharks regularly lose their hard, enamel-covered creatures and, while not unique to them, this sense
teeth but replace them from extra rows lying flat is developed to an extraordinary extent in this group.
behind the active teeth—a characteristic that
helps make sharks some of the most formidable r e p ro d u c t i o n
predators on Earth. The skin of all three groups— All cartilaginous fishes mate and have internal
sharks, rays, and chimaeras—is protected by fertilization. Chimaeras and many smaller sharks and
toothlike scales called dermal denticles. rays lay eggs, each protected by a tough egg capsule
and commonly known as a “mermaid’s purse.”
s e a rc h i n g f o r p r e y However, about sixty percent of cartilaginous fishes
The ocean is home to most cartilaginous fishes and give birth to well-developed live young that are
while many live on the seabed, the larger predatory nourished in the uterus by egg yolk or by a placental
shark species and plankton feeders roam in open connection to the mother. Unlike mammals, the
A white shark, Carcharodon carcharias,
water. The bullshark and over 100 other species young are independent from birth and neither shows the razor-sharp, replaceable rows of
can enter estuaries and swim up rivers and a few parent takes any interest in their offspring. teeth that make it such a fearsome predator.

CHIMAERAS SI X- A N D
Also known as rabbitfishes because of their 4 1/4 ft SEVEN-GILL
fused, platelike teeth, the Holocephali are PACIFIC
SPOOKFISH
1.3 m
SHARKS
a small class of cartilaginous fishes with one Rhinochimaera pacifica Most sharks have five pairs of gill slits, but those
order, Chimaeriformes, and about 56 species. f: Rhinochimaeridae
The long, conical snout of this chimaera in the order Hexanchiformes have either six or
A strong, venomous spine in front of the first is covered in sensory pores that detect seven pairs. All six known species live in deep
of two dorsal fins provides protection in their the electrical fields of its prey. water. Two of them, called frilled sharks, have
deepwater habitat. a long, soft, eellike body and have recently
SPOTTED 4 1/4 ft been placed in their own order,
large eye provides RATFISH 1.3 m Chlamydoselachiformes. 18 ft
better vision in Hydrolagus colliei 5.5 m
deep, dark water f: Chimaeridae
Like most chimaeras,
this N.E. Pacific
species uses its large ELEPHANT FISH CHIMAERA
pectoral fins to glide Callorhinchus milii
and flap along while f: Callorhinchidae
searching for food. Using its long fleshy snout as a plow, BLUNTNOSE SIXGILL SHARK
this chimaera unearths shellfish from Hexanchus griseus
muddy seabeds in southern Australia f: Hexanchidae
and New Zealand. Rocky seamounts around the world are
the haunt of this huge, green-eyed shark
that weighs up to 1,325 lb (600 kg).

6 1/2 ft
2m

RATFISH FRILLED SHARK

Chimaera monstrosa Chlamydoselachus anguineus
f: Chimaeridae f: Chlamydoselachidae
Typically living below 1,000 ft Scattered records indicate this shark
(300 m) in the Mediterranean and has a worldwide distribution. It has
pectoral fins flap E. Atlantic, ratfishes swim in small groups brilliant white teeth that may attract its
for swimming and search for seabed invertebrates. prey of fishes and squid.
 DOGFISH SHARKS CARPET SHARKS
328 A N D R E L AT I V E S The 46 or so sharks of the Orectolobiformes have two dorsal fins, one anal fin,
A large and varied order, the Squaliformes contains at least 143 species and and sensory barbels hanging down from the nostrils. With the exception of the
includes gulper, lantern, sleeper, and kitefin sharks. These all have two whale shark, they live quietly on the seabed, feeding on fishes and invertebrates.
FISHES • CARTILAGINOUS FISHES

dorsal (back) fins and no anal fin. All species so far studied bear live young.

white
spots mouth at
end of snout
5 ft
1.5 m PIKED DOGFISH
Squalus acanthias
f: Squalidae
Found worldwide in temperate waters, this once hugely
abundant shark is endangered by overfishing. Living up
to an age of 100, it grows and breeds very slowly.
22 in WHALE SHARK
56 cm Rhincodon typus
f: Rhincodontidae
40–65 ft
COOKIECUTTER This, the largest known fish, cruises tropical 12–20 m
SHARK 18 in oceans, feeding on plankton and tiny fishes.
Isistius brasiliensis 45 cm Each shark has its own unique pattern of spots.
f: Dalatiidae
This widespread, tropical
shark is an ectoparasite of
dolphins and large fishes. VELVET BELLY LANTERN SHARK 31/2 ft
Sucking on with thick lips, it Etmopterus spinax 1.1 m
twists around and bites out a f: Etmopteridae
round plug of flesh. This lantern shark lives in the deep E. Atlantic. It has tiny
light organs on its belly, which help in finding a mate.
EPAULETTE CATSHARK
8–14 ft Hemiscyllium ocellatum
2.4–4.3 m f: Hemiscylliidae
The bold eyespot mark of this long-tailed shark
may deter predators. It clambers among coral
in the South Pacific using its fins.
GREENLAND SHARK
Somniosus microcephalus TASSELLED
f: Somniosidae WOBBEGONG
One of only a few sharks that live in Arctic Eucrossorhinus dasypogon
waters, this huge, sluggish shark often f: Orectolobidae
scavenges for drowned land animals. With its fringe of skin
spines in sail-
rough skin like dorsal fins tassels, flattened body, and
camouflage pattern, this
S.W. Pacific coral reef
resident is hard to spot.
4 ft
1.2 m
5 ft “beard” of
1.5 m branching tassels NURSE SHARK
Ginglymostoma cirratum
ANGULAR ROUGHSHARK f: Ginglymostomatidae
Oxynotus centrina By day this shark lies hidden in
f: Oxynotidae rock crevices, emerging at night
Recently placed in its own order Echinorhiniformes, this to hunt in the warm coastal
deep-water shark has two saillike dorsal fins and rough 10 ft/3 m waters of the Atlantic and
skin. It lives in the E. Atlantic. the E. Pacific.

S AW S H A R K S BULLHEAD SHARKS
Sawsharks have a flat head with gills on the The Heterodontiformes are small bottom-living sharks
sides and a long snout (rostrum) with teeth with a blunt, sloping head, crushing teeth, and two
on the underside and edges. Two long sensory dorsal (back) fins each preceded by a sharp spine.
barbels hang from the rostrum and help They lay unique spiral-shaped eggs.
find buried food. Most of the nine species
spine
of Pristiophoriformes live in the tropics.
51/2 ft
1.7 m

LONGNOSE SAWSHARK
Pristiophorus cirratus 41/2 ft
f: Pristiophoridae 1.4 m PORT JACKSON SHARK
Sandy seabeds off southern Heterodontus portusjacksoni
Australia are home to this f: Heterodontidae
sawshark. It may use its rostrum to Using its paddlelike front fins, this shark crawls over the
deter predators as well as kill prey. seabed off southern Australia in search of sea urchins to eat.
 flattened head
and body
5 ft
1.5 m
SAND DEVIL
Squatina dumeril ANGELSHARKS
f: Squatinidae Flattened from top to bottom, angelsharks have gill slits on the sides
329
Sandy seabeds in the
N.W. Atlantic are home to of their large head. This distinguishes them from the similarly shaped
this well-camouflaged rays, which have gills underneath. The order Squatiniformes contains

FISHES • CARTILAGINOUS FISHES

predator. It strikes at passing just one family, which has 25 members. Angelsharks rear up from the
fishes with lighting speed. seabed on their pectoral fins to ambush prey.
gill
slit

GROU N D SH A R K S
There are about 295 species of Carcharhiniformes, making this
the largest and most diverse order of sharks. Most are large predators,
although the small yet numerous cat sharks are also included in the
order. All have two dorsal fins and one anal fin.

13 ft
4m BLUE SHARK
Prionace glauca
f: Carcharhinidae
Streamlined and elegant, this species travels
vast distances between feeding and breeding
grounds. More blue sharks are caught
than any other shark.

61/2 ft
2m

WHITETIP REEF SHARK

Triaenodon obesus
f: Carcharhinidae
This is perhaps the most common shark on the
coral reefs of the Indian and Pacific oceans. It
13 ft hunts fishes at night, sometimes in frenzied packs.
4m

OCEANIC sharksucker
WHITETIP
SHARK
Carcharhinus longimanus
f: Carcharhinidae
Heavily fished for its long,
white-tipped dorsal and
pectoral fins, this magnificent
oceanic predator is now 11 ft
critically endangered. 3.4 m

TIGER SHARK BULL SHARK

pelvic fin Galeocerdo cuvier Carcharhinus leucas
f: Carcharhinidae f: Carcharhinidae
Named for its stripes, which fade with One of the ocean’s most dangerous
25 ft age, the tiger shark will eat anything sharks, this top predator inhabits
7.5 m
and has attacked humans. Its serrated tropical coastal waters worldwide
teeth are shaped like a cockscomb. and also swims up rivers.
vertical dark
rear bars and spots broad, blunt
dorsal fin snout

anal fin
 GROU N D SH A R K S
330 white spots
on upperparts

3 1/4 ft
1m
SMALL-SPOTTED CATSHARK
Scyliorhinus canicula
41/2 ft f: Scyliorhinidae
STARRY SMOOTH-HOUND 1.4 m
Common in the N.E. Atlantic, this small shark is
Mustelus asterias one of more than 100 similar species in its family.
f: Triakidae Its egg cases are called mermaid’s purses.
This small, common shark of the N.E. Atlantic
has shiny skin and flat, slablike teeth, ideal
for crushing crabs and shellfish.

larger dorsal
portion of tail

hammer-shaped head

13 ft
4m SMOOTH HAMMERHEAD
Sphyrna zygaena
f: Sphyrnidae
With its eyes at both ends of a T-shaped
head, this shark has all-around vision. It
is found worldwide in warm water.

M AC K E R E L
SHARKS
The 16 species of shark in this order are
large with a cylindrical body and conical
head. The Lamniformes are (mostly)
13 ft 10 ft
fearsome hunters and many of them GOBLIN SHARK 3.9 m SAND TIGER SHARK 3.2 m
can maintain a high body temperature, Mitsukurina owstoni Carcharias taurus
allowing them to sustain speed in f: Mitsukurinidae f: Odontaspididae
cold waters. This shark has jaws that shoot forward to grab its prey. Living in The menacing, daggerlike teeth of this large
the dark depths of the Atlantic, Pacific, and W. Indian oceans, it warm-water shark belie its docile nature. It is
detects prey using its flat, electro-sensitive, bill-like snout. often kept in large public aquariums.

pointed
18 ft snout
5.5 m
SHORTFIN MAKO
Isurus oxyrinchus
MEGAMOUTH SHARK f: Lamnidae
Megachasma pelagios Capable of reaching 31 mph
f: Megachasmidae (50 kph), this may be the
Undiscovered until 1976, this giant shark is a filter fastest of all sharks. Its range 13 ft
feeder. It sucks in plankton-rich water and strains out is worldwide except for 4m
tiny shrimps and other plankton. The 100 or so polar waters, but it is
records suggest that it lives throughout the tropics. now endangered.

tall, triangular
powerful tail first dorsal fin conical snout

18 ft
5.5 m

WHITE SHARK
Carcharodon carcharias
f: Lamnidae THRESHER SHARK
One of the best-known top ocean Alopias vulpinus
predators, the white shark, or 24 ft large pectoral fin f: Alopiidae
the great white, is endangered. It 7.2 m Found almost worldwide, the thresher
travels long distances and ranges shark has a tail as long as its body, which it
through most oceans. uses to lash into fish shoals and stun its prey.
R AYS A N D S T I NG R AYS
Most rays and skates (Rajiformes) and stingrays (Myliobatiformes) live on the
tail as long
as body 331
seabed, athough they can swim by flapping their winglike pectoral fins. Their
flat, disk-shaped body is well-suited to this way of life. Stingrays have a long,

FISHES • CARTILAGINOUS FISHES

thin tail armed with a venomous spine for protection. Some stingrays spend
all or most of the time swimming and a few are found in fresh water.

41/2 ft pectoral
1.4 m
fin

brownish 91/2 ft
35 in 35 in 2.9 m
90 cm 90 cm upperparts

THORNBACK RAY BLUE SKATE

COMMON STINGRAY BLUE-SPOTTED Raja clavata Dipturus flossada
Dasyatis pastinaca RIBBONTAIL RAY f: Rajidae f: Rajidae
f: Dasyatidae Taeniura lymma One of the most common European One of two very similar, large European
While most stingrays live in f: Dasyatidae rays, this species has a row of skates, this species is still sometimes known
the tropics, the range of this Some individuals of this stingray, distinctive curved spines with wide as Dipturus batis. It is distinguished by its
fish is from the Mediterranean commonly found on coral reefs in the bases running down its back. long, pointed snout and blue underparts.
into N. Europe, where it lives Indian Ocean and the W. Pacific, have
inshore on sediment. two instead of one venomous tail spine.
HALLER’S ROUND RAY
Urobatis halleri
huge, pointed f: Urotrygonidae
GIANT MANTA RAY/REEF MANTA RAY “wings” At home on shallow, sandy flats, the
30 ft Mobula birostris/Mobula alfredi venomous spine on this stingray poses
9m f: Myliobatidae a threat to bathers. It is found in coastal
The largest of all rays, these two tropical species are waters from California to Panama.
filter-feeders and funnel plankton into the mouth,
using special flaps called cephalic horns. 23 in
58 cm

11 ft 13 ft
3.3 m 4m
cephalic
horn

SPOTTED EAGLE RAY SPINY BUTTERFLY RAY

Aetobatus narinari Gymnura altavela
f: Myliobatidae f: Gymnuridae
This tropical stingray noses out mollusks from This ray glides over the seabed in the warmer
the sand. It swims well by beating its pectoral parts of the Atlantic, searching for
fins like a bird’s wings. invertebrates and fishes.

S AW F I S H E S A N D ATLANTIC GUITARFISH
Pseudobatos lentiginosus E L E C T R IC R AYS
GU I TA R F I SH E S f: Rhinobatidae The pectoral fins of electric rays, order Torpediniformes,
A shovel-shaped snout allows this fish to dig out
Sawfishes and guitarfishes live and hunt on mollusks and crabs from the sand. Found mainly in have special organs that produce enough electricity to
and near the seabed. They have a flattened the Gulf of Mexico, it is now increasingly rare. stun prey and discourage predators. These
body, but swim with their tail. rays have a disklike body and
Sawfishes have a tough, bladelike snout, a powerful tail fin.
armed with regular-sized teeth on both
edges. Superficially they resemble 31/4 ft
sawsharks, but, like rays, they have gills on 1m
their underside. Recent research places the
30 in
seven species of sawfishes with guitarfishes 75 cm
in the order Rhinopristiformes.
SMALLTOOTH SAWFISH
Pristis pectinata
25 ft f: Pristidae
7.6 m Becoming more rare, this coastal warm-water
species uses its saw to slash shoals of fishes
and to stir up invertebrates from the seabed. MARBLED ELECTRIC RAY
Torpedo marmorata
f: Torpedinidae
When it pounces on a seabed fish, this electric ray can shock
its prey with up to 200 volts. Electricity can be produced by
even the newborn of this species.
332 BLUE-SPOTTED
R I B B O N TA I L R AY
Taen iu ra ly mma
FISHES • CARTILAGINOUS FISHES

size 28–35 in (70–90 cm), including tail

Stingrays are notorious for the painful wounds they can inflict with habitat Sandy patches in coral reefs
their barbed tail, which can, in exceptional circumstances, be lethal. distribution Indian Ocean, W. Pacific
However, the tropical blue-spotted ribbontail ray, like other stingrays, diet Mollusks, crabs, shrimp, worms
uses its sting just to defend itself. Much of its time is spent resting
motionless on sandy patches among coral, hidden under overhangs. MOUTH
The mouth is on
Often only its blue-edged tail gives it away to divers—if disturbed, it the underside of the
will swim away, flapping its two winglike pectoral fins. The best time stingray, allowing it
to see one is on a rising tide when the stingrays swim inshore to feed to extract mollusks
on invertebrates in shallow water. and crabs hidden
beneath the surface of
the sand. Two plates
of small teeth within
PELVIC FINS the mouth are used
In this female the to crush the shells
urogenital opening of its prey.
is visible between
the pelvic fins on the
underside. After mating,
females produce up
to seven live young
following a few months’
to a year’s gestation.
GILL SLITS
After passing over
the gills, water leaves
the body through five
pairs of gill slits on
the underside.

TOP HOLE BACK SPINES

Water is drawn in through two This species has
spiracles on top of the head behind relatively smooth skin,
the eyes, and leaves through but has two parallel
gill slits on the underside. rows of tiny spines
The elevated position of the running down its
spiracles helps prevent sand back, as well as other
from getting in. scattered spines.

ARMED TAIL
spiracle The tail is armed with one or two sharp, barbed
behind eye spines that cause physical wounds and inject
venom if the ray is attacked or stepped on.

barbed tail
 HIDDEN SPOTS
Unlike most of its relatives,
the blue-spotted ribbontail
ray rarely buries itself in
sand, relying instead on its
camouflage colors. Although
pectoral fin bright, its blue spots break up
the ray’s outline when seen
from above in the shifting
mouth sunlight of a shallow coral reef.

tail spine
THE FLIP SIDE
The mouth, nostrils, and gill slits
are all on the underside. The skin FRONT VIEW
is plain white with no blue spots, The eyes are raised up on top of
as this side is usually hidden. the head so that the ray can watch
for predators and prey, even when
it is partially buried in sand.
eye

pelvic fin
334 R AY- F I N N E D F I S H E S
These fishes are bony, with a hard, coral reefs to the waters beneath the ice-shelves of PHYLUM CHORDATA
calcified skeleton. Their fins are Antarctica, from the depths of the ocean to shallow
F I SH E S • R AY-F I N N ED F ISH E S

CLASS ACTINOPTERYGII

supported by a fan of jointed rods desert pools. Herbivores, carnivores, and scavengers ORDERS 47–59

called rays, made of bone or cartilage. are all represented in the group and its members FAMILIES 495–523

exhibit many ingenious hunting and defense SPECIES About 32,400

Ray-finned fishes are able to swim with much strategies, as well as cooperation between species.
greater precision than cartilaginous fishes. Using
their highly mobile and versatile fins, they can sa fet y i n n u m bers
execute maneuvers such as hovering, braking, and Most ray-finned fishes shed eggs and sperm into
even swimming backward. The fins themselves can the water and fertilization is external. In some
be delicate and flexible or strong and spiny and cases, fewer eggs are laid and parental care is
they often have important secondary uses, such provided. For example, jawfishes and some cichlids
as in defense, display, and camouflage. protect their eggs and young in their mouth, while
With the exception of many bottom-living sticklebacks and many wrasses build nests of weed
and deep-sea species, most ray-finned fishes have and debris. Some species protect their eggs with
a buoyancy aid in the form of a gas-filled swim such vigor that even scuba divers are warned off.
bladder. This allows them to adjust their buoyancy Most species, however, lay eggs in vast numbers.
at different depths by adding or removing gas to The millions of floating eggs and fish larvae are an
and from the bladder via the bloodstream. important food source for other aquatic creatures,
but those that survive drift and disperse the species.
m y r i a d a da p tat i o n s Populations of fishes reproducing in this way are less
The vast majority of fishes are ray-finned and the vulnerable to overfishing, as numbers can recover
group is hugely diverse—ranging from tiny gobies when fishing stops, but stocks—even of such
A pair of bluecheek butterflyfishes
to the gigantic ocean sunfishes. Species have evolved prolific breeders as Atlantic cod—will eventually (Chaetodon semilarvatus) swim in unison
to inhabit every conceivable aquatic niche from tropical succumb if intense levels of fishing are continued. as they patrol their patch of coral reef.

ST U RGEONS A N D R E L AT I V E S
Among the 28 members of the order Acipenseriformes, only the sturgeon AMERICAN PADDLEFISH
family includes marine species. The skull and some fin supports are Polyodon spathula
made of hard bone, but most of the remaining skeleton is made of pliable f: Polyodontidae
cartilage. As in sharks, which these fishes resemble, the tail is 6 ft Inhabiting N. American wetlands,
1.8 m this species has a long, paddlelike upper
asymmetrical with a longer upper lobe. jaw. It is one of the few freshwater
fishes that feed by straining plankton.
bony scutes
flattened,
bony head
EUROPEAN
STURGEON
Acipenser sturio
f: Acipenseridae
Heavily protected with rows
of bony plates, this critically 11 ft
endangered species, once prized for 3.5 m sensitive barbels
its caviar, inhabits coastal waters, help locate prey
but swims into rivers to breed.

GARS TA R PONS A N D
Fishes in the order Lepisosteiformes are primitive, TENPOUNDERS
freshwater predators from North America, with a long, The order Elopiformes is small and its members are 81/4 ft
cylindrical body protected by heavy, close-fitting scales. 2.5 m
silvery with only one dorsal fin and a forked tail. They TARPON
Their long jaws have needlelike teeth. resemble giant herring. They have special throat bones Megalops atlanticus
(gular plates). Though these are marine fishes, some f: Megalopidae
species also swim up estuaries and rivers. Found along Atlantic coasts, the tarpon
sometimes enters rivers. In stagnant
water, it gulps air from the surface using
LADYFISH 31/4 ft its swim bladder as a primitive lung.
6 ft LONGNOSE GAR Elops saurus 1m
1.8 m Lepisosteus osseus f: Elopidae
f: Lepisosteidae The ladyfish moves in large
A skilled predator, this long, thin fish hangs schools, close to W. Atlantic
motionless in the water, hidden by vegetation, shores. When alarmed, it skips
before thrusting forward to capture its prey. over the water’s surface.
BONYTONGUES
A N D R E L AT I V E S 335
As their name suggests, the fishes of the order
Osteoglossiformes have many sharp teeth on the 15 ft

F I SH E S • R AY-F I N N ED F ISH E S
tongue and roof of the mouth, which help them 4.5 m
ARAPAIMA gray to
to seize and hold prey. These fishes live in fresh dorsal fin set Arapaima gigas green body
water, mainly in the tropics. The group has many far back on body f: Arapaimidae
fishes with unusual shapes. Weighing up to 440 lb (200 kg), this S. American
species is one of the largest freshwater fishes. Its swim
bladder acts as a lung and it must gulp air regularly.
9 in ELEPHANTNOSE FISH CLOWN KNIFEFISH
23 cm Gnathonemus petersii Chitala chitala
f: Mormyridae 34 in
87 cm
f: Notopteridae
This African fish finds its way This slender, humpbacked
in its murky water habitat by fish lives in wetlands of
generating weak electrical S.E. Asia. In stagnant water, it
pulses. Its long lower jaw gulps air and absorbs oxygen
probes the mud for food. from its swim bladder.

EELS 5 ft
ZEBRA MORAY
Gymnomuraena zebra
The Anguilliformes have a long, thin, snakelike body with smooth skin, 1.5 m f: Muraenidae
This tropical, boldly
in which the scales are either absent or deeply embedded. The fins are long striped eel has dense
often limited to one long fin runing down the back, around the tail, and dorsal fin pebblelike teeth to eat
along the belly. Eels are found in marine and freshwater habitats. tough-shelled crabs,
mollusks, and sea urchins.

231/2 in
60 cm

spotted, JEWEL MORAY EEL large jaws

camouflaging Muraena lentiginosa
skin f: Muraenidae
Inhabiting coral reefs in the E. Pacific,
this eel rhythmically opens and closes
its mouth for respiration.
SPOTTED GARDEN EEL 41/4 ft
Heteroconger hassi RIBBON EEL 1.3 m
f: Congridae 16 in
CONGER EEL thick Rhinomuraena quaesita
40 cm
Sandy patches near coral reefs house colonies Conger conger body f: Muraenidae
or “gardens” of this eel. With their tails f: Congridae Juvenile ribbon eels are black with yellow
hidden in burrows, these eels sway like This eel of the North Atlantic fins and mature into bright blue males.
plants, retreating if disturbed. and Mediterranean finds an ideal These later change sex and become yellow
home in shipwrecks. It hides females. They are found in the Indian
in crevices during the day, and W. Pacific oceans.
emerging mostly at night
to hunt other fishes.
10 ft
3m

smooth EUROPEAN EEL 41/4 ft

BANDED SNAKE EEL 38 in skin Anguilla anguilla 1.3 m
Myrichthys colubrinus 97 cm f: Anguillidae
f: Ophichthidae Spending most of its life in fresh water,
Closely resembling a venomous sea snake, this harmless this endangered, snakelike eel migrates
eel of the Indian and W. Pacific oceans is left alone by across the Atlantic to the Sargasso Sea
predators. It searches in sand burrows for small fishes. to spawn and then die.

S WA L L OW E R S
AND GULPERS 31/4 ft
1m long, whiplike tail huge, loosely
hinged mouth
Living in the deep ocean, these bizarre, eellike fishes, PELICAN GULPER EEL
belonging to the order Saccopharyngiformes, have no tail Eurypharynx pelecanoides
fin or pelvic fins, and no scales. They do not have ribs and
f: Eurypharyngidae
This deep-sea, eellike fish has huge jaws and
their large jaws are modified for a wide gape. Like true an expandable stomach, which enable it to
eels, they are believed to spawn once and then die. swallow prey almost as large as itself.
 M I L K F I SH E S A N D R E L AT I V E S SA R DI N E S A N D R E L AT I V E S
336 With only two exceptions, including the milkfish itself, This predominantly marine order includes many commercially
the Gonorynchiformes live in fresh water. They have a pair important species. The Clupeiformes are silvery, with loose scales,
of pelvic fins that are set well back on the belly. one dorsal fin, a forked tail, and a keel-shaped belly. Most live in large
schools and are preyed on by sharks, tuna, and other large fishes.
streamlined body 8 in
20 cm
PERUVIAN ANCHOVETA
Engraulis ringens
f: Engraulidae
This tiny plankton-eater lives in enormous
shoals along the western coast of
S. America, where it is a major source of
food for humans, pelicans, and larger fishes.
18 in
MILKFISH 45 cm
Chanos chanos 6 ft
f: Chanidae 1.8 m
This fast swimmer has a large
forked tail. Feeding solely on pelvic fins
plankton, it is farmed BEAKED SALMON ALLIS SHAD 33 in ATLANTIC HERRING
in S.E. Asia. Gonorynchus grayi Alosa alosa 83 cm Clupea harengus
20 in f: Gonorynchidae f: Clupeidae f: Clupeidae
50 cm Native to Australia and New In spring, adults of this migratory Huge shoals of these silvery fishes swim
Zealand, this fish lives in species swim from the sea into in unison, eating planktonic copepods.
shallow bays and dives headfirst European rivers to spawn, often Highly prized, many N.E. Atlantic stocks
into the sand if threatened. swimming very long distances. are very heavily fished.

CA R P A N D R E L AT I V E S
This is one of the largest freshwater fish orders, with more than red-edged tigerlike
4,000 species worldwide. The Cypriniformes have the standard dorsal fin black stripes
“fish shape,” with a single dorsal fin. They typically have large
scales. The teeth are in the throat instead of the jaws. Many are
familiar aquarium fishes, including loaches,
minnows, and carp.

12 in
30 cm
deeply
forked tail

CLOWN LOACH TIGER BARB 23/4 in

Chromobotia macracanthus Puntigrus tetrazona 7 cm
f: Cobitidae f: Cyprinidae
Native to wetlands in S.E. Asia, this loach Native to the Indonesian islands of
is a bottom-feeder. It has sharp spines by Sumatra and Borneo, this fish has been
its eyes, which it uses to defend itself. widely introduced elsewhere and is
bred for the aquarium trade.

4 in
10 cm
BITTERLING 41/4 in 5 ft GRASS CARP GOLDFISH
11 cm 1.5 m
Rhodeus amarus Ctenopharyngodon idella Carassius auratus
f: Cyprinidae f: Cyprinidae f: Cyprinidae
This European fish lays its eggs A native of Asia, this species feeds on aquatic plants. Originally native to C. Asia and China, the
inside the mantle cavity of a For this reason, it has been introduced in Europe and goldfish has been introduced all over the
freshwater mussel, where they large, the US to keep drainage channels clear of weeds. world and there are now many varieties.
eventually hatch and the fry escape. silvery scales
COMMON CARP
Cyprinus carpio
f: Cyprinidae
4 ft protrusible With a mouth that
1.2 m mouth can be thrust out and
sensory barbels, a carp 21/4 in
finds food by grubbing 6 cm ZEBRAFISH
through the bottom Danio rerio
mud. Now introduced f: Cyprinidae
worldwide, it was This active little fish spawns frequently and
originally from China is common in ponds and lakes in S. Asia.
and C. Europe. It is bred in aquaria and laboratories.
C H A R AC I NS A N D R E L AT I V E S MEXICAN TETRA
Astyanax mexicanus
These freshwater fishes are mostly carnivores, with well-developed teeth. f: Characidae 337
While the normal form of the
As well as a normal dorsal fin, most also have a small, fatty adipose fin 43/4 in
Mexican tetra lives in streams
12 cm
near the tail. Of the 23 families in the order Characiformes, and can see well, this variant

F I SH E S • R AY-F I N N ED F ISH E S
the piranhas are the most notorious predators. lives in cave pools and is blind.

dark gray
head

adipose fin
13 in
33 cm

spots on body

RED PIRANHA
Pygocentrus nattereri
f: Serrasalmidae
Native to S. American rivers, this
fish usually eats invertebrates and
fishes, but schools of them, frenzied
red belly when by blood, can kill large mammals
fully grown with their razor-sharp teeth.

21/2 in
6.5 cm 16 in
40 cm
RIVER 31/4 ft
HATCHETFISH 1m
Gasteropelecus sternicla TIGERFISH LONGSNOUT DISTICHODUS
f: Gasteropelecidae Hydrocynus vittatus Distichodus lusosso
This insectivorous S. American f: Alestidae f: Distichodontidae
fish can leap out of the water Native to African rivers, this large Unlike some of its piranha relatives, this is
to escape predators and to predator has fanglike teeth and can a peaceful herbivorous fish. It lives in
catch aerial prey. eat fishes half as long as itself. streams in equatorial Africa.

CAT F I SH E S
The predominantly freshwater catfishes
have a long body and many whiskery
mouth barbels used to help find food. 121/2 in
32 cm
There is a sharp spine in front of the STRIPED EEL CATFISH 31/4 ft TIGER SHOVELNOSE
dorsal fin. Most species in this order Plotosus lineatus 1m CATFISH
f: Plotosidae Pseudoplatystoma fasciatum
(Siluriformes) grub around on the Juveniles of this tropical marine species form dense, f: Pimelodidae
bottom, searching for algae and ball-shaped shoals for protection. Solitary adults The long barbels of this S. American fish
small invertebrates. defend themselves with their venomous fin spines. help it find food as it hunts over riverbeds
at night searching out small fishes.

16 ft
5m SHEATFISH
Silurus glanis
f: Siluridae
From wetlands in C. Europe and Asia, this huge fish has
been known to weigh more than 660 lb (300 kg), but
due to heavy fishing, none this size exists today. long anal fin
sensory mouth
spine visible through barbels help
BROWN BULLHEAD transparent body 6 in GLASS CATFISH find food
15 cm
Ameiurus nebulosus Kryptopterus bicirrhis
201/2 in
f: Ictaluridae f: Siluridae
52 cm This N. American catfish has venomous A native of S.E. Asia, this small fish hangs
fin spines that help ward off predators motionless in the water; its transparent
when it is guarding its nest. body makes it hard for predators to see.
 SALMON AND adipose fin
SOCKEYE SALMON
Oncorhynchus nerka
338 R E L AT I V E S 33 in f: Salmonidae
84 cm In red breeding color, this
This order of marine and freshwater N. Pacific fish migrates into
fishes includes many anadromous N. American and Asian
F I SH E S • R AY-F I N N ED F ISH E S

members (moving from sea to fresh rivers to spawn and die. The
salmon are hunted by brown
water to breed). Powerful predators, bears while on their
the Salmoniformes have a large tail, difficult journey upriver.
a single dorsal fin, and a much
smaller adipose fin. hooked jaw on breeding male

221/2 in 31/4 ft 4 ft
CISCO 57 cm ARCTIC CHAR 1m 1.2 m RAINBOW TROUT
Coregonus artedi Salvelinus alpinus Oncorhynchus mykiss
f: Salmonidae f: Salmonidae f: Salmonidae
Widespread in N. American lakes and Clean, cold water is a must for this fish. Although native to N. America, this fish
large rivers, this fish forms shoals that Some individuals live in high-altitude lakes, has been introduced into fresh waters for
feed on plankton and invertebrates. while others migrate from sea to river. food and sport throughout the world.

PIKES AND single dorsal fin

R E L AT I V E S set far back
Found in cool, fresh waters across the northern
hemisphere, the Esociformes are fast and agile.
The dorsal and anal fins are set far back, near
the tail, to give these predatory fishes an instant
forward thrust.

61/2 in
17 cm

MUD MINNOW distinctive

Umbra krameri markings
f: Umbridae
The European mud minnow is now rare, as the
small ditches and canals of its home—the Danube
and Dniester river systems—disappear.

L I Z A R DF I SH E S A N D R E L AT I V E S LANTERNFISHES
This diverse order of marine fishes inhabits shallow coastal waters as well as the elongated A N D R E L AT I V E S
deep sea. The Aulopiformes have a large mouth, with many small teeth, and can pectoral fins
Lanternfishes are small, slim fishes with
catch large prey. They have a single dorsal fin and a much smaller adipose fin. many photophores (light organs) that help
triangular head
them communicate in their deep, dark
ocean habitat. Members of the
order Myctophiformes, these species
have large eyes and many migrate
toward the surface at night to feed.
16 in
40 cm
41/4 in
long pelvic fin VARIEGATED LIZARDFISH 11 cm
used as prop Synodus variegatus
f: Synodontidae
This resident of tropical reefs of the Indian and
16 in Pacific oceans perches on a coral head, remaining
40 cm completely still, and then darts out to catch fishes.

SPOTTED LANTERNFISH
BOMBAY DUCK FEELER FISH Myctophum punctatum
Harpadon nehereus Bathypterois longifilis 141/2 in f: Myctophidae
f: Synodontidae f: Ipnopidae 37 cm Living in the dark depths of the Atlantic,
During the monsoon season in the Indo-Pacific, Perching on its pelvic fins and tail this lanternfish uses its impressive array
schools of this small fish gather near river deltas above the deep, muddy ocean floor, this fish uses elongated of photophores for camouflage as
to feed on material that is washed down. its filamentlike pectoral fins to detect prey. pelvic fins well as communication.
DR AG ON FISH ES
A N D R E L AT I V E S large eyes 339
A majority of these deep-sea fishes have photophores to help row of
them hunt, hide, and find mates. Most members of the order photophores
Stomiiformes are fearsome-looking predators, with large
teeth and sometimes a long chin barbel. 2¾ in
7 cm
14 in
long, slender body 35 cm

SLOANE’S
91/2 in VIPERFISH
24 cm Chauliodus sloani
f: Stomiidae PACIFIC
NORTHERN STOPLIGHT LOOSEJAW Viperfishes have long, HATCHETFISH
Malacosteus niger transparent fangs that protrude Argyropelecus affinis
f: Stomiidae when the mouth is closed. Light f: Sternoptychidae
Found globally in temperate, tropical, and subtropical from photophores confuses The silvery, thin body of this fish helps to camouflage
oceans, this fish pinpoints its shrimp prey with a beam their prey in the depths of and hide it from predators. It is found in temperate,
of red bioluminescent light invisible to the shrimp. tropical and subtropical oceans. tropical, and subtropical waters.

duckbill-shaped KNIFEFISHES
snout
Knifefishes have a sideways flattened, knifelike body, and a single
long anal fin, which they use to glide backward and forward. The
electric eel is atypical, with a long, round body. The Gymnotiformes
live in fresh water and can produce electrical impulses.
231/2 in
60 cm

BANDED KNIFEFISH
CHAIN PICKEREL Gymnotus carapo
31/4 ft Esox niger ELECTRIC EEL 81/4 ft
f: Gymnotidae
99 cm f: Esocidae Electrophorus electricus 2.5 m Found in murky wetlands in
When hunting, this N. American f: Gymnotidae C. and S. America, this fish
pike uses delicate movements of This large S. American fish can produce an produces mild electric
its fins to hover motionless before electric shock of up to 600 volts—enough currents, which it uses to
striking with lightning speed. to kill other fishes and stun a human. sense its surroundings.

S M E LT S A N D R E L AT I V E S OA R F I SH E S A N D R E L AT I V E S
crest of
Smelts resemble small, slim salmon and, like them, most The 23 members of this marine order, called Lampriformes, are elongated
have an adipose fin on the back near the tail. Some of the colorful fishes of the open ocean, the adults having crimson fins. dorsal fin rays
Osmeriformes have a distinctive smell, like the European In many species, the rays of the dorsal fin extend as long streamers.
smelt, which smells like fresh cucumber. Most fishes in this order are ocean wanderers that are rarely seen.

18 in OARFISH
45 cm Regalecus glesne
f: Regalecidae
EUROPEAN SMELT The longest bony fish in the world, blue juvenile 36 ft
Osmerus eperlanus this huge fish has been the subject 11 m
f: Opisthoproctidae of many stories about sea serpents.
Resembling a cross between a herring and a trout, It is found in tropical to temperate
this fish is common in North Sea estuaries. In waters worldwide. pectoral fin
spring, smelt swim up rivers from the sea to spawn.

61/2 ft
2m

OPAH
Lampris guttatus
CAPELIN 10 in f: Lampridae
Mallotus villosus 25 cm This rarely seen fish beats
f: Osmeridae its long pectoral fins like
Found in cold Arctic and nearby waters, this small fish forms large wings. Inhabiting tropical
shoals that are a crucial food source for many seabirds; its abundance moderately to temperate oceans, it eats
or scarcity determines the breeding success of these birds. forked tail fin squid and small fishes.
 ANGLERFISHES COD AND
340 A N D R E L AT I V E S R E L AT I V E S
The 376 or so Lophiiformes include The Gadiformes include many important marine commercial species.
some of the most bizarre of all marine Most have two or three soft dorsal fins on their back and many have a
F I SH E S • R AY-F I N N ED F ISH E S

fishes. A modified fin ray on top of the chin barbel. Grenadiers live in deep water and have a long thin tail.
head acts as a fishing lure (bioluminescent 8 in
20 cm
in deep-water species), which attracts
prey toward the fish’s cavernous mouth. RED-LIPPED
BATFISH
Ogcocephalus darwini
9 in f: Ogcocephalidae
COFFINFISH 22 cm The oddly shaped
Chaunax endeavouri batfish scrambles over
f: Chaunacidae the seabed using
Coffinfishes lie on its paired pectoral and 61/2 ft
2m
muddy seabeds in the pelvic fins as props.
S.W. Pacific, waiting Its red lips are ATLANTIC COD chin barbel
for small fishes to a mystery. Gadus morhua
stray within reach. f: Gadidae
Overfishing has reduced the average weight of
ANGLER Atlantic cod to 24 lb (11 kg) from an historical
Lophius piscatorius maximum weight of over 200 lb (90 kg).
f: Lophiidae
61/2 ft A fringe of seaweed-shaped
2m flaps around its mouth helps
disguise this N.E. Atlantic
angler. It can strike
with lightning speed.

41/2 in 3 ft
11.5 cm 91 cm
ALASKA POLLOCK
Gadus chalcogrammus
8 in
f: Gadidae
20 cm FANFIN ANGLER The lack of a chin barbel and a protruding lower
Caulophryne jordani jaw help distinguish this fish from cod. It lives
f: Caulophrynidae in cold Arctic waters.
In the dark ocean depths it is
difficult to find a mate. Once
successful, the tiny male WARTY FROGFISH
fanfin angler latches onto Antennarius maculatus
the female permanently. f: Antennariidae
The well-camouflaged warty frogfish
clambers over coral reefs using 20 in
its limblike pectoral fins. 50 cm
SHORE ROCKLING
Gaidropsarus mediterraneus colorful
f: Lotidae tail fin with
Equipped with three sensory mouth barbels, rounded edge
this eellike rockling searches for food
in N.E. Atlantic rock pools.
8 in
20 cm

4 ft
SARGASSUMFISH 1.2 m
BURBOT
Histrio histrio Lota lota
f: Antennariidae f: Lotidae
While most frogfishes live Unlike all other Gadiformes, the burbot lives in
on the seabed, this one fresh water. It is found in deep lakes and rivers
hides among rafts of across the northern hemisphere.
floating Sargassum seaweed, 31/4 ft
which it closely resembles. 1m

PACIFIC GRENADIER
Coryphaenoides acrolepis
f: Macrouridae
skin flaps for A long, scaly tail and bulbous head
camouflage give this abundant deep-water
large pectoral fins cod relative its alternative name
used for scrambling of Pacific rattail.
CUSK EELS G R AY
Most members of the order Ophidiiformes live MULLETS 341
in the ocean and are long, slim, eellike fishes. They Gray mullets are
have thin pelvic fins and long dorsal and anal fins, silvery striped fishes
which in many species join onto the tail fin. with two widely
separated dorsal fins;
the first has sharp 30 in
8½ in
spines, the second has 75 cm
PEARLFISH 21 cm soft rays. Members of the GOLDEN GRAY MULLET
Carapus acus Mugiliformes are distributed Liza aurata
f: Carapidae f: Mugilidae
The adult pearlfish finds refuge inside a sea worldwide. They are vegetarian, Harbors, estuaries, and coastal waters in the
cucumber, entering tail-first through the anus, feeding on fine algae and detritus. N.E. Atlantic are all likely haunts of the this mullet,
exiting at night to feed. which often lives in shoals.

CORAL TOADFISH
Sanopus splendidus
15 in
f: Batrachoididae
38 cm The rare coral toadfish is found on reefs around
only one island off the coast of Mexico. It hides
under coral and in crevices.

mouth barbels

8 in
TOA DFISH ES 20 cm

A N D R E L AT I V E S
Toadfishes are broad, squat fishes with a
wide mouth and eyes on top of the head. They PLAINFIN
have two dorsal fins, the first short and spiny MIDSHIPMAN
and the second long and soft. Many species Porichthys notatus
f: Batrachoididae
of the Batrachoidiformes, especially the Living on the rocky shores along the west
midshipmen, are known for the remarkable way large pectoral fins
used for “walking” coast of N. America, this fish can breathe
they use their swim bladder to produce sound. air to survive when the tide goes out.

SI LV E R SI D E S NEEDLEFISHES
A N D R E L AT I V E S A N D R E L AT I V E S
These small, slim, silvery fishes often live in large shoals. There are more than With a long, thin, rodlike body and extended beaklike jaws, these
330 Atheriniformes, occuring both in marine and freshwater habitats. Most silvery fishes are well camouflaged in the open ocean. Flying fishes—with
have two dorsal fins, the first of which has flexible spines, and a single anal fin. their large, paired pectoral (side) and pelvic (belly) fins—also belong to
this order, the Beloniformes.
1½ in THREADFIN
4 cm RAINBOWFISH
Iriatherina werneri
f: Melanotaeniidae 3 ft
Mature males of this GARFISH
93 cm Belone belone
species diplay to females f: Belonidae
with their long fins. These
fishes live in S.E. Asia Remaining close to the ocean surface in the
and northern Australia N.E. Atlantic, this needlefish chases small fishes,
in weedy fresh waters. especially members of the herring family.
ATLANTIC
FLYINGFISH
CALIFORNIA GRUNION Cheilopogon heterurus
Leuresthes tenuis f: Exocoetidae
f: Atherinopsidae Pursued by predators or
Grunion spawn together in 7½ in startled by boats, this fish
thousands, depositing eggs in sand 19 cm escapes by powering up into
on the shore at night during the air and gliding away 16 in
high spring tides. on outspread fins. 40 cm
 KILLIFISHES SQUIRRELFISHES
342 A N D R E L AT I V E S A N D ROUGH I E S
Most members of the Cyprinodontiformes are small freshwater fishes These marine fishes are deep-bodied with large scales, a forked tail, and sharp
with a single dorsal fin and a large tail. Of the ten families, the guppies fin spines. Most of these fishes (Beryciformes and Trachichthyiformes) are
F I SH E S • R AY-F I N N ED F ISH E S

are perhaps the best known, as they produce live young and are popular nocturnal and are often a red color—a good disguise at night. Red light is
aquarium fishes. absorbed first by water and red fishes appear black below a certain depth.
23/4 in
7 cm CROWN
SQUIRRELFISH
Sargocentron diadema
f: Holocentridae
This is a typical
squirrelfish and one of
many similar species
found on tropical reefs. By
day it hides in crevices. 6 1/2 in
17 cm
AMIET’S LYRETAIL
tail important Fundulopanchax amieti 9 in
PINEAPPLEFISH
in display f: Nothobranchiidae 22 cm
Cleidopus gloriamaris
This small, colorful fish lives in rainforest streams f: Monocentridae
in Cameroon, Africa. There are many other Few predators would attack this spiny
similar species, known collectively as killifishes. fish, which has an armor of thick scales.
FOUREYED FISH Its color warns that it is unpalatable.
Anableps anableps
f: Anablepidae
The bulging eyes of this
S. American fish are divided
so that it can see clearly both 43/4 in
above and below water. 121/2 in 12 cm
32 cm EYELIGHT FISH
Photoblepharon palpebratum
f: Anomalopidae
AMERICAN FLAGFISH At night this fish signals to
Jordanella floridae others using a light organ
f: Cyprinodontidae under each eye. It has a black
Swamps and streams in Florida membrane that it uses to
are home to this peaceful cover the light, creating
vegetarian. The male courts a flashing effect.
a female and guards their eggs.
3 in COMMON FANGTOOTH
7.5 cm Anoplogaster cornuta
f: Anoplogastridae 7 in
The huge fangs of this deep-ocean 18 cm
BLACK PEARLFISH predator make escape impossible for
Austrolebias nigripinnis its prey, which it swallows whole.
f: Rivulidae
Subtropical rivers in S. America are
home to this species and all the other
members of its colorful family.
2 3/4 in
7 cm

30 in
2 in SAILFIN MOLLY 75 cm
5 cm Poecilia latipinna
f: Poeciliidae
The large dorsal fin of this ORANGE ROUGHY
N. American fish is displayed Hoplostethus atlanticus
in courtship. The female f: Trachichthyidae
bears live young. This is one of the longest-lived and
slowest-growing of any fish and can
live to at least 150 years old.

DORIES
AND
R E L AT I V E S protrusible
jaw
The 41 species belonging
to the order Zeiformes are
all marine, deep-bodied, but
thin fishes with long dorsal
and anal fins. Dories have JOHN DORY
protrusible jaws—they can Zeus faber
be shot out to capture prey. f: Zeidae
They feed on a wide range Viewed face on, this ultra-thin
fish is hard to see and can 3 ft
of small fishes that they stalk and strike its prey 90 cm
stalk head on. very effectively.
ST IC K L E BAC K S A N D SE A MOT H S CLINGFISHES
Most sticklebacks inhabit still and slow-moving fresh water, but some live in the sea, AND 343
as do seamoths. The Gasterosteiformes have a long, thin, stiff body protected by bony R E L AT I V E S
scutes (plates) along the sides and sharp spines along the back. Clingfishes are small, usually marine,

F I SH E S • R AY-F I N N ED F ISH E S
spines bottom-dwelling fishes. Most of the
Gobiesociformes possess a sucker formed
from modified pelvic fins with which they
4¼ in cling to rocks. Their eyes are set high on
11 cm the head and they have one dorsal fin.
3 ¼ in
8 cm

THREE-SPINED
STICKLEBACK
Gasterosteus aculeatus bony scutes males have red belly
f: Gasterosteidae when breeding CONNEMARA CLINGFISH
This little fish is widespread in SEAMOTH Lepadogaster candolii
fresh water and shallow seas in Eurypegasus draconis f: Gobiesocidae
the northern hemisphere. The f: Pegasidae Living in rocky shallows in the
male performs an elaborate Unlike the closely related sticklebacks, the N.E. Atlantic, this little fish
courtship dance. tropical marine seamoth is flat and has large 2 3/4 in is exposed to strong waves, but
7 cm can cling on tightly.
winglike pectoral (side) fins.

PIPEFISHES
AND SEAHORSES 61/2 in
16 cm
Seahorses and other similar fishes in the order
Syngnathiformes are encased in a body armor of tiny pectoral
bony plates, making their bodies very stiff. The group fin helps
maintain ROBUST GHOST PIPEFISH
includes both marine and freshwater species. Seahorses position Solenostomus cyanopterus
have a small mouth at the end of a tubular snout, which f: Solenostomidae
they use to feed on tiny planktonic crustaceans. Large pelvic fins enable the ghost pipefish to
drift and swim slowly among weeds and
seagrasses as it hunts minute invertebrates.

long,
tubular
snout
12 in
YELLOW 30 cm
SEAHORSE
Hippocampus kuda
18 in f: Syngnathidae
46 cm As in all seahorses, the
male of this species has
WEEDY a belly pouch in which
SEADRAGON he broods eggs laid
Phyllopteryx taeniolatus there by the female.
f: Syngnathidae 6 in
15 cm
This large, bizarrely shaped Australian seadragon
hides among seaweeds on rocky reefs, RAZORFISH
camouflaged by its many leaflike skin flaps. Aeoliscus strigatus
f: Centriscidae
Hanging head down
amongst urchin spines
disguises the razorfish,
which habitually swims in
RINGED PIPEFISH this position. This is an
Dunckerocampus dactyliophorus Indo-Pacific species.
f: Syngnathidae 7 in
18 cm
The long, thin body of this coral-reef
dweller is typical of pipefishes. It hovers
under and between corals and rocks.

TRUMPETFISH 32 in
Aulostomus chinensis 80 cm
skin flaps
provide f: Aulostomidae
camouflage Trumpetfishes often shadow moray
in seaweed eels, which hunt over coral reefs in
order to snap up small fishes that
the eels have flushed out.
 S WA M P E E L S A N D R E L AT I V E S TRIGGERFISHES, PUFFERFISHES,
344 These tropical and subtropical freshwater fishes have an eellike body A N D R E L AT I V E S
and most Synbranchiformes have tiny or no fins. This makes it easy for This diverse group of marine and freshwater fishes includes the huge
them to slither through swamps and marshes and burrow in mud. ocean sunfish and the poisonous pufferfish. Instead of normal teeth,
the Tetraodontiformes have fused tooth plates or just a few large teeth.
FIRE EEL Their scales are modified to form protective spines or plates.
Mastacembelus erythrotaenia
f: Mastacembelidae
31/4 ft Living in flooded lowland plains and
1m slow rivers in S.E. Asia, this edible spiny
eel feeds on insect larvae and worms. 20 in CLOWN
50 cm TRIGGERFISH
Balistoides conspicillum
MARBLED SWAMPEEL f: Balistidae
Synbranchus marmoratus This brightly colored coral-
f: Synbranchidae 5 ft
reef fish can wedge itself in a
Able to breathe air if necessary, this crevice by erecting its dorsal
1.5 m spines, which lock in position.
almost finless fish survives in very small
bodies of water in C. and S. America.

F L AT F I SH E S
Pleuronectiformes start life in the plankton as normal
upright juveniles. As they grow, they become flattened large tail
sideways and lie on one side on the seabed. The
underneath eye migrates around to join the other on
the upper side.
EUROPEAN PLAICE
Pleuronectes platessa
f: Pleuronectidae
This North Atlantic
commercial flatfish lies
camouflaged on the seabed
with its right side
uppermost. It emerges 10 in
1m at night to feed. 25 cm
3 1/4 ft

ATLANTIC HALIBUT 8 1/4 ft

Hippoglossus hippoglossus 2.5 m
f: Pleuronectidae
One of the largest flatfishes, the Atlantic SPOTTED
halibut lies on its left side with both BOXFISH
eyes on the upward-facing right side. Ostracion meleagris
f: Ostraciidae
Encased in a rigid box
of fused bony plates, and males have violet
with poisonous skin, this blue sides
Indo-Pacific reef fish is left
alone by predators.

28 in
COMMON 70 cm top is the left
SOLE side of the fish
Solea solea 6 in
f: Soleidae 15 cm
Although they can live to
be 30 years old, most sole WHITE-SPOTTED PUFFER
do not survive this long, Arothron hispidus
as they are a valuable f: Tetraodontidae
commercial fish. The neurotoxin contained within
the skin and organs of this fish
can easily kill a human and deters
natural predators.
20 in
right eye has 50 cm
moved to top
of the fish

TURBOT
Scophthalmus maximus
f: Scophthalmidae LONG-SPINE PORCUPINEFISH
The ability to alter its color Diodon holocanthus
to match the seabed helps f: Diodontidae
the turbot escape predators’ Found in all tropical seas, this spiny fish can suck in
attention. It is a valuable 31/4 ft water to inflate its body and turn itself into a prickly
North Atlantic species. 1m ball—a very effective deterrent against predators.
 41/4 in
11 cm SCORPIONFISHES
A N D R E L AT I V E S 345
Mostly bottom-living and marine, fishes in this
large order have a large spiny head with a unique

F I SH E S • R AY-F I N N ED F ISH E S
bony strut across the cheek. Most Scorpaeniformes
have sharp, sometimes venomous
spines in their dorsal fins and many
MIMIC FILEFISH are camouflage experts.
Paraluteres prionurus 14½ in
f: Monacanthidae 37 cm
The mimic filefish is avoided by
predators because it closely SMALLSCALED
SCORPIONFISH TUB GURNARD
resembles the saddled Scorpaena porcus Chelidonichthys lucerna
pufferfish, which has toxic flesh. f: Scorpaenidae juvenile f: Triglidae
Well concealed by skin flaps Walking over the seabed on three 30 in
flat back forms on the head and its ability to mobile rays on each of its 75 cm
top of bony change color, the scorpionfish pectoral fins, the tub gurnard can
protective box is hard to spot. probe for hidden invertebrates.

RED LIONFISH
18 in Pterois volitans
45 cm f: Scorpaenidae
The striped colors warn predators that even juveniles of
this coral-reef fish have venomous dorsal fin spines.
Adults darken with age and can be almost black.

8½ in
21 cm

23½ in
60 cm LUMPSUCKER BIG BAIKAL OILFISH
Cyclopterus lumpus Comephorus baikalensis
f: Cyclopteridae f: Comephoridae
A strong sucker on the belly allows this About a quarter of the body of this fish
rotund North Atlantic fish to cling onto is oil, which provides it with buoyancy.
wave-battered rocks and guard its eggs. It is endemic to Lake Baikal, Russia.

venomous dorsal
fin spines
small mouth has
stout teeth to
tear off sponges
large, upturned
mouth
OCEAN SUNFISH 10 in
Mola mola 25 cm
f: Molidae
Often lying on its side
at the surface, this fish
is nearly the heaviest
bony fish. The record is
held by the bump-head
sunfish M.alexandrini at
5,070 lb (2,300 kg). LONGSPINED BULLHEAD STONEFISH
Taurulus bubalis Synanceia verrucosa
f: Cottidae f: Synanceiidae
The color of this coastal It is extremely difficult to
and shore fish varies widely, spot this well camouflaged
according to its background. For tropical reef fish. A sting 16 in
example, fishes living among from its venomous spines 40 cm
red seaweeds are red. can be fatal to humans.
20 in
50 cm
7 in
18 cm

BULLHEAD
Cottus gobio
f: Cottidae
The bullhead lives
among stones and FLYING GURNARD
11ft vegetation in Dactylopterus volitans
3.3 m freshwater streams f: Dactylopteridae
and rivers throughout This species uses its huge fanlike
much of Europe. The pectoral fins to “fly” through the water. It
male guards the eggs. “takes off” from the seabed if disturbed.
346 RED LIONFISH
P t erois volit an s
The red lionfish is a night hunter, patrolling tropical coral size 18 in (45 cm) in length
habitat Coral and rock reefs
and rocky reefs in the western Pacific in search of small distribution Pacific Ocean,
fishes and crustaceans. It corrals its prey against the reef by introduced into W. Atlantic
spreading out its wide pectoral fins—one on each side of its diet Fishes and crustaceans

body—before gulping in the prey with lightning speed.

Sometimes it stalks its prey out in the open, relying on stealth
and a final quick rush, similar to a lion hunting on an African
plain. Protected by venomous spines, it will often face a diver
or potential predator head-on if approached. One male may VIBRANT WARNING
collect a small harem of females and will charge other males The contrasting striped pattern of
that stray too close. When ready to spawn, the male displays the red lionfish acts as a warning to
to the female, circling around her before they swim up toward predators that the fish is venomous
and they should keep away. On
the surface to shed eggs and sperm into the water. After a few land, stinging wasps use a similar
days the eggs hatch into planktonic larvae that drift in the method to avoid being eaten.
plankton for about a month before settling on the seabed.

MENACING PREDATOR eye is disguised

This efficient predator has been by a dark stripe
released from aquariums into the that confuses
Caribbean, where it is a threat to potential predators
native reef fishes. It hunts at dusk
using its large eyes and acute
sense of smell.
head
tentacle

VARIED STRIPES
The striped pattern differs
between individuals,
and becomes much less
obvious in breeding males,
which can be very dark.

PECTORAL FINS
The soft rays of the pectoral
fins are joined together
for part of their length by
a fine membrane, which
is marked with circular
splotches of color.
VENOMOUS SPINES
Sharp spines in this fish’s fleshy “whiskers”
dorsal, anal, and pelvic fins may help disguise
can be used to inject venom the fish’s large,
that causes extreme pain open mouth when
in humans, but is rarely approaching prey
fatal. The spines are
purely defensive.
TAIL UP
Lionfishes often hang slightly
head down and tail up, ready
to pounce on passing prey.
The tail helps the fishes to
maintain this position,
rather than to swim fast.
dorsal fin made up
of sharp spines
wide rays of pectoral
fins spread out fully
when the lionfish
ambushes prey

tail

anal fin
 PE RC H A N D
348 R E L AT I V E S
With at least 162 families and
35 in
over 11,500 species, the 90 cm
F I SH E S • R AY-F I N N ED F ISH E S

Perciformes form the largest and

most diverse vertebrate order.
Most have both spines and soft
rays in their dorsal and anal fins. 231/2 in 61/2 in
60 cm 16 cm
The pelvic fins are set well
forward, close to the pectoral LONGFIN SPADEFISH FOURSPOT BUTTERFLYFISH BLUE-SPOTTED SEABREAM
fins. New research means that Platax teira Chaetodon quadrimaculatus Pagrus caeruleostictus
some families included here f: Ephippidae f: Chaetodontidae f: Sparidae
are likely to be moved into Living on coral reefs in the Many different species of Found in the E. Atlantic, this is a typical
Indo-Pacific, this flattened species butterflyfishes are found on coral seabream with a steep head profile,
existing or new orders. patrols in small groups, feeding reefs around the world. This forked tail, and long dorsal fin.
on algae and invertebrates. fish lives in the W. Pacific.

43/4 in
12 cm 32 in
80 cm RED BANDFISH
Cepola macrophthalma
f: Cepolidae
Living in vertical mud
burrows in the N.E. Atlantic,
the red bandfish feeds on
passing plankton.
OCHER-STRIPED CARDINALFISH
Ostorhinchus compressus RED MULLET 16 in
f: Apogonidae Mullus surmuletus 40 cm
This coral-reef resident of the W. Pacific is a f: Mullidae
small, nocturnal species with two dorsal fins Common in the Mediterranean and N.E Atlantic,
and large eyes. The male incubates this species detects buried prey with its mobile
the eggs in his mouth. barbels. It is a relative of the tropical goatfish.

12 in 12 in
31 cm 30 cm
GREEN SUNFISH BANDED ARCHERFISH
Lepomis cyanellus Toxotes jaculatrix
f: Centrarchidae f: Toxotidae
Well known in N. America, this Living mainly in brackish mangrove estuaries
large species is one of the most in S.E. Asia, Australia, and the W. Pacific, this
common fishes in lakes and rivers. fish catches insects from overhanging branches
by shooting a jet of water from its mouth.

juvenile has brown

body with black-edged 4 ft
white spots 1.2 m
28 in
72 cm

BLUEFISH
Pomatomus saltatrix 20 in
f: Pomatomidae 51 cm
This widespread, voracious, and
aggressive predator roams tropical
and subtropical oceans in schools
that herd and attack smaller fishes.

HARLEQUIN SWEETLIPS 4 in
Plectorhinchus chaetodonoides 10 cm EUROPEAN PERCH
f: Haemulidae YELLOWHEAD JAWFISH Perca fluviatilis
The adult harlequin sweetlips Opistognathus aurifrons f: Percidae
is cream with black spots. f: Opistognathidae Native to Europe and Asia, this widespread,
The color and movement After the female of this Caribbean predatory freshwater fish has been introduced
of the juvenile shown here fish has laid her eggs, the male as a sport fish for anglers in Australia and
mimics a toxic flatworm. broods them in his mouth. elsewhere, where it has become a pest.
 16 in bold striped ROYAL ANGELFISH
40 cm pattern Pygoplites diacanthus
f: Pomacanthidae 349
The striking colors
of the royal angelfish
and of other coral reef
fishes help recognition
and communication
between species.
EMPEROR ANGELFISH
Pomacanthus imperator
f: Pomacanthidae 10 in
25 cm
This species lives on coral reefs in the Indo-Pacific.
Juveniles and adults have different patterns, which
protects juveniles from territorial aggression by adults.

single, long
dorsal fin

4 in 61/2 ft
10 cm 2m

10 in
25 cm

AMAZON LEAFFISH WRECKFISH SILVER MOONY

Monocirrhus polyacanthus Polyprion americanus Monodactylus argenteus
f: Polycentridae f: Polyprionidae f: Monodactylidae
With its large mouth and habit of Young wreckfishes live a nomadic life among This flattened Indo-Pacific
drifting like a dead leaf, this S. American surface-drifting debris, while the adults favor species is most at home in
freshwater predator can quickly shipwrecks, caves, and rocky areas. They are brackish-water estuaries,
engulf unsuspecting prey. found throughout the world’s oceans. where it swims in small shoals.

elongated, slender
head, rising
sharply at nape

6 in
15 cm 28 in
SEA GOLDIE 70 cm BROWN MEAGRE
Pseudanthias squamipinnis Sciaena umbra
f: Serranidae f: Sciaenidae
This small fish gathers plankton from Belonging to a family known as drums and
the waters near steep coral cliffs croakers, this fish of the N.E. Atlantic and
and ledges. The males defend Mediterranean communicates with loud
a harem of females. sounds generated by its swim bladder.

28 in 4 ft
70 cm 1.2 m

high
HUMPBACK GROUPER dorsal fin
Cromileptes altivelis
f: Serranidae
Like most other groupers, this increasingly JACK-KNIFEFISH
rare Indo-Pacific coral-reef species changes Equetus lanceolatus
sex from female to male as it matures. f: Sciaenidae
MALABAR GROUPER Deep coral reefs in the
Epinephelus malabaricus tropical W. Atlantic are
f: Serranidae home to this unusually
16 in 4 ft This Indo-Pacific species is found in shaped fish. Its shape and
40 cm 1.2 m a variety of habitats from reefs to color provide camouflage.
estuaries. Like many groupers, it
changes sex from female to male,
at about 10 years old.

COMMON BLUESTRIPE GRAY TILEFISH

SNAPPER Caulolatilus microps
Lutjanus kasmira f: Malacanthidae
f: Lutjanidae Living in mud and sand off the east
This fast-swimming snapper forms coast of N. America, this fish mostly
shoals in coral reefs during the day, remains within a depth of 655 ft 10 in
dispersing at night to feed. (200 m) to avoid very cold water. 25 cm
 PE RC H A N D stout tail with
350 R E L AT I V E S convex fin

8 in
forked tail 20 cm
SMALL SANDEEL
F I SH E S • R AY-F I N N ED F ISH E S

Ammodytes tobianus
5 ft 6 in f: Ammodytidae
1.5 m 15 cm This and other small sandeel
species are vital food source for
seabirds. It is found in large
shoals in shallow, sandy bays of
the N.E. Atlantic Ocean.
LESSER WEEVERFISH
WOLF-FISH Echiichthys vipera
Anarhichas lupus f: Trachinidae
f: Anarhichadidae This venomous N.E. Atlantic fish lies buried
A large mouth and strong teeth enable 16 in in the sand and can inflict a painful sting
this North Atlantic species to tackle 40 cm from its spiny, black, first dorsal fin.
hard-shelled invertebrates.
BLACK SWALLOWER
Chiasmodon niger
blue coloration f: Chiasmodontidae
in male Found in deep tropical and
subtropical waters, where food
is scarce, this fish, with its huge
mouth and distendable stomach, 10 in
25 cm
can tackle prey larger than itself.

CUCKOO
WRASSE
Labrus mixtus 10 in HARLEQUIN TUSKFISH CLEANER WRASSE 51/2 in
f: Labridae 25 cm Choerodon fasciatus Labroides dimidiatus 14 cm
Blue-marked males of the species f: Labridae f: Labridae
(shown here) develop from large, This member of the wrasse family This tiny wrasse forms a mutualistic
pink females with black and white tail from the W. Pacific uses its protruding relationship with other reef fishes. It cleans
markings, who change both color and caninelike teeth to move stones in and eats parasites off these fishes at specially
sex. This fish is found in the N.E. its search for invertebrate prey. set up “cleaner stations” on the reef.
Atlantic and Mediterranean.

71/4 ft
2.2 m DOLPHINFISH
15 in Coryphaena hippurus
ANTARCTIC TOOTHFISH MAORI COD 7 ft
Dissostichus mawsoni Paranotothenia magellanica 38 cm f: Coryphaenidae 2.1 m
f: Nototheniidae f: Nototheniidae This fast predator of the open ocean can
Like many Southern Ocean fishes, this species grows Found as far north as New Zealand and reach a speed of 37 mph (60 kph). It is
slowly, eventually reaching a large size. Its blood Argentina, this Southern Ocean species found worldwide in warm waters.
contains natural antifreeze, which helps is commercially important as food.
it to survive icy waters.

small mouth SHARKSUCKER

for picking out Echeneis naucrates 31/4 ft
invertebrates f: Echeneidae 1m
This species has a ridged sucker on top of
its head that allows it to cling onto large
fishes, dolphins, and turtles. It benefits
from scraps as its host feeds.

pattern
LONGNOSE HAWKFISH 5 in provides
Oxycirrhites typus 13 cm camouflage
f: Cirrhitidae
This Indo-Pacific species stays well
camouflaged among sea fans and black corals,
where it preys on small invertebrates.
BIGEYE TREVALLY
Caranx sexfasciatus
f: Carangidae
Built for speed, this predator
hunts other fishes at night over
reefs in the Indian and Pacific reddish coloration overall
4 ft oceans. Juveniles live inshore
1.2 m and may enter estuaries.
 white stripes
with thin 351
black outline

F I SH E S • R AY-F I N N ED F ISH E S
6 in BLUE CHROMIS 9 in SERGEANT MAJOR
15 cm Chromis cyanea 23 cm Abudefduf saxatilis
f: Pomacentridae f: Pomacentridae
Found in the tropical W. Atlantic, this The most common member of
is one of the most common fishes on the damselfish family, this brightly
coral reefs. It lays its eggs using striped, small fish can be easily spotted
an orange ovipositor tube. on coral reefs in the Atlantic Ocean.

single, long dorsal fin

CLOWN
41/4 in ANEMONEFISH
11 cm Amphiprion ocellaris
f: Pomacentridae
Protected by a special slime, 5 in
this colorful fish of the tropical juvenile
191/2 in 13 cm
W. Pacific finds a safe home 49 cm
21/2 in among the stinging tentacles NILE TILAPIA CHIPOKAE
6.5 cm of giant sea anemones. Oreochromis niloticus Melanochromis chipokae
f: Cichlidae f: Cichlidae
This common species of African This species is found only around
lakes is a locally important source the rocky shores of Lake Malawi in
of food. The female broods about Africa. Other fishes in this family
2,000 eggs in her mouth. are endemic to other lakes.

ORCHID DOTTYBACK 4 in 6 in
Pseudochromis fridmani 10 cm 15 cm
f: Pseudochromidae
One of the most brightly colored species of all FRESHWATER
coral-reef fishes, this Red Sea denizen hides ANGELFISH
beneath overhangs in steep areas. Pterophyllum scalare
f: Cichlidae
This popular aquarium
30 in fish, with its disklike and
75 cm CONVICT CICHLID
Amatitlania nigrofasciata laterally compressed body,
f: Cichlidae is native to S. American
Native to C. American rivers and streams, swamps. Both parents
this cichlid has been introduced elsewhere and guard the eggs and fry.
is a potential pest competing for food and space. elongated
pelvic fins

COMMON STARGAZER
Kathetostoma laeve 10 in GUNNEL
f: Uranoscopidae 25 cm Pholis gunnellus
Found around southern Australia, the common f: Pholidae
stargazer buries itself in sand with just its 28 in Found in rocky pools in the North Atlantic,
eyes and mouth exposed. 72 cm BLACKFIN ICEFISH this eellike fish with no scales can be very
Chaenocephalus aceratus slippery, sliding away easily from predators.
f: Channichthyidae
grayish A natural antifreeze in the blood hump helps to
saddlelike of this Southern Ocean fish enables break up coral
marking it to survive at temperatures as
low as 28.40F (-20C).
20 in 41/4 ft
50 cm 1.3 m

MEDITERRANEAN
PARROTFISH GREEN HUMPHEAD
Sparisoma cretense PARROTFISH
f: Scaridae Bolbometopon muricatum
This Mediterranean species is f: Scaridae
the only parrotfish in which the A tough, beaklike mouth allows this
beaklike female is more colorful than huge Indo-Pacific coral reef resident
teeth the male. Most other parrotfish to crunch up living coral, expelling
species are tropical. the remains as sand.
 PE RC H A N D R E L AT I V E S
352 8 in
20 cm
F I SH E S • R AY-F I N N ED F ISH E S

FIREFISH 31/2 in
Nemateleotris magnifica 9 cm
f: Microdesmidae
43/4 in Hovering above its coral reef burrow, this
12 cm fish gathers plankton from the water and
SPOTTED CTENOPOMA darts back into safety if it senses danger.
Ctenopoma acutirostre
f: Anabantidae elongated first
ray on dorsal fin
This tropical freshwater fish is found in BUTTERFLY BLENNY
the Congo basin of Africa. It often stalks Blennius ocellaris
its prey with its head held down. f: Blenniidae
Like many of its relatives, this fish of
the N.E. Atlantic lives on the seabed
21/2 in and guards its eggs, which are often
6.5 cm
laid in an empty shell.

broad tail fin

SIAMESE FIGHTING FISH

Betta splendens
f: Osphronemidae
The original range of this Asian freshwater fish
is unclear, but it has been bred for centuries for
its fighting ability, particularly strong in males.

21/4 in
6 cm

9 in
23 cm
MANDARINFISH
POWDERBLUE SURGEONFISH Synchiropus splendidus
Acanthurus leucosternon f: Callionymidae
f: Acanthuridae huge, saillike Native to the Pacific, this is
This Indian Ocean fish has a sharp, dorsal fin 10 ft ATLANTIC SAILFISH one of the most colorful of all
bladelike structure sheathed on either side 3.2 m Istiophorus albicans tropical reef fishes. Its vivid colors
of its tail base, which it can flick out f: Istiophoridae warn predators of its foul taste.
to slash a would-be attacker. This ocean predator uses its upper torpedo- projecting
jaw, which extends as a long spear, shaped body lower jaw
to slash into shoals of fishes
and stun them.

spear-shaped 61/2 ft
snout GREAT BARRACUDA 2m
Sphyraena barracuda
f: Sphyraenidae
Found globally in tropical and subtropical
waters, this solitary predator stalks its prey
and then accelerates rapidly as it strikes.
231/2 in
60 cm

11/2 in 31/2 in
4 cm 9 cm
15 ft ATLANTIC MACKEREL BUMBLEBEE FISH PAINTED GOBY
4.5 m Scomber scombrus Brachygobius doriae Pomatoschistus pictus
f: Scombridae f: Gobiidae f: Gobiidae
Living in large shoals in the North Found in S.E. Asia, this This and other similar gobies are common in
Atlantic, this fish feeds voraciously on bottom-dwelling goby is tolerant shallow sediment areas of the N.E. Atlantic. DNA
small fishes and plankton. Its streamlined of brackish water and lives in analysis suggests gobies might be placed in a
body makes it a fast swimmer. estuaries and mangroves. high-set, separate order, Gobiiformes.
NORTHERN BLUEFIN TUNA bulbous eyes ATLANTIC MUDSKIPPER
Thunnus thynnus Periophthalmus barbarus
f: Scombridae long tail
10 in f: Gobiidae
Found globally, this tuna is one of the 25 cm As long as it stays moist, this
world’s most valuable commercial mudskipper can stay out of
fishes. A fast, dynamic predator, it water for hours by absorbing
roams widely, hunting small fishes. oxygen through its skin.
LOBE-FINNED FISHES 353
Once considered to be the ancestors rocky areas in the western Indian Ocean. A second PHYLUM CHORDATA

of land vertebrates, lobe-finned fishes species was found in Indonesian waters in 1998.

FISHES • LOBE-FINNED FISHES

CLASSES COELACANTHI

have fins that resemble primitive They have several unique structural features DIPNEUSTI
ORDERS 3
limbs, with a fleshy base preceding including an incomplete backbone, residual
FAMILIES 4
notochord, and a tail fin with a characteristic
the fin membrane. extra middle lobe. Their scales are heavy, bony
SPECIES 8

Like the ray-finned fishes, these are bony fishes with a plates and they are not long-distance swimmers.
hard skeleton but their fins have a different structure. Unlike the egg-laying lungfishes, coelacanths bear
The fin membrane is supported on a muscular lobe live young as the eggs hatch internally. Gestation D E B AT E
that projects out from the body and is strong enough may be as long as three years, and this makes their FISHES ON LAND
to let some of these fishes shuffle along on their paired survival precarious if individuals continue to be
While it is generally accepted that
pectoral and pelvic fins. Bones and cartilage inside the caught in the nets of deep-sea trawlers.
land-living vertebrates evolved from
lobes provide attachment for muscles. There are many a primitive fish or fishlike ancestor in
fossil groups of lobe-finned fishes, but living b r e at h i n g o u t o f wat e r the sea, finding that ancestor is more
representatives are confined to the marine Although most of their fossil ancestors lived in the difficult. Recent work suggests that
coelacanths and the freshwater lungfishes. ocean, modern lungfishes are limited to freshwater lungfishes are more closely related to
habitats in South America, Africa, and Australia. tetrapods (four-limbed vertebrates such
a living fossil All can, to some extent, breathe air via a connection as mammals) than are coelacanths.
Coelacanths are nocturnal and to the swim bladder—useful when pools dry out In 2002 a fossil lobe-finned fish,
seasonally. Some species can survive buried in mud Styloichthys, was found in China. It
secretive. The first living appears to show close links between
specimen was discovered in for many months and would die if kept permanently
lungfishes and tetrapods. The debate
1938—until then only fossil submerged in water, while others still rely mainly on continues, but currently coelacanths
species over 65 million years old gills for breathing. Their shape and the fact that the are not considered direct ancestors
were known. The historic find larvae of some species have external gills led early of tetrapods.
belonged to a species living in deep zoologists to believe that lungfishes were amphibians.

AFRICAN LUNGFISHES COE L ACA N T H S

triple-lobed tail fin
All four species of African lungfishes have a long body and The two primitive fishes from 41/2 ft
threadlike pectoral and pelvic fins, collectively called paired fins. the order Coelacanthiformes 1.4 m
The Lepidosireniformes breathe through a pair of lungs derived have limblike, fleshy bases to their
from their swim bladder. pectoral and pelvic fins, and large
bony scales. In life, these fishes are a
metallic blue with pale spots, but the
color fades on death. Interesting
61/2 ft
‘head-standing’ behavior has been
2m WEST AFRICAN LUNGFISH observed from a submersible. INDONESIAN COELACANTH
Protopterus annectens Latimeria menadoensis
f: Protopteridae f: Latimeriidae
When the lakes in which it lives dry out, this Molecular studies have shown this to be
body flecked a separate species from the coelacanth,
lungfish survives by burying itself in mud, with white
forming a cocoon with an inlet for air. speckles although they are physically similar.
It lives in the Celebes Sea.

AU ST R A L I A N
LUNGFISHES
The single species in the order Ceratodontiformes
has a long body, large scales, paddlelike paired fins,
and a tapering tail. It can breathe through its lungs
for short periods, but cannot survive if its habitat
dries out.

COELACANTH
AUSTRALIAN LUNGFISH 61/2 ft Latimeria chalumnae
Neoceratodus forsteri 6 ft 2m f: Latimeriidae
f: Neoceratodontidae 1.8 m This fish lives off the coasts of southern
Living in deep pools and rivers, this Africa and Madagascar in steep, rocky,
lungfish can survive in stagnant water underwater terrain and hides in deep
by gulping air into its swim bladder. limblike fins fin with fleshy stalk ocean caves during the day.
354 A M PHIBIANS
Amphibians are cold-blooded vertebrates that thrive in freshwater
ANIMALS • AMPHIBIANS

habitats. Some spend their whole lives in water; others only require
water to breed. On land, they must find damp places, as their skin
is permeable and does not protect them from drying out.

PHYLUM CHORDATA The three orders of living amphibians are thought

CLASS AMPHIBIA to share a common ancestor, but their origins are
ORDERS 3 still uncertain. There is an enormous gap in the fossil
FAMILIES 74
SPECIES 8,212
record between the first land animals to evolve from
fishes—the tetrapods, some 375 million years ago—
and a froglike creature that lived 230 million years ago.
Amphibians have a unique and complex life cycle
and occupy very different ecological niches at different
stages of their lives. For most amphibians, their eggs
hatch into larvae—called tadpoles in frogs and toads—
that live in water, often in very high densities, and feed
mostly on algae and other plant material. As larvae,
they grow rapidly and then undergo a complete change
Large numbers of male and female of form—metamorphosis—to become land-living
Costa Rican golden toads gather in a adults. As adults, all amphibians are carnivores, most
pool to mate. The species is now extinct.
feeding on insects and other small invertebrates.
They typically lead secretive and solitary lives, except
when they return to ponds and streams to breed.
Most amphibians thus require two very different
habitats during their lives: water and land. During
metamorphosis, they undergo a wide variety of
anatomical and physiological changes—from aquatic
creatures that swim with a tail and breathe through
A female of the North American gills, to terrestrial animals that move with four limbs
Jefferson’s salamander attaches a mass of
eggs to a submerged twig in early spring. and breathe through lungs.

d i v e r s e pa r e n ta l c a r e
Some amphibians produce huge numbers of eggs that
are left to fend for themselves, so that only a very few
will survive. Many others have evolved various forms
of parental care. This is generally associated with the
production of far fewer offspring, so that reproductive
success is achieved by caring for a manageable number
When the eggs hatch, the male
red-backed poison frog carries the of young, rather than by producing as many eggs as
tadpoles to bromeliad pools in the canopy. possible. Parental care takes many forms, including
defending eggs or larvae against predators, feeding
D E B AT E tadpoles with unfertilized eggs, and carrying tadpoles
OUR BIGGEST CHALLENGE? from one place to another. In some species, such as
the midwife toads and some poison frogs, parental
A third of the world’s amphibians face
extinction in the near future and this
care is the duty of the father. In many salamanders
poses a huge conservation challenge. and caecilians, the mother is the sole protector of
This danger of extinction is largely due the young. In a few frog species, mother and father
to the destruction and pollution of establish a durable pair-bond and share parental duties.
freshwater habitats, but amphibians are
also threatened by the global spread of
the disease chytridiomycosis, caused by READY TO HATCH
a fungus that invades their soft skin. Tadpoles of the Tanzanian Mitchell’s reed frog wriggle
within their egg membranes, just prior to hatching.
a m p h i b i a n g ro u p s
The three orders have very different
body forms: the tailless frogs
and toads, the caecilians, which
resemble earthworms, and the
lizard-like salamanders and newts.
f ro g s a n d t oa d s

356
ca e c i l i a n s

369
sa la m an der s
and newts

370
356 F RO G S A N D T OA D S
With powerful hind limbs tucked Not all anurans jump, however. Many have hind PHYLUM CHORDATA
under its body, a wide mouth, and limbs adapted for other kinds of locomotion,
AMPHIBIANS • FROGS AND TOADS

CLASS AMPHIBIA

protruding eyes, the typical frog has a including swimming, burrowing, climbing, and, in ORDER ANURA

unique and unmistakable body shape. a few species, gliding through the air. Most frogs FAMILIES 54

and toads live in damp habitats, close to the pools SPECIES 7,244

Anura, the name of the order of frogs and toads, and streams in which they breed, but there are
means “animals without a tail.” As adults, all other several species adapted to life in very arid areas. D E B AT E
amphibians have a tail, but in the anurans, the tail The greatest diversity of anurans is found in the FROG OR TOAD
gradually disappears during metamorphosis from tropics, particularly in rainforests. Many are active
the larval to the adult stage of the life cycle. The by day, while others are nocturnal. Some species The distinction between frogs and
larvae, known as tadpoles, feed chiefly on plant are cleverly camouflaged; in contrast, others are “toads” is biologically meaningless and
the two words are used differently in
material and have a spherical body that contains brightly colored, advertising that they are poisonous different parts of the world. In Europe
the long, coiled gut required by such a diet. The or unpleasant to taste. and North America, for example, the
adults, in contrast, are totally carnivorous, feeding term “toad” refers to species in the
on a wide range of insects and other invertebrates, c o u rt s h i p a n d b r e e d i n g family Bufonidae, but this also includes
with larger species also taking small reptiles and Anurans differ from other amphibians in having a the South and Central American
mammals, as well as other frogs. voice and very good hearing. Males of most species harlequin frogs. In general, toads have
call to attract females, making sounds characteristic rough skins, are slow-moving, and
i n g e n i o u s a da p tat i o n s of their species. In all but a very few species, often burrow in the ground, whereas
Frogs and toads catch their prey by ambush, most fertilization is external, the male shedding sperm frogs are smooth-skinned, agile, and
fast-moving, and spend much of their
spectacularly by jumping. In many frogs and toads, onto the eggs as they emerge from the female’s
time in water. A group of smooth-
the hind limbs are modified for jumping, being body. To do this, the male clasps the female from skinned, aquatic frogs native to Africa
much longer than the forelimbs and very muscular. above—a position known as amplexus. The duration were once known as clawed toads,
Jumping is also an effective method of escaping of amplexus varies between species—from a but are now called clawed frogs.
from predators, of which frogs and toads have many. few minutes to several days.

A LY T I DA E SQUEAKERS ¾–1¼ in
Males of the Alytidae, a small family of terrestrial frogs, A N D R E L AT I V E S 2–3 cm

call at night to attract females. When mating, the male The Arthroleptidae are a large, diverse family of
midwife toad attaches fertilized eggs to his back. He frogs occurring across sub-Saharan Africa—in
carries them until they are ready to hatch, when he forest, woodland, and grassland, some at high
releases the tadpoles into water. Occasionally, a male altitudes. Its members range from tiny
may carry egg strings from more than one female. With “squeakers,” named for their high-pitched
painted frogs, a female mates with several males, producing call, to large treefrogs.
up to 1,000 eggs, which are dropped into water.
eggs attached vertical,
to male’s back slitlike pupil very long
third finger

WEST AFRICAN
RIO BENITO SCREECHING FROG
LONG-FINGERED Arthroleptis
FROG poecilonotus
Cardioglossa gracilis The female of this
An inhabitant of lowland small species lays large
forest, this frog breeds eggs in soil cavities.
1¼–1½ in in streams. Males call The male is noted
3–4 cm from nearby slopes. for his loud call.

1–1½ in
2.5–4 cm

1¼–2 in 1½–2¼ in
3–5 cm 4–5.5 cm
WEST CAMEROON
COMMON MIDWIFE TOAD FOREST TREEFROG AFRICAN TREEFROG
Alytes obstetricans Leptopelis nordequatorialis Leptopelis modestus
Found in W. and C. Europe, this midwife toad has This large treefrog lives in montane grasslands This species is found near streams
a plump body and powerful forelimbs adapted for of W. Africa. Males call to females near water when in forests of W. and C. Africa.
digging. It hides in a burrow by day. breeding, and eggs are deposited in ponds or marshes. Females are larger than males.
GL A SS F ROGS silver eyes with
black reticulations
Found in Central and South America, frogs of
357
the family Centrolenidae are known as “glass LIMON GIANT GLASS FROG
Sachatamia ilex
frogs” because many species have transparent This arboreal frog lives in wet

AMPHIBIANS • FROGS AND TOADS

skin on the underside through which their vegetation near streams. It has dark
internal organs are visible. green bones, visible through its skin.

¾–1¼ in adhesive pad

2–3 cm at tip of digits
1–1½ in
2.5–3.5 cm

WHITE-SPOTTED ¾–1¼ in
COCHRAN FROG 2–3 cm
Sachatamia albomaculata
FLEISCHMANN’S GLASS FROG This species is found in EMERALD GLASS FROG
Hyalinobatrachium fleischmanni wet lowland forests, and Espadarana prosoblepon
Males of this species are territorial, using calls breeds near streams. Males Males of this arboreal frog are fiercely territorial,
to defend their territory and to attract females. ¾–1¼ in call to females from low defending their space by calling, and occasionally
The females lay eggs on leaves over water. 2–3 cm vegetation nearby. fighting rivals while hanging upside down.

C E R AT O P H RY I DA E ORNATE HORNED 3¼–5 in

FROG 8–13 cm
These South American horned frogs have a very large head “horn” above eye Ceratophrys ornata
and wide mouth that enable them to eat animals nearly as A voracious predator living
large as themselves. They are “sit-and-wait” predators, in grasslands in Argentina,
remaining still and well-camouflaged before their prey this species breeds after
heavy rain, in temporary
comes within range. pools and ditches.

CRANWELL’S HORNED FROG

Ceratophrys cranwelli
Spending much of its life underground, this
large frog emerges after heavy rain to mate
and lay its eggs in pools.
3½–5½ in
9–14 cm
1½–4 in
4–10 cm

wide
mouth

BUDGETT’S FROG
Lepidobatrachus laevis
This frog has a flattened body, a wide mouth,
and fangs. It spends dry periods in a cocoon
below ground, emerging to breed after rain.

ROBBE R F ROGS ¾–2 in

2–5 cm
Found in North, Central, and South America, the Craugastoridae
produce eggs that develop directly into small adults, without
a tadpole stage. Eggs may be deposited on the ground or in
vegetation. Many species show parental care of the eggs.

BROAD-HEADED
RAIN FROG 1–2¼ in
Craugastor megacephalus 2.5–5.5 cm
This C. American frog hides in a
burrow by day and emerges at night.
It lays its eggs in leaf litter. ISLA BONITA FITZINGER’S ROBBER FROG
ROBBER FROG Craugastor fitzingeri
Craugastor crassidigitus In this forest species, the male calls to
Native to the humid forests of C. America, the larger female from an elevated perch.
1¼–2¾ in this ground-living frog is also found in coffee Eggs are laid in the ground and are
3–7 cm plantations and pastures. guarded by the female.
 T RU E TOA DS COMMON AFRICAN TOAD
Sclerophorus regularis
358 Distributed worldwide, the family Bufonidae is Common throughout Africa except the
driest parts of Sahara and Namib deserts,
large and diverse. Its members are characterized by horizontal this thickset toad breeds in dams and
shortened forelimbs, hind limbs that are used for pupil ponds. Males produce rasping, ducklike
walking or hopping, dry warty skin, and parotoid calls to attract females.
glands behind the eyes. However, this family also
includes the more slender and long-limbed South
and Central American harlequin frogs and the
2–4½ in
stubfoot toads. 5–11.5 cm

2–4 in
green blotches 5–10 cm
on back
MALAYAN
TREE TOAD
Rentapia hosii
Found in E. Asia, this toad
warty is unusual in living a largely
skin arboreal life. It has adhesive
pads on its toes that
enable it to climb trees.

3½–4¾ in
9–12 cm

GREEN TOAD
Bufotes viridis
A native of sandy habitats,
this colorful toad is found 2–4 in
in Europe and W. Asia. 5–10 cm
It emerges from its
burrow in spring to NATTERJACK
breed in ponds. Epidalea calamita
Compared with other true
toads, this species has
short legs and runs like a
mouse. Occurring across
Europe, it breeds from
spring to summer.

C E R ATOBAT R AC H I DA E BOM BI NATOR I DA E

Found in Southeast Asia, China, and several Pacific These small aquatic toads are found in Europe and
islands, frogs of this family produce large eggs that Asia. They have a flattened body and many are brightly prominent
hatch directly into small frogs. In many species, the colored. Fire-bellied toads are active by day, but the eyes
tips of the fingers and toes are enlarged. dull-colored barbourulas from the Philippines and
Borneo are nocturnal.

FIJI GROUND FROG

Cornufer vitianus
bright red
The populations of this species underside
on several of the Fijian Islands
have been wiped out following
the introduction of mongooses.
1–4¼ in
2.5–11 cm
hornlike projection
above eye ORIENTAL
FIRE-BELLIED TOAD
Bombina orientalis
flat, Found in China and Korea, this small,
triangular bright green 1¼–2 in squat frog can produce a toxic skin secretion.
head coloration 3–5 cm If attacked, it displays its bright belly colors.

2–3¼ in
5–8 cm
R A I N F ROGS 1¼–2 in
3–5 cm

SOLOMON ISLANDS
Members of the family DESERT RAIN FROG
HORNED FROG Brevicipitidae are found in Breviceps macrops
Cornufer guentheri eastern and southern Africa. Living away from
This species has a During mating, the much standing water, this
pointed snout and smaller male is glued onto burrowing frog lives and
hornlike projections breeds among Namibian
above its eyes. It hides a female’s back by a special sand dunes, occasionally
among dead leaves. skin secretion. moistened by sea fog.
 2–3½ in VARIABLE HARLEQUIN FROG
5–9 cm Atelopus varius
This aggressive species from Panama 359
AMERICAN TOAD and Costa Rica has vivid and variable
Anaxyrus americanus coloration. It lives near streams and
Found in eastern is active during the day.

AMPHIBIANS • FROGS AND TOADS

N. America, this toad is
quite variable in color. 1–21/4 in
large Breeding occurs in ponds 2.5–6 cm
parotoid where males produce
glands long trilling calls.

1½–31/4 in 1–1½ in 2–4 in

short, 4–8 cm 2.5–4 cm 5–10 cm
warty legs

TRUANDO TOAD
Rhaebo haematiticus GUYANAN STUBFOOT TOAD PANAMANIAN GOLDEN FROG
Living among leaf litter in Atelopus barbotini Atelopus zeteki
forests of C. and S. America, this This small toad from Guyana has a This brightly colored frog from Panama
broad-headed toad lays its eggs flattened body. It breeds throughout breeds in pools after heavy rain. It may
in long strings in rocky pools. the year in forest streams. now be extinct in the wild.

parotoid glands
secrete toxin
olive-
21/4–33/4 in brown,
5.5–9.5 cm warty skin

4–9½ in
10–24 cm
31/4–8 in
8–20 cm CANE TOAD
GREEN Rhinella marina
EUROPEAN COMMON TOAD CLIMBING FROG One of the world’s
Bufo bufo Incilius coniferus largest toads, this
Found throughout Europe and Native to C. and S. America, American species was
N. Africa, this species breeds in spring, this nocturnal toad is introduced in Australia,
with males outnumbering the much often found climbing where it is now a serious
larger females by about three to one. among vegetation. threat to native wildlife.

R H I NODE R M AT I DA E fleshy
proboscis E L E U T H E RODAC T Y L I DA E
This South American family includes two species This group of frogs, whose eggs develop directly
with a pointed snout and cryptic coloration, and into froglets, is found throughout the Caribbean,
the males carry the eggs and tadpoles in their the southern US, and northern South America.
mouth. The remaining species is rare Some species that are extremely small have reduced
and little is known about it. numbers of digits, and produce very small numbers
of eggs—sometimes only one.

eyes with
green horizontal ½–31/4 in
back pupils 1.5–8 cm

PUERTO RICAN
COQUI
Eleutherodactylus coqui
Native to Puerto Rico,
this frog gets its name
from its two-part call—
3/4–11/4 in the “co” is a warning to
2–3 cm other males and the
“qui” attracts females.

½–1 in
1.5–2.5 cm
large toe
pads

CARETTA ROBBER FROG

DARWIN’S FROG Diasporus diastema
Rhinoderma darwinii This small and very agile frog is arboreal
Found in Chile and Argentina, this frog has a and active at night. It lays its eggs in
unique form of parental care—the young develop water-filled cavities in bromeliads
from egg to froglet in their father’s vocal sacs. growing epiphytically in trees.
360 CA N E TOA D
R hinella mar ina
One of the largest toads in the world, the
AMPHIBIANS • FROGS AND TOADS

cane toad is a hardy creature with a huge

appetite. Also known as the marine toad,
it is mainly a resident of dry environments,
scrub, and savanna. It commonly lives around
human settlements and is often seen under
streetlights, waiting for insects to fall. The
female is larger than the male, and the largest
females can lay more than 20,000 eggs in a
single clutch. Males attract females with size 4–9½ in (10–24 cm)
a slow, low-pitched trill. They have few habitat Non-forested habitats parotoid gland
distribution C. and S. America;
enemies, as at all stages of their life cycle they introduced to Australia and elsewhere
are distasteful or toxic to potential predators. diet Terrestrial invertebrates adult coloration is yellow,
olive, or reddish brown
In Australia they have become a major pest—
they are poisonous to native and domestic
animals, injurious to humans, and breed so warts on male’s skin
develop dark, sharp spines
prolifically as to be out of control. in the breeding season

NIGHT HUNTER
Protected by their poisonous
skin and unafraid of predators,
cane toads emerge at night from
their daytime hiding places to
hop around in search of prey.

dark belly
markings

short legs have

PALE BELLY powerful muscles
for jumping
The cane toad’s belly and throat are
relatively smooth, and mostly pale in
color. Toads have permeable skin and
must hide by day to conserve water.

D E B AT E
PEST CONTROL
The cane toad gets its name from
Australia, where it was introduced into
Queensland in 1935 to control insect
pests on sugar cane farms. The toads
flourished in Australia, feasting on
native fauna and building up much
denser populations than in their native
habitat. Still spreading at an alarming
rate, they now occur throughout eastern
and northern Australia, and are likely to
migrate further. Scientists are working
on methods to limit their numbers and
contain their territorial expansion.
 361
nostril

AMPHIBIANS • FROGS AND TOADS

IRIDESCENT IRIS
Like most toads, cane toads have large,
protruding eyes. They have very good eyesight,
enabling them to detect small, moving objects
and to lunge accurately at their insect prey.

wide mouth used to

take prey of any size
that will fit inside it

male’s throat is distended

when he produces his loud,
trill-like mating call

GLAND
The enormous parotoid
glands—located on each side of
the head—secrete a powerful
toxin that is distasteful to some
predators and lethal to most.

EAR
Toads rely on their hearing NOSTRIL
to identify potential The cane toad relies more
enemies. At night, it is on its sense of smell to find
especially important for food than other toads—and
females, who locate males breathes more with its lungs
by their calls. than with its skin.
HIND FOOT
The long toes on the hind foot,
each with a horny tip, provide
a firm grip on the ground
when a toad pushes off to
hop or jump.

FRONT FOOT prominent

In the breeding season, warts
males develop dark, horny
nuptial pads on their first
three fingers. These enable
them to clasp females
firmly during mating.

WEBBING
Compared with many other frogs and toads,
cane toads have poorly developed webbing
between their toes. This reflects the relatively
small part of their lives spent in water. web of skin
 POI SON - DA RT F RO GS ¾ in
3⁄4 in
1.8 cm
362 Called poison or poison-dart frogs, the Dendrobatidae are
2 cm

noted for their bright coloration. This warns predators that long,
slender legs MIMIC POISON
the skin of these frogs contains powerful toxins, which are FROG
Ranitomeya imitator
AMPHIBIANS • FROGS AND TOADS

derived from their insect prey. These frogs are found in the
forests of Central and South America. They are active by day. Found in Peru, this frog
is variable in color and
is similar in appearance
white stripe on to at least three other
dark brown back frog species. DORIS SWANSON’S
POISON-DART FROG
Andinobates dorisswansonae
Discovered in 2006, this species
is found only in a fragment of
1–1½ in ¾–1 in 1¼–1¾ in forest in the Colombian Andes.
2.5–3.5 cm 2–2.5 cm 3–4.5 cm
It lives on the forest floor and
on low-growing bromeliads.

LOVELY GOLDEN POISON FROG

POISON-DART FROG Phyllobates terribilis
BRILLIANT-THIGHED POISON-DART FROG Phyllobates lugubris Possibly the most toxic
Allobates femoralis This poisonous frog is found in of all poison-dart frogs,
The male of this S. American species guards leaf litter of lowland forests from this terrestrial species
the eggs laid by the female in leaf nests. Later, Nicaragua to Panama. The male occurs in lowland
he carries the tadpoles to water on his back. cares for the eggs and tadpoles. forests of Colombia.

3/8–¾ in 1½–1¾ in
1–2 cm 3.5–4.5 cm

RAINFOREST THREE-STRIPED
ROCKET FROG POISON FROG
Silverstoneia flotator Ameerega trivittata
This frog is found in This S. American frog
Costa Rica and Panama. calls by day, especially
Its tadpoles have an after rain. It lays its
upward-tilting mouth, eggs in leaf litter and
which helps them feed is common near
from the water surface. human settlements.

M A R SU PI A L F ROGS R E E D A N D SE D GE F ROGS 1–1½ in

2.5–3.5 cm
Found in South and Central America, the Also called African treefrogs, this large family, the Hyperoliidae, includes many agile
Hemiphractidae carry their eggs on their back, climbers that gather in trees, bushes, or reeds near water to find mates and lay eggs.
where they hatch directly into froglets. Some Some of the species are brightly colored, with marked differences between the sexes.
also carry the eggs in a pouch, hence the name
“marsupial frogs.” RED-LEGGED KASSINA
prominent eyes
Kassina maculosa
This is an aquatic E. African frog with adhesive
disks on its toes. Its eggs, laid on submerged
1¾–2½ in vegetation, hatch into large tadpoles.
4.5–6.5 cm

FOULASSI BANANA
FROG
Afrixalus paradorsalis
Found in W. Africa, this
red patch frog lays its eggs in a
on leg
folded leaf above water.
The male attracts females
SUMACO HORNED TREEFROG with a clicklike call.
Hemiphractus proboscideus 2¼–2½ in
5.5–6.5 cm
This species is found in Colombia, Ecuador,
and Peru. The female carries her eggs on
her back, but does not have a pouch.
 GREEN AND BLACK 1¼–1½ in bright
POISON-DART FROG 3–4 cm blue skin
Dendrobates auratus 363
The males of this species fight
to secure a territory. The male
defends the eggs and, when

AMPHIBIANS • FROGS AND TOADS

they hatch, carries tadpoles 1¼–1¾ in
to small pools in bromeliads. 3–4.5 cm

1–2¼ in
2.5–6 cm

bright long
red body forelimbs
YELLOW-HEADED
POISON FROG
Dendrobates leucomelas
Found in wet forests DYEING POISON FROG
of northern S. America, Dendrobates tinctorius
this frog derives its skin This is the blue, or azureus, form of this variable
toxin from the ants S. American species. Both sexes protect the eggs and
on which it feeds. are very aggressive while defending their territory.
rounded
snout

½–¾ in
1.5–2 cm

¾–1 in
2–2.5 cm 1¼–1½ in
3–4 cm

toes with SPLASHBACK POISON FROG RIO MADEIRA POISON FROG

adhesive pads Adelphobates galactonotus Adelphobates quinquevittatus
Living in leaf litter in Brazilian forests, Found in Brazil and Peru, this tiny frog carries
STRAWBERRY this frog lays eggs on the ground and its tadpoles to water-filled holes, where the
POISON FROG carries the tadpoles to water. female feeds them with unfertilized eggs.
Oophaga pumilio
The female of this species cares ¾ in ¾–1 in
for the young by carrying tadpoles to 2 cm 2–2.5 cm
water-filled bromeliads and feeding
them with unfertilized eggs. BRAZIL-NUT
GRANULAR POISON FROG
POISON FROG Adelphobates castaneoticus
Oophaga granulifera This species is found in
This species is found in Brazil. The males of this frog
Costa Rica and Panama. place tadpoles individually
Females care for the into small, water-filled
young by feeding them tree-holes. The tadpoles
with unfertilized eggs. are voracious feeders.

SMITH’S REED FROG

AU ST R A L I A N
Hyperolius tuberilinguis large GROU N D F ROGS
This agile frog is noted for its loud call. In the mating adhesive disk
season, thousands of males may gather around a on toes The Limnodynastidae, found in Australia and
pond to produce a deafening chorus. New Guinea, include many terrestrial and burrowing
frog species. Two recently extinct species uniquely
brooded their eggs in the stomach.

¾–1½ in
2–3.5 cm
1¼–1¾ in
3–4.5 cm
large
eyes

1¼–2¼ in
3–6 cm
BOLIFAMBA
REED FROG
Hyperolius bolifambae
This small W. African BROWN-STRIPED MARSH FROG
frog occurs in bushland, Limnodynastes peronii
and breeds in pools. This Australian frog survives dry periods by burying
The male’s call is a itself in the ground. It emerges to breed after heavy
high-pitched buzz. rain, and deposits its eggs in floating foam nests.
 T R E E F ROGS
364 The Hylidae are a large family widespread
brown
upperparts with
worldwide. They are especially well represented dark patches
in the New World. These frogs have long, slender
AMPHIBIANS • FROGS AND TOADS

limbs and adhesive disks on their fingers and toes.

Most are arboreal and nocturnal. Many gather
to breed in noisy choruses. 2–2¾ in
5–7 cm

PARADOX FROG
¾–1¼ in Pseudis paradoxa
2–3 cm This aquatic frog is so named
because its tadpoles are four times
longer than the adult. It is found
in S. America and Trinidad.
2–3½ in 1–1½ in SPRING PEEPER
5–9 cm 2.5–4 cm Pseudacris crucifer
Found in moist woodland in the
SPLENDID LEAF FROG RUFOUS-EYED STREAM FROG E. US and Canada, the spring
Cruziohyla calcarifer Duellmanohyla rufioculis peeper’s distinctive, high-
This frog occurs in C. and northern Found in forests of Costa Rica, this pitched call indicates that spring
S. America and lives high up in trees. It frog breeds in fast-flowing streams. Its is underway.
glides from one tree to another, using its tadpoles have a modified mouth with
extended webbed feet as parachutes. which they attach themselves to rocks.

2¾–3½ in
7–9 cm

2¼–3 in 1½–2 in
5.5–7.5 cm 4–5 cm

MISSION GOLDEN-EYED TREEFROG ROSENBERG’S TREEFROG ORANGE-LEGGED LEAF FROG

Trachycephalus resinifictrix Boana rosenbergi Phyllomedusa hypochondrialis
Living high in the canopy of S. American Males of this C. and S. American frog A native of dry habitats in northern
forests, this frog deposits its eggs in dig pools in damp ground in which the eggs S. America, this climbing frog reduces
water-filled tree-holes, where the are laid. They defend them against rivals water loss by rubbing a waxy
tadpoles then develop into froglets. in fights that can be fatal. secretion over its skin. adhesive
disks on toes
2¼–4¼ in
5.5–11 cm

1½–2¼ in 1–4 in 1¼–2 in

3.5–5.5 cm 2.5–10 cm 3–5 cm

COPE’S BROWN TREEFROG BOULENGER’S SNOUTED TREEFROG CUBAN TREEFROG EUROPEAN TREEFROG
Ecnomiohyla miliaria Scinax boulengeri Osteopilus septentrionalis Hyla arborea
This large C. American treefrog has fringes Found in C. America and Colombia, this frog breeds in Native to Cuba, the Cayman Islands, Males of this European treefrog
of skin on the limbs that may enable it to temporary pools created after rain. Males return night and the Bahamas, this treefrog has gather in spring, calling in loud
glide from one tree to another. after night to the same calling perch to attract mates. been introduced to Florida, where choruses to attract females.
it preys on native frogs, reducing Mating pairs descend to lay
their population. eggs in nearby ponds.

large, prominent,
red eyes 1¼–2 in 2¾–4 in
3–5 cm 7–10 cm

1½–2¾ in
4–7 cm

pale
underparts
LEMUR FROG MANAUS SLENDER-
RED-EYED TREEFROG Agalychnis lemur LEGGED TREEFROG
Agalychnis callidryas This C. American Osteocephalus taurinus
An excellent climber, this treefrog is nocturnal, This arboreal frog
treefrog mates in trees sleeping on the underside lives in S. American
overhanging water. It lays its of leaves during the day. forests. It mates after
eggs on leaves and, upon hatching, It lays its eggs on rain, and lays its eggs on
the tadpoles fall into the water. leaves over water. the surface of pools.
 SMALL-HEADED
TREEFROG NEW T ROPICA L
Dendropsophus microcephalus ZEALAND GR A SS F ROGS 365
Found in C. and S. America and
Trinidad, this frog breeds in F ROGS The Leptodactylidae are a large, varied family of
pools. It is pale yellow by day, This family, the Leiopelmatidae, mostly ground-dwelling frogs from North, Central,
and red-brown by night. and South America, and the West Indies, with a
is composed of four species of
¾–1¼ in
frogs that are all confined to New pointed snout and long, powerful hind limbs. Their
2–3 cm
Zealand. They are characterized breeding habits are varied and many lay their eggs
horizontal by extra vertebrae and, unlike most in nests of foam.
pupils frogs, they kick their legs alternately
when swimming. They live in humid warty skin
forests and are nocturnal.
1–1½ in
2.5–3.5 cm

2–4 in
5–10 cm

AUSTRALIAN COROMANDEL 1¼–1½ in

GREEN TREEFROG NEW ZEALAND FROG 3–4 cm
Litoria caerulea Leiopelma archeyi TÚNGARA FROG
Found in N.E. Australia Found only in New Zealand’s North Island, Engystomops pustulosus
and New Guinea, this terrestrial frog lays its eggs under logs. During mating, the female of this C. American
this agile climber is It is critically endangered, as a result species produces a secretion that the male whips
often found close to of habitat loss and disease. into a floating foam nest in which the eggs are laid.
human habitations.

MANTELLAS
Found only on the islands of Madagascar and
Mayotte, the Mantellidae are active by day.
Many are brightly colored, warning
predators of powerful toxins in their
SPINY-HEADED
skin. Most species are threatened
TREEFROG by the loss of habitat and by the
Anotheca spinosa international pet trade.
Native to Mexico and
C. America, this large frog
lives in bromeliads and
banana plants, laying its
eggs in water-filled cavities.
1¾–3¼ in 2–2¼ in
4.5–8 cm 5–6 cm
2¼–3¼ in
1½–3¼ in 6–8 cm
4–8 cm

MASKED TREEFROG WHITE-LIPPED BRIGHT-EYED FROG ELEGANT MADAGASCAN FROG

Smilisca phaeota Boophis albilabris Spinomantis elegans
¾–1 in
Active only at night, This large arboreal frog, found only in An inhabitant of rocky outcrops, this 2–2.5 cm
this inhabitant of Madagascar, lives near the streams in which frog is found at high altitudes, even
humid forests in it breeds. It has fully webbed hind feet. above the tree line. It breeds in streams.
C. and S. America lays
its eggs in small pools. rough, moist skin

bony
projection

2–3 in ¾–1¼ in
5–7.5 cm 2–3 cm

YUCATECAN SHOVEL- MADAGASCAN PAINTED

HEADED TREEFROG GOLDEN MANTELLA MANTELLA
Triprion petasatus Mantella aurantiaca Mantella madagascariensis
An inhabitant of lowland The bright color of this This Madagascan frog
forests in Mexico and tiny frog from Madagascan breeds in forest streams.
C. America, this frog retreats rainforests warns It is under threat due
into tree-holes, using the potential predators to loss of habitat. The
bony projection on its that its skin secretes male’s call consists
head to seal the opening. a powerful toxin. of short chirps.
 NA R ROW- MOU T H E D 2–3 in
5–7.5 cm
M E G O P H RY I DA E hornlike
366 F ROGS Found across Asia, this is a small family of
projections
on eyelids
Frogs of the large and diverse family Microhylidae frogs whose body shape and color patterns
are found in the Americas, Asia, Australia, and enable them to be camouflaged among
Africa. Most are ground-dwelling and some live leaves. They walk rather than jump,
in burrows. Most have a short snout, a plump, and most are ground-living.
often teardrop-shaped, body, and stout hind legs.
cryptic
coloration with
PAINTED TOAD black markings
Kaloula pulchra
Widespread in Asia, this ASIAN
species has adapted well to HORNED FROG
human settlements. It protects Pelobatrachus
itself with a noxious, sticky nasuta
skin secretion. This frog conceals
itself among dead
1¼–2¼ in
leaves, while it waits
3–6 cm for prey. Females lay
their eggs under
rocks and logs
in streams.

3¼–4¾ in
8–12 cm

TOMATO FROG
Dyscophus antongilii BLACK-SPOTTED
Native to Madagascar, this NARROW-MOUTHED FROG
frog stays buried in soil by day, Kalophrynus pleurostigma 2¾–5½ in
emerging to feed at night. A This frog from the Philippines protects 7–14 cm
sticky skin secretion protects itself by producing a sticky secretion.
it against predators. It breeds in small pools after rain.

PA R S L E Y F RO G S
Made up of only four species, the family
Pelodytidae is confined to Europe and
the Caucasus. Named for the green
¾–1½ in ¾–1 in markings on their skin, parsley frogs
2–3.5 cm 2–2.5 cm breed after rain, laying their eggs in
broad strips.
1¼–2 in
3–5 cm
EASTERN NARROW-MOUTHED TOAD GIANT STUMP-TOED FROG COMMON PARSLEY FROG
Gastrophryne carolinensis Stumpffia grandis Pelodytes punctatus
Found in the S.E. US, this burrowing toad This small terrestrial frog is When climbing smooth, vertical surfaces,
breeds in water bodies of all sizes. The male’s found in leaf litter in high-altitude this European frog uses its underside as a
call sounds like a bleating lamb. forests in Madagascar. suction cup. Both sexes call during breeding.

TONGU E L E SS F ROGS claws used for S PA D E F O O T

These aquatic frogs are well adapted to life in 1¼–2 in
tearing food
TOA DS
water: they have a flattened body, fully webbed 3–5 cm A small family found in Eurasia and North
hind feet, and eyes that protrude upward, Africa, the Pelobatidae are characterized by
enabling them to see above the surface. As horny projections on their hind feet. They
their common name suggests, the Pipidae lack use them to burrow into the ground, where
a tongue. They feed on a wide range of prey they wait for rain.
and scavenge on dead animals.
eggs on
female’s back
FRASER’S
CLAWED FROG
Xenopus fraseri
Found in W. and C. Africa, this wholly
aquatic frog thrives in human-altered
habitats and is harvested by people for food.
1½–3¼ in
4–8 cm

DWARF SURINAM TOAD COMMON SPADEFOOT TOAD

Pipa parva Pelobates fuscus
1–1¾ in In this wholly aquatic species found Found in Europe and Asia, this species is
2.5–4.5 cm in Venezuela and Colombia, the eggs variable in color. It has a plump body,
muscular develop on the female’s back. which it inflates when attacked.
hind legs
DIC ROGLOSSI DA E T RU E F RO GS
This diverse family of frogs is found across Africa, Asia, Known as true frogs, this large family is found in most parts of the world. Most
367
and several Pacific islands. Most are ground-living, but of the Ranidae have powerful hind limbs that enable them to jump athletically on
are found close to water. Many lay their eggs in water land and to swim powerfully in water. They typically breed in early spring, with

AMPHIBIANS • FROGS AND TOADS

and have free-living tadpoles. many laying their eggs communally.
INDIAN BULLFROG
Hoplobatrachus tigerinus
This large, voracious feeder from
S. Asia breeds during the monsoon.
The male has a particularly loud call.

2½–6½ in
6.5–17 cm

3¼–4¾ in 2¼–2¾ in
RAJAMALLY WART FROG 8–12 cm 6–7 cm
Fejervarya kirtisinghei
Found only in Sri Lanka, this frog lives
in leaf litter near streams and thrives in EDIBLE FROG PICKEREL FROG
plantations and gardens. Pelophylax esculentus Rana palustris
This frog is a hybrid between the widespread Found over much of N. America, this frog breeds in
1–1¾ in European Pelophylax lessonae and other more spring, with females laying their eggs in clumps
1½–2½ in 2.5–4.5 cm
4–6.5 cm local species. It lives in and close to water. containing two to three thousand eggs.

COMMON SKITTERING FROG

Euphlyctis cyanophlyctis 1½–3¼ in
3.5–8 cm
Widespread in S. Asia, this aquatic
frog is noted for its ability to skitter
over the surface of water.

3½–8 in
9–20 cm

WOOD FROG AMERICAN BULLFROG

MARTEN’S PUDDLE FROG Rana sylvaticus Rana catesbeianus
Occidozyga martensii The only American frog found north of the This voracious predator has tadpoles that may take
½–¾ in Arctic Circle, it breeds in early spring in four years to develop, reaching a large size. It is the
Found in China and S.E. Asia, this small frog 1.5–2 cm
occurs in pools near forest streams and rivers. temporary pools free of fishes. largest N. American frog.

black spots on
green to brown body
G OL I AT H F ROGS
The family Conrauidae comprises six species of large,
semiaquatic frogs from Africa that live and breed in
fast-flowing rivers and streams. Males are thought not
to call, and females lay their eggs among gravel and
rocks on the riverbed.

4–16 in
10–40 cm
white vocal sac
long,
webbed toes
male has
thick forelimbs
powerful hind legs
for swimming
EUROPEAN
GOLIATH BULLFROG COMMON FROG
Conraua goliath Rana temporaria
The world’s largest frog, this W. African Also known as the grass frog, this species 2–4 in
species has an aquatic lifestyle; its powerful legs lives mostly on land, migrating to ponds in 5–10 cm
and webbed feet make it a strong swimmer. spring to breed. It lays eggs in clumps.
 P H RY N O BAT R AC H I DA E warty skin ½–¾ in
1.5–2 cm
OR NAT E F ROGS
368 This family of small, terrestrial A N D GR A SS F ROGS
or semixaquatic frogs is confined Found in open country in Africa, Madagascar, and the
to sub-Saharan Africa. Most breed Seychelles, the Ptychadenidae include many brightly
AMPHIBIANS • FROGS AND TOADS

throughout the year, laying their colored frogs. A streamlined body and strong
eggs in water. They reach hind legs make them prodigious jumpers.
maturity in five months.
1¾–2¾ in
4.5–7 cm
MASCARENE
GOLDEN PUDDLE FROG RIDGED FROG
Phrynobatrachus auritus Ptychadena mascareniensis
This frog is so named because Common on agricultural land,
it breeds in very small pools. A this frog has long legs and a
ground-dwelling species, it is pointed snout. It breeds in puddles,
found in C. African rainforests. wheel ruts, and ditches.

A F RO -A SI A N T R E E F ROGS shiny green

1½–2¼ in coloration 3½–4 in
The Rhacophoridae range across Africa and much 4–6 cm 9–10 cm
of Asia, and are mostly arboreal frogs. The family
also includes flying frogs that glide from one
tree to another. Many lay their eggs in foam
nests, where the eggs and tadpoles stay
protected against predators.

1¾–2¼ in SOUTHERN
4.5–6 cm WHIPPING FROG
Taruga longinasus
Endangered by the loss of much
of its habitat, this arboreal frog
occurs in Sri Lanka’s remaining
patches of rainforest.

MOSSY FROG
AFRICAN FOAM-NEST TREEFROG Theloderma corticale WALLACE’S FLYING FROG long, fully
Chiromantis rufescens The warty skin and green color of Rhacophorus nigropalmatus webbed fore-
Found in the forests of W. and C. Africa, this Vietnamese frog camouflages This arboreal species from the rainforests and hind feet
this frog lays its eggs in a foam nest 2¾–3½ in it against moss. It curls into a ball of S.E. Asia has webbed feet that
attached to a branch overhanging water. 7–9 cm when threatened. enable it to glide between trees.

PY XICEPH A LIDA E
olive-green body 3¼–9 in
Inhabiting a variety of habitats in sub-Saharan Africa, members of with dark markings 8–23 cm
this family range in size from huge bullfrogs to typical pond frogs very wide
mouth
and tiny moss frogs. Most lay eggs in water, although some small
species lay eggs on land.

AFRICAN BULLFROG
Pyxicephalus adspersus
Males of this large African species aggressively
defend their eggs and tadpoles. They dig channels
in soil to help the tadpoles reach open water.

powerful
limbs
MEXICAN ST R A BOM AN TIDA E
BU R ROW I NG TOA D 2¼–3¼ in
6–8 cm MEXICAN Many small frogs native to South America and the
369
BURROWING TOAD Caribbean make up this family. They are all direct
The only member of the family Rhinophrynus dorsalis
Rhinophrynidae, the burrowing toad This unusually shaped
developers: there is no tadpole stage and the egg

AMPHIBIANS • CAECILIANS
specializes in digging in soil and toad spends most of hatches straight into a miniature adult.
eating ants. It has a long, thin tongue its life burrowing
that it sticks out of its narrow mouth. underground, emerging
only after rain to breed in LIMON ROBBER FROG
temporary pools. Pristimantis cerasinus
Hiding in leaf litter by
day, but arboreal by night,
this small frog is found in
AMERICAN 2¼–3½ in
5.5–9 cm humid lowland forests
S PA D E F O O T of C. America.
TOA DS ½–1½ in ¾–1½ in
This family consists of toads that live on 1.5–3.5 cm 2–4 cm
dry land and stay inactive underground
CHIRIQUI
for long periods. The Scaphiopodidae ROBBER FROG
emerge after rain to breed in temporary Pristimantis cruentus
pools. These may evaporate quickly, so This small, terrestrial
the tadpoles develop very rapidly. frog, found in C. and
S. America, lays its eggs in
crevices on tree trunks.
1½–2¼ in
4–6 cm
mottled greenish ½–1 in
brown skin 1.5–2.5 cm PYGMY RAIN FROG
PLAINS SPADEFOOT COUCH’S SPADEFOOT Pristimantis ridens
Spea bombifrons Scaphiopus couchii Found in the forests
Found in arid plains of N. America This N. American frog lives in arid of C. and S. America,
and Mexico, this frog burrows by areas, spending much of its time this tiny nocturnal frog
day. It gathers to breed in large underground. It emerges at night to thrives in gardens and lays
numbers after heavy rain. feed and, after heavy rain, to breed. its eggs in leaf litter.

CAECILIANS
Caecilians are long-bodied, limbless termites, and other insects. Others live in water PHYLUM CHORDATA
amphibians with little or no tail. and resemble eels, rarely moving onto the land. CLASS AMPHIBIA

Ring-shaped folds (annuli) in the skin These have a fin on the tail. With only rudimentary ORDER GYMNOPHIONA
FAMILIES 10
give them a segmented appearance. eyes, all rely on smell to find food and mates. A
SPECIES 214
pair of retractable tentacles between the eyes and
All caecilians live in the tropics. They vary in length the nostrils transmit chemical signals to the nose.
from 4¾ in (12 cm) to 5¼ ft (1.6 m). Most live In all caecilians, the eggs are fertilized internally.
underground, burrowing in soft soil, using their Some species lay eggs, but in others the eggs are
pointed, bony head as a shovel. They emerge at retained inside the female’s body. The young
night, especially after rain, to feed on earthworms, emerge either as gilled larvae or as small adults.

IC H T H YOPH I I DA E DER MOPHIIDA E 20 in

50 cm
Found in Asia, these caecilians lay eggs in soil near water. These thick-set, cylindrical caecilians from Africa PURPLE CAECILIAN
Gymnopis multiplicata
Females remain with their clutches, defending them until live beneath the surface and are mostly brown, gray, This terrestrial caecilian from C. America
the larvae have made their way to open water. or dark purple in color but may be yellow. Some lives in a wide range of habitats. The
species lack eyes. They give birth to live young. eggs hatch inside the female.

13 in
33 cm
CA ECILIIDA E 14½–36 in
37–91 cm
Most species in this family are burrowers. Found in
most tropical regions of the world, they vary greatly
in length, some growing to more than 5 ft (1.5 m).
In some the eggs hatch into larvae; in others the
larvae develop inside the female.
yellow stripe
along body SANTA ROSA CAECILIAN
KOH TAO CAECILIAN Caecilia attenuata
Ichthyophis kohtaoensis A rarely seen, grayish blue, burrowing species from
Found in a variety of habitats in S.E. Asia, the moist lowland forests of Amazonian Ecuador,
this caecilian lays its eggs on land, this caecilian also occurs in degraded habitats
but its larvae live in water. including plantations and gardens.
370 SA L A M AN DERS AN D NEW TS
Unlike their fellow amphibians, the the most important sense used in finding food and PHYLUM CHORDATA
frogs, salamanders and newts normally in social interactions. Most species, especially the
AMPHIBIANS • SALAMANDERS AND NEWTS

CLASS AMPHIBIA

have a slender, lizardlike body, a long terrestrial ones, are nocturnal, hiding under a log ORDER CAUDATA

tail, and four legs similar in size. or rock during the day. FAMILIES 10
SPECIES 754

Newts and salamanders, also known as urodeles, r e p ro d u c t i o n

are generally found in damp habitats and are In the majority of species, the eggs are fertilized
largely confined to the northern hemisphere. inside the female. Males, however, do not have
They are numerous in the Americas, ranging from a penis but package sperm in capsules called
Canada to northern South America. They vary spermatophores which are passed to the female
considerably in size, from species over 31/4 ft (1 m) during mating. In many species sperm transfer
in length, to tiny creatures about ¾ in (2 cm) long. is preceded by elaborate courtship in which the
male induces the female to cooperate with him.
a m ph i bious li fest y les She may, of course, reject his advances. In many
Some species, notably the newts, spend part of newts, males develop dorsal crests and bright
their life in water, part on land. Some salamander colors in the breeding season.
species live their entire lives in water, while others Many species lay their eggs in water. When
are wholly terrestrial. Most have smooth, moist the larvae hatch, they have a long, slender body,
skin through which they breathe to a greater or a deep, finlike tail, and large, feathery external
lesser extent. gills. The larvae are carnivorous, feeding on tiny
The salamanders of one family, the plethodontids, water creatures. The exceptions to this rule are
have no lungs and breathe entirely through their the wholly terrestrial species of salamander.
skin and the roof of their mouth. Urodeles have These lay their eggs on land and the larval stage
A male alpine newt sniffs a female
a relatively small head, compared to frogs and is completed within the egg, which hatches to before courting her. Odor helps to identify
toads; they also have smaller eyes, smell being produce a miniature adult. the gender and species of potential partners.

SIRENS N E W T S A N D E U ROPE A N
The wholly aquatic salamanders of the family SALA M ANDERS
Sirenidae are found in the southern US and The Salamandridae are small- and medium-sized
Mexico. They retain larval features in the adult salamanders and newts found in Europe, North Africa, 4¾–8 in
stage and resemble eels. Asia, and North America. Eggs are fertilized inside 12–20 cm
They have external gills CALIFORNIA NEWT
the female, with sperm being transferred Taricha torosa
and no hind limbs. from the male in a spermatophore This nocturnal newt enters
(sperm capsule) during an ponds in spring to mate and lay
elaborate courtship. eggs. It secretes a lethal nerve
smooth, toxin to discourage predators.
slimy skin
GREATER orange poison
20–35 in SIREN glands
50–90 cm Siren lacertina
This species with tiny
forelimbs and a finlike tail inhabits
shallow rivers, lakes, and ponds in
the S.E. US and N.E. Mexico.

TORRENT SALA M ANDERS 4¾–7 in

12–18 cm
Confined to the extreme northwestern US, the CROCODILE
Rhyacotritonidae are a small family of four species. These NEWT broad
stocky, semiaquatic salamanders lay their eggs on rocks Tylototriton verrucosus head with
Found in C. Asia, this rounded jaws
under water and have aquatic larvae. robust newt breeds in ponds after
the monsoon. Its orange markings
signal distasteful secretions.

COLUMBIA TORRENT SALAMANDER 6–12 in SHARP-RIBBED SALAMANDER

Rhyacotriton kezeri 3–4½ in 15–30 cm Pleurodeles waltl
7.5–11.5 cm flattened tail
Confined to the forests of Oregon and Washington, This large Spanish and Moroccan species has a
this salamander lays its eggs in springs. Intensive unique mode of defense. When grasped, it pushes
logging has caused its numbers to decline considerably. out the sharp tips of its ribs through its skin.
 blue
markings
cylindrical tail on tail 371

AMPHIBIANS • SALAMANDERS AND NEWTS

2¼–4¾ in
6–12 cm
orange ALPINE NEWT
underside Ichthyosaura alpestris
2¾–4 in Found across much of
7–10 cm N. Europe, this newt breeds
in early spring. The female
wraps eggs individually in
the leaves of water plants.

7–11 in
18–28 cm SPECTACLED SALAMANDER
Salamandrina terdigitata 5–6½ in
This secretive salamander is found only 13–17 cm
in Italy and lives in streams in hilly areas.
It has a long, slender, and flattened body. SPOTLESS STOUT NEWT
Paramesotriton labiatus
Found in mountain streams in China,
this newt is equipped with a large tail for
swimming. It attaches its eggs to rocks.
large,
prominent eyes

4–5½ in SARDINIAN BROOK

10–14 cm SALAMANDER
Euproctus platycephalus
Found only in Sardinia, this slender
salamander lives in streams, laying its
FIRE SALAMANDER eggs under rocks. It is endangered
Salamandra salamandra due to degradation of its habitat.
This European salamander spends most large glands
of its time on land, entering water only secrete toxin
3½–5 in
to lay eggs. The glands on its head can 9–13 cm
spray a toxic secretion at enemies.

EASTERN NEWT
Notophthalmus viridescens 2½–5½ in
6.5–14 cm LORESTAN NEWT
This pond-breeding newt is from eastern Neurergus kaiseri
N. America. Juveniles, or “efts,” are
terrestrial, bright red in color, and very toxic. Found only in Iran, this species lives
in streams. It is critically endangered orange and
due to loss of its habitat and its black limbs
popularity as a pet.

JAPANESE NEWT 3½–4¾ in 2¾–4 in

Cynops pyrrhogaster 9–12 cm 7–10 cm
This newt spends much of its life in water. Its
brightly colored belly warns potential predators
that the glands in its skin secrete a toxin.
SMOOTH NEWT
Lissotriton vulgaris
Common across Europe and W. Asia,
this small amphibian breeds in ponds.
The male mates with the female
after elaborate courtship.

GREAT CRESTED NEWT MARBLED NEWT

Triturus cristatus Triturus marmoratus
Found across Europe and C. Asia, males of this large, Found in France and Spain, this newt lives
pond-breeding species develop a spectacular dorsal crest 4–7 in in woodland, heathland, and hedgerows. 4–5½ in
in spring, and display it vigorously to potential mates. 10–18 cm It enters ponds to breed in spring. 10–14 cm
 LUNGLESS SALAMANDERS
372 Containing more than 390 species, the Plethodontidae are
the largest salamander family. Members of this family have no 3¼–5 in
8–13 cm
lungs, but breathe through their mouth and skin. Apart from
AMPHIBIANS • SALAMANDERS AND NEWTS

six European species, all live in North, Central, and South small legs
America, inhabiting a wide variety of habitats and feeding CUKRA CLIMBING
mainly on small invertebrates. SALAMANDER
Bolitoglossa striatula
A small salamander
2¾–4¼ in with webbed digits,
7–11 cm this species is nocturnal,
SEAL SALAMANDER hiding by day among
Desmognathus monticola banana leaves. It is found
This stout-bodied species lives in a in Costa Rica, Honduras,
burrow by day and is active at night. long, slender and Nicaragua.
It often perches on a rock. tail and body

ALLEGHANY MOUNTAIN
DUSKY SALAMANDER
Desmognathus ochrophaeus
Mostly terrestrial, this salamander
is often found foraging in forests in 3¼–5 in
large numbers after heavy rain, and 8–13 cm
sometimes climbing trees and shrubs.

ALLEN’S 4–6½ in THREE-LINED SALAMANDER

WORM SALAMANDER 10–16 cm Eurycea guttolineata
Oedipina alleni Found in and around water, the slender
4¼–6 in Found in leaf litter in lowland three-lined salamander is a very good
11–15 cm forests of Costa Rica, this swimmer, but spends much of
salamander coils up its elongated its time in a burrow.
body and tail when attacked.

BLUE RIDGE TWO-LINED

SALAMANDER
Eurycea wilderae
Found around springs and streams,
this small species is common in
the wooded mountains of the
black stripe along 2¾–4¼ in S. Appalachians. It mates in the fall
the side of the belly 7–11 cm and lays its eggs in winter.

2¾–4¾ in
7–12 cm

MISSISSIPPI SLIMY SALAMANDER REDBACK SALAMANDER

Plethodon mississippi Plethodon cinereus
4½–8½ in Found in hardwood forests, this terrestrial salamander A terrestrial species, this salamander hides
11.5–21 cm protects itself from predators by producing a sticky under bark by day and hunts for insects and
skin secretion. It lays its eggs on land. other prey in foliage after dark.

GIANT SALAMANDERS HELLBENDER

Cryptobranchus alleganiensis
12–30 in
30–75 cm
The Cryptobranchidae are three large, wholly aquatic species—one each This N. American species has a
flattened head that helps it to dig
from Japan, China, and North America. They feed on a wide variety of prey, a burrow under rocks, where
from worms to small mammals. The family includes the world’s largest the males guard the eggs. It
salamander—the Chinese giant salamander, which is about 6 ft (1.8 m) long. has very wrinkled skin.

flattened body

3¼–4½ ft
1–1.4 m JAPANESE
GIANT SALAMANDER
Andrias japonicus
This aquatic species is threatened
by degradation of its habitat. Some
males, called “den masters,” defend
splayed legs burrows in which females lay eggs.
 MOLE SALAMANDERS
These large creatures mostly live in burrows and
emerge to forage by night. The Ambystomatidae
4¾–7½ in comprises 33 species, all found in North America. Some
12–19 cm AXOLOTL
species, notably the Mexican axolotl, are aquatic as Ambystoma mexicanum 4–12 in
SPRING SALAMANDER adults and retain larval features, such as external gills. 10–30 cm
Gyrinophilus porphyriticus Adult axolotls never leave the water and
resemble very large salamander larvae, with
An agile inhabitant of mountain streams their deep tail and feathery external gills.
and springs, this colorful salamander is most TIGER SALAMANDER
commonly found hiding under a log or rock. Ambystoma tigrinum
A thickset salamander found 7–10 in
throughout much of N. America, 18–25 cm
this species migrates in spring
to ponds to mate and lay eggs.
broad head
with small
eyes
2¾–4¾ in
7–12 cm
ITALIAN CAVE SALAMANDER
Hydromantes italicus yellow or white
Found near streams and springs in the markings on brown 3½–4¼ in
mountains of N. Italy, this salamander or black skin 9–11 cm
lives in caves and rock crevices.
MARBLED SALAMANDER
Ambystoma opacum
This short-tailed, stocky salamander
breeds in the fall and lays its eggs in
dried-up ponds that fill with rain in winter.

3–6 in ENSATINA SALAMANDER AMERICAN GIANT CALIFORNIA GIANT SALAMANDER

Dicamptodon ensatus
7.5–15.5 cm Ensatina eschscholtzii SALAMANDERS This large, nocturnal salamander is threatened
This smooth-skinned, N. American by loss and degradation of its forest habitat.This
salamander has a plump tail, which is The four large and aggressive species of the species has an aquatic larval stage.
narrow at the base. It waves its tail at Dicamptodontidae are found in the damp conifer
enemies in a defensive posture. forests of western North America. They breed in massive head
clear, unpolluted, and permanent streams, where
their larvae live and develop for two to four years
before they metamorphose.

2–3½ in
5–9 cm 6½–12 in
17–30 cm
marbled coloration

FOUR-TOED SALAMANDER
Hemidactylium scutatum
A terrestrial salamander, the adults of this species MUDPUPPIES AMPHIUMAS
live in mosses, but the larvae grow in water. There A N D R E L AT I V E Occurring in eastern North America,
is a marked narrowing at the base of its tail.
Five of the six species in the Proteidae are the Amphiumidae are a family of three
found in North America; the sixth one—the large, wholly aquatic salamanders with
an eellike body and tiny limbs. Females
A SI AT IC cave-living olm—lives in Europe. Members
guard the eggs in a nest on land.
of this family retain the larval form into
SALAMANDERS their adult life, with a long, slender body, Amphiumas can survive drought by
About 50 small- to medium-sized species external gills, and small eyes. burrowing into mud and forming
form the Hynobiidae. They are confined to a cocoon. They feed on worms,
Asia, with some found in mountain streams. 8–20 in mollusks, fishes, snakes, and
20–50 cm small amphibians.
They lay their eggs in ponds or streams, and
their larvae have external gills. Some species 8–12 in
have claws to grasp rocks. 20–30 cm

4–6½ in MUDPUPPY THREE-TOED 16–43 in

10–16 cm Necturus maculosus EUROPEAN OLM AMPHIUMA 40–110 cm
OITA SALAMANDER A voracious predator, the mudpuppy, Proteus anguinus Amphiuma tridactylum
Hynobius dunni or waterdog, feeds on a variety of The wholly aquatic olm lives in This large salamander, with slimy skin
Females of this endangered Japanese species invertebrates, fishes, and amphibians. dark, flooded caves in Slovenia and a long tail, can give a painful bite.
produce eggs in sacs, and males then The female aggressively defends and Montenegro. It is blind and Males breed every year, while females
compete to fertilize them externally. her developing eggs. has white, pink, or gray skin. breed in alternate years.
374 REPTILES
Reptiles are a sophisticated, diverse, and successful group of
ANIMALS • REPTILES

ectothermic (cold-blooded) vertebrates. Although commonly

associated with hot, dry environments, they are found in a
wide range of habitats and climates around the world.

PHYLUM CHORDATA The first reptiles evolved from amphibians more

CLASS REPTILIA than 295 million years ago. They are the ancestors
ORDERS 4 not only of modern-day reptiles but also of mammals
FAMILIES 92
SPECIES 11,050
and birds. During the Mesozoic Era, reptiles such
as dinosaurs, aquatic ichthyosaurs and plesiosaurs,
and aerial pterosaurs dominated Earth. Reptile
groups that still exist today evolved in this period
and survived the mass extinction event that killed
off the dinosaurs 66 million years ago.
All reptiles share certain characteristics, such as scaly
skin and behavioral thermoregulation—using outside
sources to maintain a constant body temperature, by
basking in the sun, for example. But the various groups
exhibit obvious differences. Turtles and tortoises are
Reptile scales are a protective layer of protected by a unique, heavily armored shell. Lizards,
skin with overlapping patches thickened
with keratin and sometimes bone. crocodilians, and tuataras have four limbs and a long tail.
Many squamates, or scaled reptiles—lizards, snakes,
and amphisbaenians—have evolved limbless forms.
Hot desert ecosystems are typically dominated by
lizards and snakes but reptiles of all types are found in
all habitats in the tropics and subtropics. Fewer species
are found in cooler temperate climates. Within each
ecosystem reptiles are important both as predators
and as prey. Most are carnivores, feeding on a wide
The shelled egg allows reptiles to range of other animals. A few large lizards and
reproduce out of water. The outer layer tortoises are exclusively herbivores, but many
protects the embryo from dehydration.
species are opportunistic omnivores.

b e h av i o r a n d s u rv i va l
Some species are solitary whereas others are highly
social. Although sluggish and inactive when cool, once
their preferred body temperature is reached by basking
or pressing against a warm rock, they can be very active.
Reproductive behavior can be complex, with males
Crocodiles can walk with their body actively defending territories and courting females.
raised well off the ground, whereas Although some squamates give birth to live young, in
turtles manage only a sprawling gait. most cases the male reptile fertilizes the female, who
then lays eggs in an underground nest. All reptiles are
independent and self-feeding from birth, although some
crocodilians give parental care for two or more years.
Reptiles are exploited by humans for their skin or
as food. This, along with habitat loss, pollution, and
climate change, threatens the survival of many species.

Reptiles modify their behavior to SEA TURTLE IN ITS ELEMENT

regulate body temperature. They bask in The green turtle may appear primitive but, with efficient
the morning sun to absorb heat energy. flippers and a flattened shell, is well adapted to its aquatic life.
r e p t i l e g ro u p s
By the Mesozoic Era, reptiles had
diversified. Turtles and crocodilians
had diverged early on, and later the
squamates—lizards, snakes, and
amphisbaenians—appeared.
t u rt l e s a n d
t o rt o i s e s

376
t u ata r a

383
lizards

384
a m ph i s ba e n i a n s

393
s na k e s

394
c ro c o d i l e s a n d
a l l i g at o r s

404
376 T U RT L E S A N D T O RT O I S E S
With their bony shell, stout limbs, forelimbs into flippers. Although they need to PHYLUM CHORDATA
and toothless, beaklike mouth, turtles breathe air, many turtles are tolerant of low oxygen
REPTILES • TURTLES AND TORTOISES

CLASS REPTILIA

and tortoises have changed little from levels and can remain submerged for hours. Turtles ORDER TESTUDINES

species that lived 200 million years ago. have a slow metabolism and are generally long-lived. FAMILIES 14

Some species are carnivores, others herbivores, SPECIES 353

The order includes sea turtles, freshwater turtles, but most turtles are omnivorous. Animal prey is
and terrestrial tortoises. Some extinct turtles were either slow-moving or ambushed as the turtle lies
gigantic, but most modern ones are more modestly hidden. Food is broken up by the “beak,” a sharp,
sized. The exceptions are the large sea turtles and a keratinous jaw covering.
few terrestrial tortoises inhabiting isolated islands.
n esti ng an d br eedi ng
p ro t e c t i o n a n d l o c o m o t i o n Turtles do not defend territories, but they can
A turtle’s shell is formed of numerous fused bones, have extensive home ranges and may develop social
overlaid by scutes. The domed upperpart is known hierarchies. Otherwise turtles often congregate
as the carapace and the lower part, the plastron. on river banks or lakesides to bask or nest.
Scutes are horny plates and new ones form beneath Both in water and on land, males of some species
the old ones each year. The degree of protection engage in elaborate courtship of females before
provided by the shell varies between species and copulation. Fertilization is internal and the females
is much reduced in some aquatic turtles. Not all lay shelled eggs, like those of other reptiles and
species can retract the head into the shell—some birds. These are spherical or elongated, with rigid
tuck their head along one shoulder under the or flexible shells. Clutches are laid in nests dug up
edge of the shell. by the female in terrestrial sites. Nearly all marine
Tortoises are slow animals, but some sea species come to land only to nest. In many species,
A green sea turtle hatchling enters
turtles can reach speeds of 19 mph (30 kph) when but not all, the gender of the hatchlings depends the sea. Many of its breeding beaches are
swimming, as a result of the modification of their on incubation temperature. protected, but it remains endangered.

AU ST RO -A M E R ICA N 13½ in
34 cm
AFRICAN
SIDE-NECKED SIDE-NECKED
T U RT L E S MACQUARIE
TURTLE T U RT L E S
From South America and Australasia, the Chelidae Emydura macquarii Most of the Pelomedusidae are carnivorous
Widely distributed
include carnivorous and omnivorous species. Their through the Murray River and occupy freshwater habitats. When
characteristically long neck cannot be retracted, basin of Australia, this threatened, they can hide their head and
so it is turned sideways under the edge of the shell. species eats amphibians, neck beneath the edge of the shell. Found
They lay elongated eggs with leathery shells. fishes, and algae. Males throughout Africa and Madagascar, they
are smaller than females.
survive dry conditions by burying
themselves in mud.

brown carapace
10 in
25 cm
REIMANN’S 30 in
SNAKE-NECKED TURTLE 75 cm
Chelodina reimanni COMMON SNAKE-NECKED TURTLE
This turtle from New Guinea eats Chelodina longicollis
crustaceans and mollusks. When scalloped A shy, freshwater turtle of Australia, this species
threatened, it tucks its large head carapace with has a long neck, which enables it to raise its
under the side of its carapace. keel (central ridge) head out of water and capture prey.

20 in MATAMATA
50 cm Chelus fimbriatus
This S. American turtle uses its
unusual appearance as camouflage
when ambushing prey, which it 8 in scales on head
sucks into its mouth. 20 cm resemble helmet
AFRICAN HELMETED TURTLE
Pelomedusa subrufa
long Widespread in sub-Saharan Africa,
snout
this carnivorous turtle is highly
sociable and often hunts in packs
to bring down large prey.
 BIG - H E A DE D T U RT L E AMERICAN SIDE-NECKED
The sole member of the Platysternidae, the big-headed turtle is an R I V E R T U RT L E S 377
endangered species from shallow forest streams in southern China and Closely related to African side-necked turtles, most species of the family
Southeast Asia. It forages in streams, bottom- Podocnemididae are found in tropical South America, barring one in

REPTILES • TURTLES AND TORTOISES

walking rather than swimming. Madagascar. These are herbivorous turtles found in various freshwater
habitats. They cannot retract their neck into their shell.
7 in
18 cm
RED-HEADED 12½ in
BIG-HEADED AMAZON RIVER 32 cm
TURTLE TURTLE
Platysternon Podocnemis
megacephalum erythrocephala
This small, carnivorous turtle This species is found in
has a flattened body, a very swamps of the Rio Negro
large head with powerful region of the Amazon
jaws, and a long tail. basin in S. America.

SNA PPI NG T U RT L E S
22 in
Native to North and Central America, these large, aquatic turtles are noted for their 55 cm
aggression. The Chelydridae have a rough shell and a powerful head, with heavy,
crushing jaws. They are effective predators, ambushing a variety of animals, but
they also eat plants.

COMMON SNAPPING
TURTLE
Chelydra serpentina
A robust turtle, it often lies in
wait for prey half-buried in mud.
It occupies freshwater habitats
from eastern N. America as far
south as Ecuador.

massive head with

strong jaws and
sharp, pointed beak

tongue with
wormlike lure

32 in
80 cm

ALLIGATOR
SNAPPING TURTLE
Macrochelys temminckii
One of the largest
freshwater turtles in the world,
heavily built carapace with this N. American species has a
three rows of conical scutes worm-shaped lure on its tongue
to attract prey as it lies in wait.

SOF T SH E L L T U RT L E S grayish-green
carapace with
raised ridges
These aquatic predators inhabit freshwater habitats in North America, Africa,
and southern Asia. The Trionychidae have a flattened shell covered with
leathery skin rather than keratinous scutes. The adult shell length ranges in
size from 10 in (25 cm) to more than 31/4 ft (1 m).

22 in
55 cm

10½ in 14 in
SPINY SOFTSHELL TURTLE INDIAN FLAPSHELL TURTLE 27 cm 35 cm CHINESE SOFT-SHELLED TURTLE
Apalone spinifera Lissemys punctata Pelodiscus sinensis
A native of eastern N. America, This Indian turtle has a rear flap on either side Hunted for its meat, this turtle of
this species eats mainly insects and of the plastron that protects the hind limbs E. Asia is rare in its native habitat, but tens
aquatic invertebrates. when they are withdrawn into the body. of thousands are bred annually on farms.
 PIG -NOSE D T U RT L E AMERICAN
378 The only species in the family Carettochelyidae, this turtle MUD AND
is an omnivore. Its carapace lacks hard scutes, but is still rigid. M U SK T U RT L E S
Its snout is adapted for breathing air while submerged. These turtles from the New World
REPTILES • TURTLES AND TORTOISES

emit a strong scent when threatened.

PIG-NOSED RIVER clawed flippers The Kinosternidae tend to walk along 5 in EASTERN MUD TURTLE
TURTLE the bottom of lakes and rivers, rather 13 cm Kinosternon subrubrum
Carettochelys insculpta than swim, and are opportunistic An omnivore, this freshwater
This nocturnal turtle is turtle feeds on the bottom of
omnivores. They lay elongated, slow, shallow watercourses
found in New Guinea and
N. Australia. Like sea turtles, hard-shelled eggs. in the S.E. US.
it has forelimbs modified as
flippers for aquatic flight. 5 in
13 cm
COMMON MUSK TURTLE
Sternotherus odoratus
This freshwater turtle from eastern
28 in
70 cm N. America is an omnivore. When
threatened, it not only exudes a
nauseating musk, but also bites.

leathery carapace
L E AT H E R BAC K with seven keels
SE A T U RT L E LEATHERBACK
5 ft SEA TURTLE
There is only one species in the Dermochelyidae. 1.5 m Dermochelys coriacea
This turtle is capable of maintaining an elevated Feeding mainly on jellyfish, this
body temperature, which allows it to swim in cold oceanic sea turtle is the world’s
largest turtle. It has a worldwide
waters. The shell has no scutes; the leathery skin distribution that includes
covers a layer of insulating oily tissue. subarctic waters.
clawless flippers

SE A T U RT L E S
Sea turtles occur throughout the world’s oceans,
mainly in coastal waters. They are highly adapted
to the marine environment, with a streamlined
body and broad, paddlelike limbs.
The Cheloniidae come to land
to nest on beaches.
Most species in
this family are 4 ft
1.2 m
endangered.

large head with

prominent eyes
vertebral keel and beak
in younger
turtles

3¼ ft
LOGGERHEAD SEA TURTLE pale coloration 1m
Caretta caretta on underside 30 in
This carnivorous species occurs in coastal waters 75 cm
worldwide, but undertakes long migrations
between feeding and nesting grounds.

almond-shaped
eyes

4¼ ft
1.3 m OLIVE RIDLEY
SEA TURTLE HAWKSBILL
Lepidochelys olivacea SEA TURTLE
GREEN SEA TURTLE A species of mainly Eretmochelys imbricata
Chelonia mydas tropical, shallow coastal This turtle uses its horny
This is the only sea turtle known to bask waters, this turtle jaws to forage for mollusks and
flattened carapace on land. It is found in temperate and tropical eats a wide range of other prey. It is found in tropical
with large scutes oceans worldwide and is a strict herbivore. invertebrates and algae. oceans throughout the world.
PON D T U RT L E S row of spines
on carapace
FALSE MAP TURTLE
Graptemys pseudogeographica
These turtles range from being fully aquatic This N. American turtle occurs in 379
freshwater habitats with abundant
to fully terrestrial. Most of them are found in vegetation. Females are almost
North America, but one species in this family twice as large as males.

REPTILES • TURTLES AND TORTOISES

comes from Europe. Diet varies between
species, although many are herbivores. 10½ in
The Emydidae are often brightly colored 27 cm
and intricately marked.

11 in
28 cm pattern of
yellow lines on
strong, clawed neck and head
10½ in front legs
27 cm
DIAMONDBACK
TERRAPIN
Malaclemys terrapin
This is a diurnal species of
9 in brackish waters in eastern
23 cm N. America. It has powerful
RED-EARED SLIDER YELLOW-BELLIED SLIDER jaws adapted for eating
Trachemys scripta elegans Trachemys scripta scripta crustaceans and mollusks.
This N. American species is mainly herbivorous Named for its habit of sliding into water
and is common in the pet trade. Its ability to when disturbed, this turtle of the S. US distinctively
colonize new habitats has allowed it to become is a diurnal omnivore. patterned shell
widely established in Europe and Asia.
8 in
10 in 20 cm
25 cm

PAINTED TURTLE CAROLINA BOX TURTLE

Chrysemys picta Terrapene carolina
Widely distributed in N. America, Males of this N. American
this small, freshwater turtle is active species have evolved highly curved
during summer. During winter, claws that help grip the female’s 5½ in
it lies torpid under water. domed shell while mating. 14 cm
ORNATE BOX TURTLE
5 in 10 in Terrapene ornata
13 cm 26 cm strong claws used This is an omnivorous, terrestrial turtle
for burrowing of central N. America. It digs burrows
to escape extreme heat or cold.
8½ in 5 in
21 cm 13 cm

SPOTTED TURTLE CHICKEN TURTLE

Clemmys guttata Deirochelys reticularia
Identifiable by its spots, this small turtle This shy turtle of swamps in eastern
feeds on aquatic invertebrates and plants N. America extends its long neck to WOOD TURTLE
in the marshes of eastern N. America. strike at crayfish and other prey. EUROPEAN POND TURTLE Glyptemys insculpta
Emys orbicularis This turtle occupies the damp forests
Widespread in Europe, this is a highly in northeastern N. America. Unusually,
aquatic turtle. It basks on logs or rocks, but males and females perform an elegant
olive skin rapidly dives into the water if disturbed. dance together during courtship.
with irregular
yellow spots 15 in 16 in
38 cm 40 cm

10 in
26 cm FLORIDA REDBELLY TURTLE RED-BELLIED TURTLE
BLANDING’S TURTLE Pseudemys nelsoni Pseudemys rubriventris
Emydoidea blandingii Restricted to Florida, this turtle Restricted to the N.E. USA, this
Found mainly across the Great Lakes region of lives in lakes and slow-moving streams. omnivorous, diurnal species prefers
N. America, this turtle is an omnivore and During courtship, the male caresses the large, deep bodies of water. Females
especially skilled at capturing crayfish. female’s head with his forelimbs. are larger than males.
380 A L DA BR A
G I A N T T O RT O I S E
A ldabrachelys gigant ea
The last remaining species of giant tortoise on islands in the Indian
Ocean, the Aldabra giant tortoise can weigh more than 660 lb (300 kg).
Though resident on three islets of the Aldabra atoll, over 90 percent
of the tortoises live on Grande-Terre—the largest islet by far—
despite the scarcity of water and inadequate supply of vegetation.
Poor conditions there have inhibited the growth of individual tortoises,
with many not reaching sexual maturity. However, they are highly
sociable, more so than the larger tortoises on other islets. Males are
size 4 ft (1.2 m)
larger than females, but courtship is a gentle affair. Their eggs are habitat Grassy areas
buried underground, and hatch in the rainy season. The whole distribution Aldabra, Indian Ocean
population is vulnerable to natural disasters and rising sea levels. diet Vegetation

HORNY BEAK
The tortoise’s large mouth
contains no teeth. It cuts
vegetation with its sharp,
horny beak, takes it into
the mouth with its tongue,
and swallows it whole.
EAR
Tortoises have no
external ear flap;
so the eardrum is
situated in a hollow.

EYE
The eye is relatively large
and has a well-developed
eyelid. Tortoises can see
in color, particularly in
LEATHERY VISAGE the red and yellow parts
The skin is tough, leathery, and— of the spectrum; this
depending on islet—either gray or probably helps them
brown. The skin is creased into to find colorful fruit.
folds around the neck. If it senses
danger, the tortoise pulls its head
back inside its shell.

horny scales
cover the front
and hind limbs

DIGGING CLAWS
Tortoises have sturdy, elephantlike
hind limbs, with five claws on each
foot. Females have larger claws
than males, which they use when
excavating a nesting site.
hind limbs are
elephantine, with
strong claws

FORELEG TAIL
The front limbs are cylindrical The giant tortoise has
in shape, and longer than the hind a short tail, and can tuck
limbs. This allows the tortoise it to one side under the
to raise its body well above the back of the carapace.
ground as it walks. The legs are Males have longer tails
covered in large, tough scales. than females.
 scutes on the MADE TO LAST
upper shell show Compared to giant tortoises from the
growth rings Galapagos Islands, this species has a
381
more rounded head, a pointed snout,
and a smiling mouth. It can live for
more than a hundred years.
 POND AND RIVER TORTOI SE S
382 T U RT L E S Found in the Americas, Africa, and
Found both in the Old and New Worlds, the Geoemydidae southern Eurasia, members of the family
are freshwater and terrestrial turtles, ranging in adult Testudinidae can reach great sizes. They
REPTILES • TURTLES AND TORTOISES

shell length from 51/2–20 in (14 to 50 cm). They vary have a dome shell into which they can 12 in
in their dietary preferences from being herbivorous to retract their head. Fully terrestrial, 30 cm
carnivorous. Many exhibit sexual dimorphism, with tortoises are characterized by elephantine
females being larger than males. limbs. They lay hard-shelled eggs. DESERT TORTOISE
Gopherus agassizii
This tortoise lives in small
9 in BROWN burrows in deserts of southwestern
23 cm WOOD TURTLE N. America. Mainly vegetarian,
Rhinoclemmys 13 in it sometimes preys on animals.
annulata 33 cm
A herbivorous
tortoise, this species
lives in tropical forests of
C. America. It is mainly
active in the mornings
and after rains.
ELONGATED TORTOISE
Indotestudo elongata
12 in Found in tropical S.E. Asia, this tortoise
30 cm eats fruits and carrion. It hides in moist
leaf-litter when the weather is dry.

16 in
40 cm

GOLDEN COIN TURTLE

Cuora trifasciata
This is a carnivorous turtle from S. China. Its name
derives from its value in the illegal wildlife market.
Its use in traditional medicine threatens its survival.
pronounced
spinal keel SERRATED HINGE-BACK
TORTOISE
61/2 in Kinixys erosa
17 cm This is an omnivorous species found in
swamps in tropical W. Africa. Older
tortoises develop hinges in the rear
of their carapaces.
16 in
40 cm

28 in
70 cm

yellow
stripes
behind
eyes RED-FOOTED TORTOISE RADIATED TORTOISE
Chelonoidis carbonaria Astrochelys radiata
Although known to eat carrion, this tortoise Restricted to S. Madagascar, this endangered
YELLOW-MARGINATED BOX TURTLE is mainly vegetarian. It is found in a variety of species eats mainly plants and is active during
Cuora flavomarginata habitats of northeastern S. America. the early part of the day.
An omnivore of China’s rice paddies,
this species avoids deep water and
spends hours basking on land. highly INDIAN STAR
domed scutes TORTOISE
5 in
Geochelone elegans
5 in 13 cm Found in dry areas of
13 cm India and Sri Lanka,
this herbivorous species
mates and breeds in
the monsoon season.

15 in
38 cm

ASIAN LEAF TURTLE

Cyclemys dentata BLACK-BREASTED LEAF TURTLE
This omnivorous turtle Geoemyda spengleri
is found in slow-moving An inhabitant of wooded mountains
waters in S.E. Asia. It in S. China, this turtle eats small
defends itself by emitting invertebrates and fruits. Its carapace
foul-smelling liquid. is rectangular, keeled, and spiky.
 PANCAKE TORTOISE
Malacochersus tornieri 383
Found in E. Africa, this
omnivorous species inhabits
7 in rocky areas. Its flat shape 4 ft

REPTILES • TUATAR A
18 cm allows it to hide in cracks. 1.2 m

growth rings on the

carapace
of juveniles
flattened
scute

ALDABRA GIANT TORTOISE

Aldabrachelys gigantea
Restricted to the Aldabra atoll in the large forelimbs
GALAPAGOS TORTOISE Indian Ocean, this large herbivore is able
Chelonoidis nigra to drink through its nostrils.
One of the world’s largest tortoises,
this mainly vegetarian species has
11 subspecies, from different islands
in the Galapagos archipelago. 7½ in
19 cm

HERMANN’S TORTOISE
Testudo hermanni
This herbivorous species inhabits dry forests
jaw with sharp of coastal Italy and S. France. It hibernates
cutting edge for during the cold winter months.
eating vegetation
4 ft
1.2 m STEPPES TORTOISE
Agrionemys horsfieldii
A herbivore from the dry
deserts and steppes of
C. Asia, this species escapes
11 in the daytime heat by
28 cm sheltering in burrows.

T U ATA R A
Superficially like a lizard, New Zealand’s Tuataras are very long lived, but are vulnerable to PHYLUM CHORDATA

tuatara belongs to a much more ancient introduced ground predators. Inhabiting coastal CLASS REPTILIA
ORDER RHYNCHOCEPHALIA
order of reptiles. Its closest relatives forests, they are active at low body temperatures FAMILIES 1
became extinct 100 million years ago. and emerge from their burrows at night to hunt for SPECIES 1
invertebrates, and birds’ eggs and chicks. Males are
The tuatara has many anatomical features that set it territorial and nesting is communal. Eggs take up
apart from lizards. Most striking are its wedge-shaped to four years to form and are incubated for 11 to 16
“teeth”—actually serrations of the jawbone. months before hatching. Incubation
The upper jaw has a double row that fits temperature determines the
over a single row on the lower jaw. sex of the hatchlings.

T U ATA R A
Often portrayed as “living fossils,”
the Sphenodontidae are modern
representatives of reptiles that
coexisted with dinosaurs. This stout tail
primitive species is found only on TUATARA
offshore islands of New Zealand. Sphenodon punctatus
This reptile has clawed limbs to dig burrows.
It can shed its tail to escape predators. The 23½ in powerful,
male uses its dorsal crest for display. 60 cm clawed limbs
384 LIZARDS
Typically, lizards have four legs and a mechanisms depend on agility, camouflage, PHYLUM CHORDATA
long, thin tail, but there are also many and bluff. Many species are able to shed their
REPTILES • LIZARDS

CLASS REPTILIA

legless species. They have scaly skin tail to distract the attention of predators as ORDER SQUAMATA

and firm jaw articulation. they escape. The tail grows back. In some FAMILIES (LIZARDS) 38

families of lizards, the skin can change color. SPECIES 6,687

All lizards are ectothermic, obtaining heat This ability can be used for camouflage or for
energy from the environment. Although seen sexual or social signaling. Unusually, in many D E B AT E
as predominately tropical or desert animals, they lizards the pineal gland on the top of the head S Q U A M ATA : A N
have a global distribution; lizards are found from acts as a light-sensitive “third eye.” ALL-INCLUSIVE ORDER
beyond the Arctic Circle in Europe to the tip of
South America. Highly adaptable, they occupy va r i e d l i f e s t y l e s Traditionally, lizards and snakes were
considered two distinct groups, but
a wide range of terrestrial habitats with many Although some species are solitary, many lizards modern genetic research makes it clear
being arboreal or rock-dwelling. Many legless have complex social structures, with the males that they are not. Primitive squamates
species are adapted for burrowing, a few lizards maintaining their territories through visual (scaled reptiles) were lizardlike animals
are effective gliders from trees, and others are signals. Many species lay eggs in underground that arose in the mid-Jurassic. Available
semiaquatic, including a marine species on nests, while others retain the eggs in the oviduct evidence suggests that snakes evolved
the Galapagos Islands. until hatching. There are also truly viviparous from lizards during the mid-Cretaceous.
species, in which the mother provides nutrition By this time various families of lizards
s t r at e g i e s f o r s u rv i va l via a placenta. Males have paired hemipenes— had branched off on their own
Lizards range in length from about 1/2 in (1.5 cm) to sexual organs used for internal fertilization— evolutionary course. Some lizard
families are thus closer to snakes
10 ft (3 m) in the case of the Komodo dragon, but although some species are parthenogenetic, with than they are to other lizards. Of living
the majority are between 4 and 12 in (10–30 cm). females reproducing without the participation lizards, the monitors are thought to be
Although most are carnivores, about two percent of a male. Some lizards tend their eggs during those most closely related to snakes.
of species are primarily herbivorous. Many lizards incubation but very few exhibit any maternal
are prey for other carnivores. Their defense care of the hatchlings.

CHAMELEONS
3¼ in
Restricted to the Old World, the members of the 8 cm
family Chamaeleonidae have long limbs with grasping
feet and a prehensile tail for gripping branches—an
adaptation for life in trees. Their eyes are capable of
moving independently in any direction to locate
insects or small vertebrates. They catch their prey BEARDED PYGMY-CHAMELEON PARSON’S CHAMELEON 28 in
with a long, sticky tongue that shoots out from the Rieppeleon brevicaudatus Calumma parsonii 70 cm
mouth. Their ability to change color is for display This unusual, small chameleon is from The largest chameleon in the world, this
E. Africa. Drab coloration and patterning species is restricted to Madagascar. It hunts
and camouflage. Many species are threatened along its body allow it to resemble a dead leaf. invertebrates in the canopy of montane forests.
with extinction. 12 in
30 cm
prehensile tail

green skin coloration

sometimes changes to brown

12 in
30 cm
MEDITERRANEAN
CHAMELEON
Chamaeleo chamaeleon JACKSON’S
This chameleon spends its CHAMELEON
time in bushes searching for Triocerus jacksonii
insects. It is found in N. This diurnal, arboreal species is found in
dorsal Africa and around the E. Africa. The male is identifiable by the three
spines Mediterranean. horns on its snout, which are for display.

grasping, 231/2 in
four-toed feet 60 cm

20 in 22 in
51 cm 56 cm PANTHER
CHAMELEON
Furcifer pardalis
GIANT SPINY CHAMELEON Found only in dry forests VEILED CHAMELEON
Furcifer verrucosus in Madagascar, this lizard Chamaeleo calyptratus
This large chameleon is a native of the humid, hunts insects in trees by This species is from the southern coast of the
coastal region of Madagascar. A shy species, it stealth. Males are Arabian Peninsula. Males have a large casque
relies on camouflage to ambush its insect prey. highly territorial. on the head, while females have a smaller one.
C H I S E L-T E E T H L I Z A R D S
Common in Africa, southern Asia, and Australasia, the small lizards
385
of the family Agamidae—which also includes flying lizards and water
AUSTRALIAN WATER DRAGON 3¼ ft
dragons—are the Old World equivalents of the iguanas. Spines, flaps, Intellagama lesueurii 1m
and crests often adorn the head and back of these lizards, which lay The largest water dragon in Australia, this
soft-shelled eggs. Males are brightly colored while the females are duller. species lives beside water into which it
dives to escape predators. Adults eat
invertebrates and small vertebrates.
16 in
40 cm

ORIENTAL 3¼ ft long limbs

GARDEN LIZARD 1m
Calotes versicolor ASIAN WATER
This agile, diurnal lizard is often DRAGON
found hunting insects in trees Physignathus cocincinus
around human habitation. It A powerful swimmer, this lizard seeks
is common in S. Asia. refuge in water when threatened. It is
long, laterally
flattened tail used found in riverside trees in S. Asia.
in swimming
long tail helps 16 in NORTH AFRICAN MASTIGURA
with balance 40 cm Uromastyx acanthinura
when climbing This herbivorous lizard inhabits
the harsh desert environments of
BOULENGER’S N. Africa. It uses its clublike,
PRICKLENAPE spiny tail in defense.
10 in Acanthosaura crucigera
26 cm This slow-moving lizard from Asia
preys on insects while perched on a
branch. Males use the long spines on
their neck when fighting each other.
chisel-shaped teeth on
outer rim of mouth
CENTRAL BEARDED DRAGON 20 in
Pogona vitticeps 50 cm
Found in arid woodland in Australia, this
lizard has a beard of spines. It lays eggs
in an underground nest.

8 in
20 cm
GREEN-STRIPED TREE DRAGON
Diploderma splendidum
This colorful lizard is from humid
montane forests of China, where it
hunts insects. It lays small clutches
of five to seven eggs.

5–7 in
15–18 cm

THORNY DEVIL
Moloch horridus
This Australian lizard feeds exclusively on ants. The
large skin frill body is covered with large, thorny spines, which make
makes head it difficult for a predator to swallow. The spines also
look larger channel rainwater or morning dew into its mouth.

35 in
90 cm
16 in
FRILLED LIZARD 40 cm
Chlamydosaurus kingii
Common in subtropical woodland
strong hind legs in Australia, this species feeds on RAINBOW LIZARD
allow this lizard to insects and other lizards in trees and Agama agama
run away bipedally on the ground. When threatened, it This common, insectivorous lizard from Africa is gray at
unfurls its frill and opens night, but becomes brightly colored in sunlight. Males
its mouth wide. use their bright skin colors during territorial disputes.
386 PA N T H E R C H A M E L E O N
Fu rcifer pardalis
This large chameleon is endemic to Madagascar off the east coast of southern Africa,
and has also been introduced to Mauritius and Réunion. This species lives in trees
in humid scrubland, and its feet are so well adapted for grasping branches that it is
difficult for the lizard to walk across a flat surface. Active during the day, it moves slowly
through the branches to hunt its insect food by stealth. When it spots its next meal,
the chameleon focuses on its victim with both eyes, then shoots out its long tongue to
grasp the insect and pull it back into its large mouth. The chameleon’s uncanny ability size 16–22 in (40–56 cm)
habitat Trees in humid scrub
to change color is an indicator of mood and social status only, and is not used for distribution Madagascar
camouflage. When faced with a rival, it rapidly inflates its body and changes color, diet Arthropods, crustaceans
putting on a display of dominance that is usually enough to decide a dispute.
a ridge of protective
spines runs along the casque, or bony shield,
midline of the back at the back of the head
 EYES TONGUE
Uniquely, chameleons The tongue is very long
are able to move each eye and is fired from the mouth
387
independently of the other. at great speed. This means
With eyes pointing in that the chameleon can seize
different directions, the unsuspecting prey from

REPTILES • LIZARDS
chameleon can look out for some distance away.
predators while also
searching for prey.
Chameleons do
not have ears.

CLAWED FEET
SKIN COLOR The chameleon’s toes are
The striking skin colors are caused by cells called the muscular tongue clawed, and arranged into
chromatophores, which contain a variety of pigments and its mucus envelop groups of two and three on
and reflectors. The cells’ size and dispersion of pigments the prey, pulling it to opposite sides of the foot.
change with the chameleon’s mood, often quickly and the mouth This enables the animal
dramatically, as a signal to potential rivals and mates. to grip any branch tightly.

insect prey

robust scales protrude

from the snout to form a
small horn

TAIL
Life up in the trees is made a lot
easier by the chameleon’s prehensile
tail. It acts as a fifth limb, wrapping
around branches to help with climbing.

UNDERSIDE
This chameleon was photographed
through a glass table from below.
They rarely, if ever, have to walk
on flat ground; when they do,
their feet are splayed.
wide jaw

ridge of spines

long tail

splayed foot

sharp spines run from

the mouth along the
underside of the chin
 GEC KOS
388 Widely distributed in the tropics and subtropics,
the superfamily Gekkonidae number more than
1,000 species in 100 genera. They are noted for
REPTILES • LIZARDS

being very vocal and for their ability to climb

AFRICAN
smooth surfaces. Many lay hard-shelled eggs, WESTERN BANDED GECKO 4¾ in 10 in FAT-TAILED GECKO
while others lay soft-shelled eggs. Some species Coleonyx variegatus 12 cm 25 cm Hemitheconyx caudicinctus
are viviparous. This terrestrial gecko hunts invertebrates Common in the W. Sahara, this species has
in the deserts of the W. US. Unlike many a tail that stores fat. It lacks the adhesive
geckos, it has moveable eyelids. foot pads of many other geckos.

8 in
20 cm
8½ in
21 cm

6 in
COMMON LEOPARD GECKO 15 cm MOORISH GECKO WONDER GECKO
Eublepharis macularius Tarentola mauritanica Teratoscincus scincus
Originally from southern C. Asia, Common around the To defend itself, this C. Asian species waves its tail
this gecko is a popular pet, which Mediterranean, this agile gecko slowly, causing a row of large scales on the tail
has been bred for different preys on insects. It lives on rock to rub together, making a hissing sound.
markings and colors. bright coloration faces but often enters houses.

8 in
20 cm

MADAGASCAR
DAY GECKO
Phelsuma madagascariensis 10 in
This colorful diurnal species is adhesive pads on toes 25 cm KUHL’S FLYING
typically found on trees. Like GECKO
many geckos, it lays hard-shelled TOKAY GECKO Ptychozoon kuhli
eggs that stick to a branch. Gekko gecko
Named after its harsh This gecko’s webbed toes and skin flaps
“to-kay” call, this large gecko help break its fall as it parachutes from
16 in from S.E. Asia often trees in the rainforests of S.E. Asia.
40 cm
lives in houses.

13½ in
34 cm

6 in 4 in
15 cm 10 cm

RING-TAILED GECKO BROOK’S HOUSE GECKO MEDITERRANEAN GECKO

Cyrtodactylus louisiadensis Hemidactylus brookii Hemidactylus turcicus
Found in New Guinea, this large, nocturnal This gecko lives in close proximity to humans. Often betrayed by its mewing cry, this small
gecko feeds aggressively on invertebrates Native to N. India, it now also occurs in Hong European gecko is common in houses, where
and small frogs found on the ground. Kong, Shanghai, and the Philippines. it hunts insects attracted to lights.

FLAP-FOOTED 4¾ in
12 cm ANOLES
LIZARDS Most anoles come from around the Caribbean. A diverse group, the
All lizards of this group have an elongated body, Dactyloidae are typically small, arboreal, and insectivorous. Although
no front limbs, and much-reduced hind limbs. FRASER’S DELMA often green or brown, they change skin color according to mood
Restricted to Australasia, the 36 species in the Delma fraseri and environment. Both sexes aggressively defend
family Pygopodidae hunt insects by burrowing This insectivorous, Australian their territories.
lizard inhabits spinifex grassland. It
or on the surface. They are related to geckos is well adapted to moving through the
and lay soft-shelled eggs. stiff blades of grass.
KNIGHT ANOLE
Anolis equestris 20 in
50 cm
8½ in 23½ in The largest anole, this species is
21 cm 60 cm restricted to Cuba. Adhesive pads on
its toes enable it to scale smooth walls.
COMMON
SCALY FOOT BURTON’S SNAKE-LIZARD CAROLINA ANOLE
Pygopus lepidopodus Lialis burtonis Anolis carolinensis 8 in
Snakelike in appearance, this lizard is Native to Australia, this lizard has an Males show their dominance by bobbing 20 cm
widespread in Australia. It is a diurnal predator elongated, wedge-shaped snout that it uses their heads and flaring their brightly
of insects and tunnel-inhabiting spiders. to grasp skinks, which it consumes whole. colored throat fan.
HELMETED LIZARDS
Nine species of arboreal lizards from Central and saillike dorsal crest
389
South America constitute the family Corytophanidae. 13½ in
They are closely related to the iguanas. All species have bright 34 cm
orange iris

REPTILES • LIZARDS
a well-developed head crest. Long legs and a long HELMETED IGUANA
tail allow these lizards to run at high speeds Corytophanes cristatus
to evade predators. This C. American iguana lives
on a diet of large arthropods. By
feeding infrequently, it minimizes
the time it spends in the open or
is conspicuous to predators.

hard, conelike casque at

rear of head

26 in
65 cm
GREEN BASILISK long legs
Basiliscus plumifrons
Native to C. American
rainforests, this species inhabits
riverbanks. The crests on its EASTERN
head, back, and tail are CASQUE-HEADED IGUANA slender, green
supported by bony spines. Laemanctus longipes feet and legs
This large, insectivorous species lives in
small groups of one male and two to three
females. It is native to C. America. 28 in
70 cm
long tail aids balance when
climbing or running

NORT H A M E R ICA N
SPINY LIZARDS COLORADO DESERT
FRINGE-TOED LIZARD 3¼ in
Restricted to North and Central America, the Phrynosomatidae are a Uma notata 8 cm
diverse family of lizards that prefer arid environments and hunt insects. Closable nostrils and ear flaps, overlapping
jaws, and interlocking eyelids help protect this
They are generally small, dull in color, and spiny. desert-living species as it burrows in sand.
Most lay eggs, but those living at high
altitudes are viviparous.

6 in
DESERT HORNED LIZARD 15 cm
GREEN SPINY LIZARD 8 in Phrynosoma platyrhinos
Sceloporus malachiticus 20 cm This lizard of N. American deserts
This diurnal, arboreal lizard from primarily eats ants. Its flattened body
C. America has stiff, keeled scales, is an adaptation to maximize heat
which give it a spiny appearance. absorption while basking in the sun.

IGUANAS MARINE IGUANA

Amblyrhynchus cristatus
Largely confined to the Americas, the Iguanidae 2¼–5 ft Native to the Galapagos
0.7–1.5 m Islands, this aquatic lizard is
number nearly 30 species in 8 genera. They are well adapted to feeding on
diverse in color. Most are diurnal, carnivorous underwater algae. Nasal
predators, although larger species are glands help remove salt.
herbivorous. They all lay eggs.
dorsal crest
GREEN IGUANA
Iguana iguana 6½ ft
Widespread in C. and 2m
S. America, this large iguana
BLACK is herbivorous. Males defend
IGUANA their territory by vigorously
Ctenosaura similis 35 in nodding their heads to
This gregarious species 90 cm display dominance.
from C. America lives in
colonies with a dominant
male. Although usually
herbivorous, it sometimes
eats small lizards.

long,
whiplike tail
 SKINKS
390 Distributed worldwide, the Scincidae are 14 in
35 cm
a diverse group of 1,400 species. While
many are active diurnal predators, several
REPTILES • LIZARDS

others are nocturnal, limbless, burrowing

lizards. They use chemical as well as visual
communication. Although typically
oviparous, many species are viviparous.
bright coloration FIRE SKINK
on flanks Mochlus fernandi
This insectivorous skink is native to
humid, forested areas of W. Africa.
Its attractive coloration makes it a
weak legs popular lizard to keep in captivity.

8½ in FIVE-LINED SKINK
EMERALD TREE SKINK 21 cm Plestiodon fasciatus PERCIVAL’S LANCE SKINK 12 in
Lamprolepis smaragdina This N. American skink Acontias percivali 30 cm
Native to islands of the coils around its eggs to protect This legless African skink burrows
W. Pacific, this arboreal them during incubation. It through leaf litter to hunt its
10 in lizard preys on insects prefers woodland and feeds invertebrate prey. It gives birth
25 cm on bare tree trunks. on ground-living insects. to up to three live young.

WA L L A N D
SAND LIZARDS
LARGE PSAMMODROMUS
Lizards of the family Lacertidae are found in a wide Psammodromus algirus
variety of habitats throughout the Old World. They are Found in the W. Mediterranean region, 3 in
this small lizard inhabits dense, bushy areas. 7.5 cm
active predators and have complex social systems, with During breeding, males develop a red
males defending their territory. Almost all species lay 8 in patch on their throat.
eggs. These lizards typically have a large head. 20 cm
OCELLATED LIZARD
Timon lepidus
The largest European member
of the family, this lizard lives in
dry, bushy places. It eats insects,
VIVIPAROUS LIZARD eggs, and small mammals.
Zootoca vivipara
Found throughout Europe and up to 9,900 ft 6 in
(3,000 m) in the Alps, this ground-dwelling lizard lives 15 cm
in a variety of habitats. It gives birth to live young. FRINGE-TOED LIZARD
Acanthodactylus erythrurus
Found in the Iberian Peninsula and
N. Africa, this species has fringes 3½ in
of spiny scales on its toes that enable 9 cm
it to cross over loose sand.

GRAN CANARIA
GIANT LIZARD ITALIAN WALL LIZARD
Gallotia stehlini 3 in Podarcis siculus
This large species is restricted to 7.5 cm A ground-living species, found around the
32 in the shrubland of Grand Canary Island. N. Mediterranean, this lizard inhabits grassy places
80 cm It is diurnal and herbivorous. and often lives in close proximity to humans.

W H I P TA I L S A N D AMAZON RACERUNNER
Ameiva ameiva
R AC E RU N N E R S Powerful jaws allow this S. American
species to feed on small vertebrates
These fast-running American lizards occupy a range of habitats. and insects that it catches in open
Smaller species of the Teiidae are insectivorous but larger ones 18 in terrestrial habitats.
are carnivorous. All 120 species are oviparous, although many 45 cm
whiptails are all-female and reproduce parthenogenetically,
laying viable, fertile eggs without mating.

long tail used

in defense

RED TEGU
Salvator rufescens
This large species of the arid
regions in central S. America is an
active predator and scavenger, but 4 ft
is also known to eat plants. 1.2 m
 BERBER SKINK EASTERN
Eumeces schneideri BLUE-TONGUED
This diurnal skink eats insects, small vertebrates, SKINK 391
and carrion. It is native to N. Africa and
16 in Tiliqua scincoides
40 cm Characterized by its
the deserts of S.W. Asia. MANY-LINED SUN SKINK
Eutropis multifasciata eponymous blue tongue,

REPTILES • LIZARDS
Found in S. Asia, this lizard hunts insects this Australian skink is a
28 in diurnal omnivore. It
in sunlit clearings within forests. 72 cm
It gives birth to live young. bears live young.

8 in
SANDFISH SKINK 20 cm
Scincus scincus 14 in
35 cm 14½ in
This insectivorous, N. African 37 cm
skink is named for its habit of
diving into loose sand to evade
predators and keep its body cool. 6 in LITTLE BROWN SKINK SOLOMON ISLANDS SKINK
15 cm Scincella lateralis Corucia zebrata
Living in leaf litter in woodland, this This is the world’s largest
N. American skink hunts insects. skink. It is strictly arboreal,
A female can store sperm to has a prehensile tail, and is
fertilize her eggs. herbivorous. It lives in groups.

PL AT E D
LIZARDS
There are 32 species of plated lizards. 14 in
36 cm
All are from sub-Saharan Africa and 19 in MADAGASCAN
lay eggs. They have a cylindrical body 48 cm GIRDLED LIZARD
Zonosaurus
with well-developed legs for hunting madagascariensis
insects in rocky areas and the savanna. ROUGH-SCALED PLATED LIZARD This insectivorous lizard
Solitary in nature, the Gerrhosauridae Broadleysaurus major is native to Madagascar.
are often aggressive to other members This omnivorous species is widely distributed It prefers to be solitary,
in the savanna of E. Africa. It occupies crevices and feeds on the ground
of their species. in rocky outcrops or termite nests. in open, dry habitats.

GIRDLED LIZARDS M IC ROT E I I D L I Z A R DS

CAPE GIRDLED LIZARD
Limited to southern and eastern Africa, the Cordylidae Cordylus cordylus There are 165 species of the Gymnophthalmidae in tropical South
are named for the rings of spiny scales that encircle their Endemic to southern Africa, America. Generally small, they are characterized by large scales
tail. Their flattened body has limited space for live this lizard lives in dense on their back. These are diurnal and secretive insectivorous lizards
colonies. Adults are aggressive,
young or eggs. In viviparous species, this restricts litter and form social hierarchies and their dull coloration provides camouflage in leaf litter.
size; oviparous species lay only under a dominant male. Most species lay eggs.
two eggs.
8½ in BROMELIAD LIZARD
21 cm Anadia ocellata
An arboreal species
spiny found in C. America,
scales the bromeliad lizard
hunts insects and takes 3¼ in
refuge in foliage. 8 cm

muscular body

glossy head scales

 C ROCODI L E L I Z A R D MONITOR LIZARDS
392 The crocodile lizard is the only representative Widely distributed throughout tropical Africa, Asia, and
of the Shinisauridae. There are Australia, the Varanidae are the largest of all living lizards.
two recognized sub-species They have an elongated body and strong legs, and many
REPTILES • LIZARDS

found in China and produce toxic saliva. Their choice of prey depends
northern Vietnam. on their size.
This little-studied armored scales
lizard is endangered along back 5 ft
1.5 m
because of the
popularity of the CHINESE CROCODILE LIZARD
Shinisaurus crocodilurus crocodilurus
lizards as pets. Restricted to the Guangxi region of S. China, HEATH MONITOR
18 in this aquatic lizard feeds on fishes and tadpoles. Varanus rosenbergi
46 cm It basks on riverside bushes. Found around the coasts of southern Australia,
this monitor has a varied diet. It is a strong
digger and often forages underground for food.
X E NOSAU R S
KNOB-SCALED LIZARD
Found mainly in Mexico, these unusual lizards Xenosaurus grandis
have the top of their head roughened by conical A flat-bodied lizard found on the
osteoderms (bony deposits) beneath the scales. The floor of Mexican rainforests, it 61/2 ft
feeds on flying insects at night. 2m
Xenosauridae bear live offspring and protect them It remains hidden during the day. ASIAN WATER
in burrows. They prey on insects and other animals. MONITOR
Varanus salvator
This monitor is found in
rainforests and wet habitats
of S. Asia. Its large size
allows it to tackle a
10 in 4¼ ft variety of prey.
25 cm 1.3 m

SAVANNA MONITOR
NIGHT LIZARDS Varanus exanthematicus
YELLOW-SPOTTED Found in the savanna of
The 30 or so Xantusiidae species are restricted to NIGHT LIZARD sub-Saharan Africa, this species
North and Central America. These secretive, viviparous Lepidophyma flavimaculatum forages for invertebrates and other long,
lizards are crepuscular (active at twilight) or diurnal Found among rotting logs in animals small enough to swallow. sinuous tail
the wet forests of C. America,
rather than nocturnal. They have a large headshield this species feeds on insects.
and rectangular belly scales.
ANGUID LIZARDS
The Anguidae include both legless forms and species with normal limbs.
Most lizards are surface dwellers and occur in a range of habitats in the
5 in
Old and New Worlds. Though often insectivorous, these lizards will feed
13 cm on a range of prey. They lay eggs.
19 in SLOW WORM
48 cm Anguis fragilis
AMERICAN CALIFORNIA
LEGLESS LIZARD
A legless lizard found in
well-vegetated habitats
LEGLESS LIZARDS Anniella pulchra across Europe, this
An insectivorous, wormlike species favors compost
Found only in the deserts lizard, this species burrows in heaps, where it preys
of western North America, sand or loose soil. It can detach on invertebrates.
the Anniellidae comprises its tail to trick predators.
4 ft
two species. They have a EUROPEAN GLASS LIZARD 1.2 m
long, cylindrical body and a 51/2 in Pseudopus apodus
small head, and burrow to 14 cm Found in the dry habitats of
S. Europe, this legless lizard
hunt invertebrates. They give hunts during the day for large
birth to one or two live young. insects and small lizards.

BEADED LIZARDS
Native to the arid areas of western North America, the Helodermatidae pink and black 20 in
have salivary glands modified to produce venom that is delivered along beadlike scales 50 cm
grooved teeth. Active at night, they hunt a wide range of
invertebrates and also eat carrion. GILA MONSTER
Heloderma suspectum
This lizard of the deserts of
southwestern N. America is
an egg-laying species. It hunts
insects and ground-dwelling
vertebrates, including
small mammals.
 PERENTIE spotted pattern of scales
Varanus giganteus
The largest lizard in Australia, this 393
shy species inhabits arid regions. GREEN TREE MONITOR
It has a varied diet and mildly Varanus prasinus
venomous saliva. This monitor from New Guinea

REPTILES • AMPHISBAENIANS
8 ft 30–39 in
2.5 m 75–100 cm
feeds on invertebrates in the forest
canopy. A prehensile tail and
sticky feet help it to grip branches.

YELLOW-SPOTTED
MONITOR
Varanus panoptes
strong claws for This species from Australia
61/2 ft digging out prey
2m 41/2 ft and S. New Guinea is rarely found
1.4 m far from a permanent source of
water. It preys on other reptiles.

NILE MONITOR
Varanus niloticus
The second-largest reptile
in Africa, this monitor
preys on various vertebrates
and mollusks, and also
scavenges on carrion. 10 ft
3m

skin folds
on neck

KOMODO
DRAGON
Varanus komodoensis
The world’s largest
lizard, this species is
restricted to a few
Indonesian islands. It
well-developed, grayish-brown is a formidable predator
strong limbs scaly skin of large mammals.

A M PHISBA ENIANS
They may resemble legless lizards, but They hunt for soil invertebrates by means of scent PHYLUM CHORDATA
the burrowing reptiles in the suborder and sound, killing them in their crushing jaws. CLASS REPTILIA

Amphisbaenia are quite distinct in Burrowing involves the worm-lizard propelling itself ORDER SQUAMATA

their anatomy and behavior. forward by contracting and extending its body while FAMILIES (AMPHISBAENIANS) 6

the specially adapted head pushes through the soil. SPECIES 195

The amphisbaenians, or worm-lizards, are reptiles The eyes are protected by tough, translucent skin.
wholly adapted to a subterranean lifestyle, Fertilization is internal, with some species laying
spending almost their entire lives underground. eggs, others bearing live young. They are found in
Most species have lost all trace of their limbs and South America, Florida, Africa, the Middle East,
have elongated bodies with smooth scales. and southern Europe.

WOR M-LIZ A R DS EUROPEAN WORM-LIZARD

Blanus cinereus
Rarely seen above ground, 12 in
Amphisbaenians are burrowing reptiles with a head specially 30 cm
this burrowing reptile from
adapted for digging. They are found in Europe, sub-Saharan Spain and Morocco forages in
Africa, and South America. They are formidable predators rudimentary leaf litter for invertebrates,
of soil invertebrates, hunting by eye particularly ants.
scent and sound.

black and LANG’S

white markings ROUND-HEADED
SPECKLED WORM-LIZARD
WORM-LIZARD Chirindia langi
18 in Amphisbaena fuliginosa 61/2 in Found in southern Africa, this
45 cm This worm-lizard hunts invertebrates 17 cm species burrows through sandy
while burrowing through leaf litter soils to forage for termites. If
in the rainforests of S. America. caught, it can shed its tail
It surfaces during heavy rains. in a bid to escape.
394 SNAKES
Snakes are predators with an large, muscular stomach. Instead of eyelids that PHYLUM CHORDATA
elongated body and skin made up can be opened and closed, each eye is covered by
REPTILES • SNAKES

CLASS REPTILIA

of overlapping scales. In a number a clear, protective scale. Eyesight can be good, but ORDER SQUAMATA

of species modified teeth, or fangs, varies according to lifestyle. Lacking external ears, FAMILIES (SNAKES) 30

snakes detect sounds as vibrations from the ground SPECIES 3,789

are used to deliver venom. as well as in the air. Their key sense is smell, not
Although most numerous in tropical regions, through the nostrils, but from airborne chemicals
snakes have adapted to live at colder latitudes and collected by the forked tongue. Some species can
at altitude and are found on all continents except also detect the body heat of mammals and birds.
Antarctica. They are usually terrestrial with many
living in trees, but some species burrow, some are k illing to live
semiaquatic, and others live entirely in the sea. All snakes are carnivorous. Their sharp teeth
The smallest are tiny, threadlike creatures, but curve backward to grasp and hold prey. Whether
the largest can reach 33 ft (10 m) in length. Most they eat their victim alive or kill it by injection of
are between 1–6½ ft (30 cm and 2 m). venom or by constriction, they consume their prey
whole. Their uniquely elastic jaws allow them to
a u n i q u e a r r ay o f s e n s e s swallow animals much larger than a snake’s own
Some snakes have external vestiges of hind limbs, head. Snakes defend themselves by camouflage,
but they move by muscular contractions creating warning coloration, or mimicry. When threatened,
traction between the scales on their undersides they may strike out and bite.
and the surface beneath. A large number of flexibly Species in cold places tend to bear live offspring,
linked vertebrae allows bending and coiling in any whereas those in warmer climates lay eggs.
direction. The scaly skin is regularly shed to allow Fertilization is internal. The male deposits sperm
by means of one of his paired sexual organs, If threatened, the Mojave rattlesnake uses
growth. Many snakes rely on just one elongated lung its characteristic rattle, bares its fangs, and
for breathing, while the gut is a simple tube with a known as hemipenes. coils up, ready to strike at the aggressor.

BOAS
The majority of Boidae are found in Central
and South America, but a few species of boas
are also present in Madagascar and New
Guinea. These snakes are found from forests jaws
dislocate
to marshes, where they prey on vertebrates, to swallow
killing them by constriction. This family large prey
includes the anaconda, the largest living
snake, and other species of varying sizes.
Most give birth to live young.
white COMMON BOA
zigzag Boa constrictor
striping This large boa lives in a variety
DUMERIL’S BOA of habitats in C. and S. America.
Acrantophis dumerili It is a nocturnal hunter of
Found in the humid forests of Madagascar, small mammals.
this snake becomes torpid when sheltering
in burrows during the dry, cool months.
5– 6½ ft
small, smooth 1.5–2 m COOK’S TREE BOA
granular scales Corallus cookii
Endemic to St. Vincent Island in
the Caribbean, this rare snake
hunts in trees for birds and
mammals under cover
of darkness.
EMERALD TREE BOA
Corallus caninus
Found in rainforests of S. America,
saddlelike dark this arboreal snake preys on small
markings mammals and sometimes birds. It
has a slender body, long prehensile
tail, and large head. The live young
are brick-red or yellow-orange in
color, changing to emerald green
at 12 months of age.
6 ft 5 ft
1.8 m 1.5 m
 EAST AFRICAN SAND BOA 3½ ft
Gongylophis colubrinus 1.1 m
This African snake, with a stout body and 395
short tail, waits in a burrow with only its head
exposed to catch unsuspecting prey. head with
small eyes

REPTILES • SNAKES
35 in small, smooth scales
90 cm assist in burrowing

headlike tail
used in defense
mottled brown
coloration for
camouflage
CALABAR GROUND BOA
Calabaria reinhardtii
Native to W. Africa, the Calabar ground
boa hunts in burrows for small mammals.
It is the only egg-laying boa.

RAINBOW BOA
Epicrates cenchria
Microscopic ridges
long, on the scales of this
muscular body S. American snake 6½ ft
refract light to give it an 2m
iridescent sheen. It hunts
for mammals in forests.

smooth, glossy scales 3¼ ft

1m

8 ft
2.5 m

NEW GUINEA
GROUND BOA
Candoia aspera
A sharply angled snout
32 in helps to identify this
80 cm RUBBER BOA New Guinea snake. It is a
Charina bottae slow-moving terrestrial
33 ft
Preferring cool, humid conditions, 10 m hunter of small vertebrates.
this burrowing species is found
at high altitudes and as
far north as British Columbia.

ROSY BOA
Lichanura trivirgata
This is a slow-moving
snake from the deserts
of the W. US. It ambushes
mammals and kills them
by constriction. dull yellow color
with prominent GREEN ANACONDA
black spots Eunectes murinus
This aquatic S. American
species is the largest of the
New World snakes. It hides
3¼ ft thick, muscular body in water and kills its prey
1m used to constrict prey by constriction.
396 COM MON BOA
B oa con s t r ic tor
The common boa is a large terrestrial snake from the tropics of Central
REPTILES • SNAKES

and South America. Although they are often found in woodland and
scrub, boas easily adapt to a variety of habitats. Lying motionless on the
ground, the snake ambushes its mammalian prey—grasping its victim
with its jaws, it coils around the body and kills by constriction,
tightening its coils with every exhalation of its prey. Once
dead, the prey is swallowed head first. The left and
right sides of the boa’s lower jaw separate in front,
and can open wide to take astonishingly large prey.
Usually a solitary species, males will actively seek out
a partner at breeding times, attracted by the scent emitted by the
female. Larger than males, female boas give birth to 30–50 live offspring,
each measuring about 12 in (30 cm) in length. This species is regularly
seen in captivity but its propensity to bite makes it an unpredictable pet.

size Up to 8 ft (2.5 m)
habitat Open woodland and scrubland
distribution C. and S. America
diet Mammals, birds, reptiles

COLOR VARIANTS CAMOUFLAGE

There are numerous subspecies of Pigmentation of some scales BELLY SCALES
common boa, which are defined forms patches of color that The robust belly scales grip most
on the basis of size and color disrupt the outline of the snake. surfaces, allowing the common boa
patterns. Color also varies This helps with camouflage when to pull itself along and climb trees.
considerably between localities. hunting prey or evading predators. It periodically sheds its skin.

distinctive,
SIZEABLE FOE saddlelike
The common boa hunts by sight and smell, markings
so the eyes and nostrils are prominent on the
triangular head. Feeding can be infrequent—
a small prey item may last a snake two
to three weeks. A large snake that kills
a deer will not need to feed for
at least six months.
 397

even when closed,

the mouth has an forked tongue
opening that allows EYE
the tongue to be The boa has keen eyesight,
extended and the pupil narrows to
a slit in bright light. It has
no eyelids. Around the
eyes, the head is covered
with numerous small
scales, which are made
of keratin.

REMNANT LIMBS
All snakes lack limbs, but
more primitive species like
the common boa still possess
the remnants of a pelvis, as
well as tiny, spurlike hind
limbs on each side of the
anal vent—these once
would have been legs.

TONGUE FLICKING
The forked tongue is an important
sense organ. Snakes flick their tongues,
collecting chemicals from the air; these
are then analyzed by the Jacobson’s
organ in the roof of the mouth.
 COLUBRIDS COMMON GARTER
SNAKE
398 With about 2,000 species worldwide, colubrids form the largest family
Thamnophis sirtalis
This N. American species is
of snakes. The members of the family Colubridae inhabit a wide range active during daytime,
of habitats, from deserts to wetlands. They have a varied diet. hunting vertebrates in a range
REPTILES • SNAKES

Many species lay eggs. of habitats. In Manitoba, after

41/4 ft hibernation it congregates in
1.3 m large numbers to mate.
WESTERN LYRE SNAKE
Trimorphodon biscutatus large eye
banded This secretive, nocturnal
coloration species of western 18 in
N. America is found in 46 cm
rocky habitats. It hunts bats
and other small mammals, 35 in
as well as lizards. 90 cm

4 ft
1.2 m

RUTHVEN’S KINGSNAKE
Lampropeltis ruthveni
An egg-laying species, GRAY EARTHSNAKE
this colubrid is from the Mexican Geophis brachycephalus
plateau, where it hunts rodents This small, C. American snake
and lizards in dry woodland. is terrestrial and nocturnal. It
feeds mainly on earthworms
and soft insect larvae.

31/4 ft
1m LONG-NOSED SNAKE
Rhinocheilus lecontei 31/2 ft
1.1 m
Characterized by a pointed snout,
this shy, nocturnal, burrowing
species is from dry grasslands
of N. America, where it CALIFORNIA
hunts lizards. MOUNTAIN KINGSNAKE
Lampropeltis zonata
This secretive snake prefers
high-altitude wooded habitats.
It is typically nocturnal, but is
diurnal when nights are cold.

4 ft
1.2 m
blotched, brown
coloration
AFRICAN EGG-EATING SNAKE
Dasypeltis scabra
This African snake has a highly
specialized diet of eggs on which it
feasts during the bird-nesting season.
It then fasts for the rest of the year.

long, 91/4 ft
powerful body 2.8 m

EASTERN PINE SNAKE

Pituophis melanoleucus
When threatened, this large,
powerful snake of woodlands in
N. America ejects foul-smelling waste
from its rear opening (cloaca).
 WESTERN INDIGO
SNAKE
31/4 ft
1m Drymarchon corais 4 ft 399
This is one of the longest 1.2 m
long, snakes in N. America.
slender body It often shares a burrow

REPTILES • SNAKES
with the gopher tortoise.

41/2 ft 10 ft
1.4 m 3m BANDED FLYING SNAKE
Chrysopelea pelias
GREEN Despite its name, this S. Asian snake
HIGHLAND RACER
Drymobius chloroticus does not actively fly. Instead, it glides
down from high branches by making its
This fast, agile snake is from underside concave.
the rainforests of C. America.
It is usually found near water, NORTHERN WATER SNAKE
where it feeds on frogs. Nerodia sipedon
This aquatic, viviparous species from 26 in 6 ft
eastern N. America is active throughout 65 cm 1.8 m
the day and the night. It feeds on
amphibians and fishes.

short
snout
MOELLENDORFF’S RATSNAKE
Elaphe moellendorffi
This species is from dry limestone
regions of China and Vietnam. It has an
FALSE CORAL SNAKE elongated snout and a relatively long tail.
7 ft Erythrolamprus mimus
2.1 m The bright coloration of the
harmless false coral snake from
S. America is similar to that
of highly poisonous
coral snakes.
MUD SNAKE
Farancia abacura REDBACK
This N. American striking red, white, COFFEE SNAKE
snake preys on aquatic
31/4 ft and black bands Ninia sebae
99 cm
salamanders, grasping them This harmless snake
with strongly curved teeth. from C. America
41/4 ft has an ability to extend 16 in
Females remain coiled 1.3 m 40 cm
around their eggs until its neck as a form
they hatch. of intimidation.

31/4 ft
1m
NORTHERN
CAT-EYED SNAKE BROWN BLUNT-HEADED BROWN TREESNAKE
Leptodeira septentrionalis VINE SNAKE Boiga irregularis
This nocturnal, arboreal species Imantodes cenchoa Originally from Australia and New
from C. America has large eyes The large eyes of this slender snake help Guinea, this snake was accidentially
to help it hunt for vertebrates it hunt lizards in the dark. It is found in introduced on the island of Guam
and eggs of tree frogs. tropical American rainforests. in the W. Pacific, where it has
decimated the native fauna.
10 ft
3m

FALSE WATER
COBRA
Hydrodynastes gigas
Native to S. American
rainforests, this is 61/2 ft
a semiaquatic snake. GREEN VINE SNAKE 2m
Like cobras, it can flatten Oxybelis fulgidus
its neck to appear Long and delicate, this arboreal snake is from
more intimidating. C. and S. American rainforests. It holds its prey in
the air until its venom immobilizes the victims.

31/2 ft characteristic
1.1 m yellow collar

GRASS SNAKE
Natrix natrix
Widely distributed
throughout Europe,
CALICO SNAKE this species often feigns
Oxyrhopus petolarius 4 ft death when threatened.
This species feeds on lizards and other small 1.2 m At home in water,
vertebrates. It is a terrestrial and diurnal olive-gray body it regularly feeds
inhabitant of S. American rainforests. color on amphibians.
 COLUBRIDS 5¼ ft 7¾ ft RED-TAILED
GREEN RATSNAKE
1.6 m 2.4 m
400 Gonyosoma
oxycephalum
This fast-moving snake
hunts for birds and
REPTILES • SNAKES

mammals in trees. It is
found in the rainforests
of S.E. Asia.
4½ ft
1.4 m long, slender
body
COMMON TRINKET ROUGH GREEN SNAKE
SNAKE Opheodrys aestivus
Coelognathus helena Native to woodlands in southeastern
This Indian species enlarges its N. America, this arboreal snake
neck and rises up to intimidate is a diurnal predator of insects.
its enemies. It hunts mammals, It is oviparous. 3¼ in 4½ ft 5¼ ft
usually at night. 1m 1.4 m 1.6 m
23½ in
60 cm

SMOOTH SNAKE
Coronella austriaca
This secretive European snake is found on
heaths and kills its prey by constriction. Females
produce eggs that hatch within their bodies.
SOUTHERN
BALKAN WHIPSNAKE DAHL’S WHIPSNAKE WATER SNAKE
Hierophis gemonensis Platyceps najadum Nerodia fasciata
Restricted to the Balkans, this From the Mediterranean region, Living in wetlands, this
snake is found in dry scrubland this snake is found in dry, stony colubrid preys on amphibians
and olive groves. It hunts during habitats. It hunts small lizards and and fishes. It is found in
BROWN-HEADED 8 in the day, preying on lizards. grasshoppers during the day. the S. US.
CENTIPEDE SNAKE 20 cm head with
Tantilla ruficeps prominent eyes brown markings
This burrowing snake on muscular body
is found in the tropical forests pale underside
of C. America. It is a mainly
diurnal insect-eater. 32 in 6 ft 6 ft 6½ ft
80 cm 1.8 m 1.8 m 2m
WESTERN
HOGNOSED SNAKE
Heterodon nasicus
This snake of the N. RED CORNSNAKE YELLOW RATSNAKE
American prairies uses its DIADEM SNAKE Pantherophis guttatus Spilotes pullatus
specialized, enlarged teeth Spalaerosophis diadema cliffordi This snake is common in A predator of small vertebrates,
to puncture the lungs of Native to deserts in N. Africa, this subspecies southeastern N. America, but this large snake is rarely found far
toads, making them is diurnal during cooler months, but turns is rarely seen. It hunts small from water. It is widely distributed
easier to swallow. nocturnal during summer months. mammals in woodland. in S. and C. America.

L A M PROPH I I DA E ASIAN PIPE SNAKES

Found mainly in Africa, the diverse lamprophiid Found in Sri Lanka and S.E. Asia, these small, burrowing snakes of the
snakes occupy varied habitats—terrestrial, Cylindrophiidae have a cylindrical body and smooth, shiny scales. They
arboreal, and semiaquatic. Predating a range live in wet habitats, sheltering in tunnels, but emerge at night to hunt
of vertebrates and invertebrates, they subdue other snakes and eels. They give birth to live young.
their prey by either constriction or venom.
CEYLONESE PIPE SNAKE 26 in
Cylindrophis maculatus 65 cm
MOLE VIPER Endemic to Sri Lanka, this harmless
Atractaspis fallax burrowing snake eats invertebrates. It
This poisonous snake from uses its flattened tail to mimic the
E. Africa has large front fangs. movements of the venomous cobra.
It hunts other burrowing
vertebrates underground. 30 in
75 cm
SEA SNAKES YELLOW-LIPPED
SEAKRAIT
These highly venomous snakes Laticauda colubrina
from the tropical coastal waters This snake hunts fishes
at night in tropical Indo-Pacific
of the Indian and Pacific oceans seas.
6½ ft 6 ft It can move on land as well.
2m 1.8 m belong to the subfamily
Hydrophiinae, part of the larger 4½ ft
1.4 m
GIANT MALAGASY Elapidae. They are well adapted
MONTPELLIER SNAKE HOGNOSE SNAKE for swimming, with a paddle-
Malpolon monspessulanus Leioheterodon madagascariensis
Inhabiting dry scrub and rocky hillsides This large, diurnal snake, with an upturned
like tail. All but a few breed in
around the Mediterranean, this long, slender snout, hunts lizards and amphibians in water, bearing live young. They
snake hunts small vertebrates during the day. grasslands and forests in Madagascar. eat eels and other fishes.
COBR A S A N D R E L AT I V E S
Widely distributed in the tropics, the venomous snakes of the small, slender head
401
32 in
Elapidae have short, permanently erect fangs at the front of the 80 cm
mouth. Varied in body forms, they occupy a wide variety of

REPTILES • SNAKES
habitats. Some species lay eggs, while others bear live young.

CENTRAL AMERICAN
CORAL SNAKE
Micrurus nigrocinctus
This venomous snake of C. America forages in leaf
litter of tropical forests. Its bright coloration
serves as a warning to potential aggressors.
4 ft
1.2 m 26 in
YELLOW-FACED WHIP SNAKE 65 cm
Demansia psammophis
Widely distributed across Australia, this
slender snake is an active diurnal hunter EASTERN SHIELD-NOSE ROSEN’S SNAKE
of lizards. It prefers dry, open habitats. SNAKE Suta fasciata
30 in Aspidelaps scutatus fulafulus Found in arid areas
75 cm This nocturnal predator of small of Western Australia,
spread lizards and mammals is found
hood acts this venomous snake
as warning in southern African savanna. hunts for lizards.
It burrows in sandy soil.

20 in 14 in
50 cm 35 cm

28 in
70 cm

RINGED BROWN SNAKE SOUTHERN DESERT

Pseudonaja modesta BANDED SNAKE
Found in dry, rocky areas in Simoselaps bertholdi
Australia, this venomous snake Widespread in Western Australia, this
feeds on small skinks. Its survival is small snake burrows in search of its lizard
DESERT threatened by the loss of its habitat. prey. The banding may confuse predators.
DEATH ADDER
Acanthophis pyrrhus coloration can vary
This predator of the western deserts depending on region
of Australia lures its prey of small
lizards and mammals by wiggling
its tail. It then ambushes them.

6½ ft 7¾ ft 16 ft
2m 2.4 m 5m

MONOCLED
COBRA
Naja kaouthia
Common in S.E. Asia, EGYPTIAN COBRA KING COBRA
this large snake hunts Naja haje Ophiophagus hannah
rats and other snakes in This large cobra hunts small A forest species of tropical Asia, the
woodland and paddy vertebrates in the deserts of massive king cobra primarily preys on
fields, often close to N. and C. Africa. When threatened, it other snakes. Unusually for snakes, its
human habitation. spreads its hood and raises its head. nest of eggs is defended by both sexes. 30 in
75 cm
smooth scales
on brown body

RED SPITTING COBRA

Naja pallida
This African species not only
raises its hood when threatened,
but also sprays streams of venom
into the face of its aggressor.
 PYTHONS
402 Members of the family Pythonidae 5 ft
can be found across Africa, Asia, and 1.5 m
Australia. Pythons detect warm- GREEN TREE PYTHON
REPTILES • SNAKES

blooded prey with their facial pits. Morelia viridis

They use their mouth to grasp prey Found in the tropical forests of
but kill by constriction. Some Australasia, this arboreal snake spends
its time looped over a branch, ready
species coil around their eggs to to ambush lizards and small mammals.
help incubate them using the heat bright green color
generated by their shivering. aids in camouflage

BLOOD PYTHON
Python curtus
This relatively short
python of S.E. Asian
rainforests coils around 6 ft
its clutch of eggs to 1.8 m
incubate and protect them.

BURMESE PYTHON
10 ft
Python molurus
3m Rare in its native homeland,
this Asian snake is becoming
superabundant in the Florida
BLACK-HEADED Everglades. Its broad diet of
PYTHON birds, mammals, and reptiles is
Aspidites melanocephalus a threat to many local animals.
Endemic to Australia, this
species is found in a variety
of habitats. It preys upon 23 ft
other snakes and reptiles. 7m

BLIND SNAKES VIPERS

The Typhlopidae have eyes covered The Viperidae are characterized by a
with scales, rendering them effectively thick body, keeled scales, and a triangular
blind. Found in leaf litter of tropical head. Long, tubular, hinged fangs at the
forests, these small, burrowing snakes front of the mouth efficiently inject
mainly eat soil invertebrates and have venom into their vertebrate prey.
teeth only on their upper jaw. Most Pit vipers have sensory 5 ft
of them lay eggs. pits between their eyes 1.5 m
and nostrils that detect
heat. Most vipers bear
live young. FER-DE-LANCE
14 in Bothrops atrox
35 cm Found in the forests
30 in of tropical America,
75 cm EURASIAN this venomous snake is
BLIND SNAKE characterized by a sharply
BLACKISH BLIND SNAKE Xerotyphlops vermicularis pointed head. It hunts birds
Anilios nigrescens This wormlike blind snake 34 in and mammals by night.
85 cm
Tough scales allow this burrowing snake from Europe inhabits dry, open
from E. Australia to resist attack from areas. It burrows to feed on
ants while eating their eggs and larvae. invertebrates, mainly ant larvae.

THREAD SNAKES
These small, slender snakes are rarely seen because of their
habit of burrowing to hunt invertebrates. Belonging to the
Leptotyphlopidae, they are found in the tropics of the 23½ in
60 cm
Americas, Africa, and southwest Asia.

12 in SENEGAL BLIND SNAKE DESERT HORNED VIPER HOGNOSE PIT VIPER

30 cm Myriopholis rouxestevae Cerastes cerastes Porthidium nasutum
This snake inhabits the soil of tropical forests Occurring in the deserts of N. Africa and This diurnal hunter of C. America prefers humid, open
in western Africa. First studied in 2004, the Sinai Peninsula, this snake lies buried in forests. Pit vipers, which have heat-detecting sensory
it is believed to eat invertebrates. sand to ambush small mammals and lizards. pits between their eyes and nostrils, bear live young.
 SUNBEAM
blotched SNAKES 403
markings
4½ ft Characterized by iridescent
1.4 m scales, the Xenopeltidae

REPTILES • SNAKES
include only two species,
both restricted to Southeast
Asia. They live in burrows
in forest habitats and hunt
SPOTTED PYTHON amphibians, other reptiles,
Antaresia maculosa and small mammals. They
This python is found on the rocky hillsides of reproduce by laying eggs.
N. Australia. It feeds on bats by catching them
at the entrance of their roosting caves.

33 ft 4¼ ft
10 m 1.3 m

netlike pattern

powerful, muscular
body useful for RETICULATED PYTHON SUNBEAM SNAKE
constricting prey Python reticulatus Xenopeltis unicolor
One of the longest snakes in the world, this A flat head helps this snake burrow into
species inhabits Asian rainforests. It kills decaying vegetation. It emerges at dusk to
large mammals by constriction. feed on amphibians and small mammals.

23½ in 4 ft
60 cm 1.2 m

4¼ ft
1.3 m
35 in
COPPERHEAD 90 cm
Agkistrodon contortrix
Banded markings help this
snake to hide in leaf litter in
rocky woodland. It is found COMMON ADDER
in eastern N. America. Vipera berus
This diurnal species hunts small mammals
and lizards. It is widely distributed across
a variety of habitats in Eurasia. short, dark-
ASP VIPER edged blotches
Vipera aspis
This European viper prefers warm,
dry habitats, where it feeds on small 7 ft
mammals. It bears up to 20 live young. 2.1 m

WEST AFRICAN WESTERN

GABOON VIPER DIAMONDBACK
Bitis rhinoceros RATTLESNAKE
6½ ft Crotalus atrox
2m Occurring in tropical
rainforests in Africa, this A nocturnal hunter of PRAIRIE RATTLESNAKE
large, heavily built snake mammals, this large snake is Crotalus viridis
preys on mammals by widespread in arid areas of This snake from the midwestern
ambushing them. western N. America. US hunts mammals at
dawn and dusk. It hides in
crevices during the day.

6 ft
1.8 m

large head
PUFF ADDER
Bitis arietans MALAYAN PIT VIPER
Most active at night, this venomous Calloselasma rhodostoma
snake usually lies still to ambush its Occurring in S.E. Asia, this species
vertebrate prey. It is found in 3¼ ft forages by night in open areas adjacent
rocky grassland in Africa. 1m to forests for rodents and lizards.
404 C RO C O D I L E S A N D A L L I G AT O R S
Crocodilians are large, carnivorous, ambush specific prey, such as migrating mammals PHYLUM CHORDATA
aquatic reptiles. Their armored and fishes. Small prey is swallowed whole but large
REPTILES • CROCODILES AND ALLIGATORS

CLASS REPTILIA

skin and powerful jaws make them prey is first drowned before the crocodilian rolls ORDER CROCODYLIA

formidable predators, yet they are over while grasping the carcass to twist off pieces FAMILIES 3

of meat. Digestion is aided by stones in the stomach SPECIES 25

sociable animals and caring parents. and highly acidic gastric secretions.
All crocodilians—crocodiles, alligators, and
gharials—have a similar body plan: an elongated s o c i a l b e h av i o r a n d b r e e d i n g
snout armed with numerous, sharp, unspecialized Behaviorally, crocodilians have more in common
teeth, a streamlined body, a long, muscular tail, with birds, their closest living relatives, than with
and skin armored with bony plates. The saltwater other reptiles. Adults form loose groupings—
crocodile is the largest living reptile. particularly at good feeding sites—and use a
wide range of vocalizations and body language
sw i m m i ng an d feedi ng to interact with each other.
Crocodilians are found in tropical regions In the breeding season, dominant males
worldwide, inhabiting a variety of freshwater control territories and actively court females.
and marine habitats. The eyes, ears, and nostrils are Fertilization is internal and clutches of rigid-
on top of the head, which enables the animal to be shelled eggs are laid in a nest constructed and
almost completely submerged while hunting. The guarded by the female. The sex of individuals
powerful tail is used for swimming and strong legs is determined by the incubation temperature.
allow crocodilians to move readily on land with Hatchlings call to stimulate the female to excavate
the body raised well above the ground. the nest and carry them in her mouth to water.
Although generalist carnivores, feeding on fishes, Mortality is high in young crocodilians but once
Exploiting a fast-flowing stream,
reptiles, birds, and mammals, crocodilians exhibit they are longer than 31/4 ft (1 m), they have few these Yacare caimans lie in wait with
sophisticated feeding behaviors to locate and natural enemies. their mouth held open, waiting for a fish
to come within reach.

GHARIALS olive-green body

The endangered Gavialidae 23 ft
are restricted to India. 7m
The slender snout has
rows of numerous sharp
teeth that help to catch GHARIAL
Gavialis gangeticus
fishes. Males develop a Among the largest of the Crocodylia, this Asian
wartlike growth on the species has a fearsome array of teeth, which help it
tip of the snout. to catch fishes, but it is not known to attack humans.

C ROCODI L E S
Relatively unspecialized in lifestyle, habitat, and diet, the
Crocodylidae are most easily recognized by the exposure 6½ ft
of the fourth tooth of the lower jaw, when the jaws 2m
are closed. These tropical reptiles occupy many
habitats around rivers and coasts. DWARF CROCODILE
Osteolaemus tetraspis
From the forests of tropical
Africa, this small crocodile has
a heavy armor of scales on its
neck and back. It is a nocturnal
hunter of fishes and frogs.

CUBAN CROCODILE
Crocodylus rhombifer 11 ft
3.5 m
Endemic to Cuba, this medium-sized crocodile
inhabits swamps, and hunts fishes and small mammals.
It lays its eggs in holes in the ground.

13 ft SIAMESE CROCODILE
4m Crocodylus siamensis
Restricted to S.E. Asia, this
large crocodile is critically
endangered in the wild. Found
in freshwater marshes, it feeds powerful tail propels
on a variety of prey. crocodile through water
A L L IGATOR S coloration
AND CAIMANS darkens with age 405
Members of the family Alligatoridae feed on fishes,
birds, and mammals that share their aquatic habitat.

REPTILES • CROCODILES AND ALLIGATORS

These reptiles are distributed throughout freshwater 16 ft
swamps and rivers of the tropical and subtropical 5m
Americas. The only species living outside the
Americas is the rare Chinese alligator. AMERICAN ALLIGATOR strong legs allow
Alligator mississippiensis smooth movement
Conservation efforts have now made on land
this N. American species quite
common. It preys on birds, small
mammals, and turtles.

8¼ ft
2.5 m
CHINESE ALLIGATOR 6½ ft
Alligator sinensis 2m
This critically endangered species of the Yangtze
SPECTACLED CAIMAN valley of China lies dormant in burrows during
Caiman crocodilus the cold winter months.
This caiman feeds on a wide range of prey. Widespread in
C. and S. America, it appears to be the only crocodilian
that readily occupies man-made aquatic habitats.
rounded head
with broad
snout
10 ft
3m

BROAD-SNOUTED CAIMAN
Caiman latirostris mottled pattern
A mound-nesting species found across much
of central S. America, this caiman is characterized
by a broad snout. It hunts mammals and birds. 5½ ft
bony armor 1.7 m

SCHNEIDER’S
DWARF CAIMAN
Paleosuchus trigonatus
CUVIER’S DWARF CAIMAN This small crocodile of
Paleosuchus palpebrosus 5 ft S. American rainforests is
This S. American species is the 1.5 m semiterrestrial. It builds
smallest of the New World members its nest against a termite
of this order. It has a doglike skull mound, which keeps
and bony-armored skin. the eggs warm.

SALTWATER CROCODILE
Crocodylus porosus
The largest of all living reptiles, this
common Indo-Pacific species readily
crosses the open ocean. Its diet is
wide-ranging and unspecialized.
23 ft
7m eyes set high on head
greenish brown body,
often covered in algae

16 ft
5m

NILE CROCODILE
Crocodylus niloticus
Widely distributed in Africa,
this large crocodile typically lives in
fresh water but has been seen around
coasts as well. Its diet varies with
age, with adults taking larger prey.
406 C U BA N C RO C O D I L E
Crocodylu s rhom bifer
This strikingly colored crocodile is native to Cuba.
It is medium-sized, robust, and armored with bony scales. More terrestrial than other
crocodilian species, it raises its belly off the ground and adopts a “high walk” posture
when moving on land. Its preferred prey are turtles, which are crushed by the strong
teeth at the rear of its mouth. Hunting pressure and habitat loss meant that, in the
1960s, the few remaining wild Cuban crocodiles were captured and released into
size 9¾–11 ft (3–3.5 m)
Zapata Swamp, a reserve in the south of the island. Although protected, the population habitat Freshwater swamps
remains small, and hybridization with American crocodiles on the reserve threatens distribution Cuba
the purity of this species. diet Fishes, turtles, small mammals

large scales armored

with bony deposits
bony projections at external ear flap,
back of the head which is closed while the
resemble horns crocodile is under water
 MUSCULAR LIMBS NOSTRILS
This crocodile can race over short Paired nostrils are located on 407
distances, powered by its strong rear the nasal disk, an elevated pad
legs. Because they are used for walking of tissue at the end of the snout.
rather than swimming, the feet have Under water, the nostrils can
no webbing between the toes. be closed by valves.

REPTILES • CROCODILES AND ALLIGATORS

FORMIDABLE JAWS
Crocodiles cannot chew their
food, but they use their powerful
jaws to grasp their prey. Sensitive
taste buds on the tongue allow
the crocodile to reject unpleasant
tasting objects. Evaporation from
the mouth cools the body.

TAIL RIDGE
Scales with bony projections
increase the depth of the BELLY SCALES
tail—useful when the The scales on the crocodile’s
crocodile is swimming. Rich underside are small and uniform
in blood vessels, they also help in size and pattern. These scales
to absorb heat when basking. are prized by the leather trade.

scales along the snout have sensory nostrils

papillae, which are sensitive to
vibrations in the water

fourth lower
tooth visible
when jaws
closed

sharp front teeth

STEALTH ATTACK
Sharp eyes and sensitive ears are located
on the top of the head, so that prey can
blunt back teeth be detected while the body is submerged.
are more robust than This enables the crocodile to take its
the front teeth victims by surprise. The long lower jaw
can be closed with tremendous force
to grasp struggling prey securely.
408 BIRDS
Birds have busy, active lives. Many are spectacularly beautiful;
ANIMALS • BIRDS

others sing complex—even musical—songs. They have the

intelligence and parental devotion of a mammal combined
with characteristics more reminiscent of a reptilian ancestor.

PHYLUM CHORDATA Birds are the only living animals with feathers. They
CLASS AVES are warm-blooded vertebrates that stand on two legs
ORDERS 40 and their forelimbs are considerably modified as wings.
FAMILIES 250
SPECIES 10,770
As well as facilitating flight, feathers insulate the body,
so many birds stay active in icy-cold conditions, just
like furry mammals. Feathers can also be colorful,
showing that birds rely on flaunting visual cues to
communicate and find a mate. Birds evolved from
a group of two-legged, carnivorous dinosaurs that
included Tyrannosaurus. It is possible—even likely—
that these ancestors were already feathered.

ta k i n g t o t h e a i r
Asymmetrical flight feathers, with No one knows for sure how—or why—the first
a narrow outer blade and broader inner birds took to the air, but the fact that they did had
blade, give birds better control in the air.
effects that would be forever imprinted upon the
bird’s body. Their wristbones became fused together
as arms evolved into wings. The breastbone evolved
a huge keel for the attachment of larger muscles for
powering the flapping wings. Already their dinosaur
ancestors had air-filled, but strong bones. But birds
coupled this with a strong cardiovascular system
powered by a mammallike four-chambered heart
Many chicks are altricial—born in and a rapid metabolism to generate lots of energy.
a helpless state. Blind and naked, they
require prolonged parental care. A system of air sacs—linked to the bone chambers—
expanded their breathing capacity to flush stale
air out of lungs more efficiently than in mammals.
Many reptilian features still remain. The bare
parts of a bird’s legs and toes have horny reptilian
scales. Birds excrete a semisolid waste product called
uric acid (not a solution of urea, as in mammals).
Waste from the kidneys and gut mix and exit via
a common reptilian opening called a cloaca. But a
Elaborate nests are built by the male
weaver bird to impress prospective mates, bird’s brain is better developed than that of a reptile,
testament to the brain power of birds. because it is supported by a warm-blooded body and
fast metabolism. This makes birds not only skilled at
D E B AT E flying, but remarkably sophisticated in the ways they
ARE BIRDS DINOSAURS? acquire food or raise a family. Young birds hatch
from reptilian hard-shelled eggs, but they are reared
Traditionally birds are in a class
separate from reptiles, including
by intelligent parents who will invest much time and
dinosaurs. But today scientists believe energy into raising them to maturity. Sixty million
that all descendants of a common years of evolution mean that a bird is no longer just
ancestor should be classified together a reptile with feathers.
to reflect evolutionary relationships—
a method known as cladistics. This
makes birds as much a part of the CROWNING GLORY
dinosaur group as Tyrannosaurus. Birds’ remarkable colors and structures often appear overly
extravagant, such as this pigeon’s elaborate head crest.
b i r d g ro u p s
The 40 orders range from heavy-bodied ground- dive-bombing falcons, and hovering hummingbirds.
dwellers—a few of which are incapable of flight— They exploit habitats from deep sea to mountain
to masters of the air, such as soaring albatrosses, peaks, from deserts to rainforests and wetlands.
t i na m o u s b i r d s o f pr e y m e si t e s

410 434 470

r at i t e s fa l c on s a n d nigh tja rs
c a r ac a r a s

410 441 471

f ow l , bu s ta r d s h u m m i ngbi r ds
ga m e bi r ds, and swifts
a n d r e l at i v e s

412 442 473

wat e r f ow l cranes, rails, t ro g on s
a n d r e l at i v e s

416 443 476

pe n g u i n s k ag u a n d mouse bi r ds
su n bi t t e rn

420 445 476

l o on s wa d e r s , g u l l s , k i ngfish ers
a n d au k s a n d r e l at i v e s

424 448 477

a l bat ro s s e s , sa n d g ro u s e c u c ko o - ro l l e r
pe t r e l s , a n d
s h e a rwat e r s

425 456 479

grebes pi g e on s a n d horn bi l ls,
d ov e s hoopoes, an d
wo o d h o o p o e s

427 457 482

f l a m i ngos pa r ro t s a n d wo o dpe c k e r s
c o c k at o o s an d toucans

428 460 483

stor k s t u r ac o s serie m as

429 464 487

i bises, bi t t e rns, h oat z i n pa s s e r i n e s
h e ron s , a n d
pe l i c a n s

430 465 488

cor mor ants, c u c ko o s
gan nets, and
r e l at i v e s

433 465
t ro pi c b i r d s ow l s

433 467
410 TINA MOUS R AT I T E S
The ground-dwelling tinamous from Central and A group that includes the largest living birds, the
South America resemble Old World partridges, but ratites are flightless and strong-footed. Some flock
BIRDS • TINAMOUS

they are most closely related to ratites. in open habitats; others live alone in forests.
Tinamous, the sole family in PHYLUM CHORDATA
Ratites evolved in the southern PHYLUM CHORDATA
the order Tinamiformes, are CLASS AVES hemisphere and there is CLASS AVES
small, ground-dwelling birds ORDER TINAMIFORMES evidence to suggest they have ORDERS 4
with a rounded body and FAMILIES 1 flying ancestors. They lack FAMILIES 4

short legs. All have a very SPECIES 47 some of the features associated SPECIES 13

short tail, which gives them a with flight, but retain others.
dumpy appearance, and some species are crested. Ratites do not have the bony ridge on their breastbone that anchors
Unlike the ratites, tinamous have a ridged breastbone for the powerful flight muscles in other birds, but they still have wings, and
attachment of flight muscles, a feature that is seen in all other the part of the brain that controls flight is well developed.
birds. Their wings are much better developed than those of The ostriches, which make up the order Struthioniformes, are
the ratites, and they can fly, although only over short distances. found in dry open areas in Africa. They are powerful-limbed birds
Usually they are reluctant to take to the air and prefer instead that can run at high speed on flat ground. Somewhat similar in
to run away from predators. Tinamous have a comparatively appearance, though less massively built, are the rheas—the order
small heart and lungs, which perhaps explains why these birds Rheiformes—from the grasslands of South America. Like most
become exhausted so easily. ratites, both these groups of birds have a reduced number of toes:
Some groups of tinamous inhabit woodlands and forest, while the rhea has three toes, the ostrich only two. Both have large wings
others are found in open grassland. All feed on seeds, fruit, insects, that they use in displays or to assist their balance when sprinting.
and sometimes small vertebrates. The cryptic plumage patterns
of most species makes them difficult to spot in the wild. They are au s t r a l a s i a n s p e c i e s
more likely to be recognized by their distinctive call. The Australasian ratites include the cassowaries and emus of the
order Casuariiformes and the kiwis, the Apterygiformes. All
b e au t i f u l e g g s are forest birds, save the emu, which inhabits scrubland and
Male tinamous mate with many different females, and it is the grassland. The tiny wings of these ratites are invisible under their
males that tend the eggs and young. The nests are made in leaf plumage, which resembles shaggy hair rather than feathers. The
litter on the ground. Tinamou eggs—brightly colored turquoise, nocturnal kiwis, familiar as the national symbol of New Zealand,
red, or purple—have a porcelainlike gloss. are the only ratites that do not have a reduced number of toes.

D E B AT E
ORIGINS OF TINAMOUS
16 in
40 cm
Traditionally, tinamous are grouped
separately from the flightless ratites.
But the two groups share a pattern
of skull bones not found in other
birds, suggesting that they have
a common ancestor. Scientists
are not certain whether this 20–26 in
50–65 cm
ancestor was flightless or able The male ostrich scrapes out a shallow
to fly; the latter is more likely. nest in the ground, in which his harem of
females lay up to 50 eggs.

NORTH ISLAND KIWI

furlike plumage Apteryx mantelli
ELEGANT CRESTED TINAMOU f: Apterygidae
Eudromia elegans One of several New Zealand kiwis,
12–14 in
31–35 cm f: Tinamidae this nocturnal ratite detects buried
This bird of high-altitude shrubland 26–28 in invertebrate prey by smell, using
occurs from S. Chile to Argentina. 65–70 cm nostrils at its probing bill tip.
Unlike other tinamou species, it
often lives in flocks.

TOKOEKA
ORNATE TINAMOU Apteryx australis
Nothoprocta ornata f: Apterygidae
f: Tinamidae This is the South Island cousin
An Andean tinamou, this species is of A. mantelli. Lighter in color
found on the high montane grasslands than its relative, it has been
of western S. America from Peru recognized as a separate
to N. Argentina. species after DNA analysis.
 helmetlike casque
coarse, shaggy
plumage 411

BIR DS • R ATITES
brown plumage
51/2–7 ft 3–31/4 ft
1.7–2.1 m 92–100 cm
red wattles

5–6 ft
1.5–1.8 m almost
EMU
Dromaius novaehollandiae bare neck
SOUTHERN f: Dromaiidae LESSER RHEA
CASSOWARY Australia’s largest living bird is found Rhea pennata
Casuarius casuarius across the continent in open grassy f: Rheidae
f: Casuariidae areas. Like the related cassowary, its Smaller than the greater rhea, its
Cassowaries are fruit-eating chicks are striped for camouflage. northern cousin, this ratite inhabits
rainforest birds. With a range that black body
plumage the S. Andes and Patagonia, where
extends through New Guinea and it lives in small flocks.
N. Australia, this is the most
widespread species of its family.

41/4–41/2 ft
1.3–1.4 m

white primary
feathers

GREATER RHEA
Rhea americana
large eye
f: Rheidae
This central S. American ratite
OSTRICH has a breeding system that does
Struthio camelus not involve pair-bonding. Males
51/2–83/4 ft f: Struthionidae tend a large nest of eggs
1.7–2.7 m The largest birds in the world, produced by many females.
ostriches live in savanna and
semidesert areas of Africa. gray neck
Breeding males mate with
26–28 in several females.
65–70 cm 51/2–83/4 ft
1.7–2.7 m
grayish-brown
plumage
GREAT
SPOTTED scaly legs
KIWI
Apteryx haastii
f: Apterygidae
One of two species of kiwis
with gray mottled plumage,
this bird is confined to
mountain areas of western
South Island.
featherless
thighs
LITTLE
14–18 in SPOTTED
35–45 cm KIWI
Apteryx owenii
f: Apterygidae
Introduced mammals SOMALI OSTRICH
have driven this small Struthio molybdophanes
kiwi to the edge of f: Struthionidae
extinction. It survives Separated from other populations of
on predator-free ostrich by the Rift Valley, this E. African
satellite islands around gray-necked form is now regarded as
New Zealand. two-toed feet a separate species.
412 F OW L , G A M E B I R D S ,
A N D R E L AT I V E S
BIR DS • FOWL , GA ME BIRDS, AND REL ATIVES

Strong footed game birds are adapted by decomposition or volcanic action to incubate PHYLUM CHORDATA
to a wide range of habitats. They are them. All other game birds take a more active CLASS AVES

primarily ground-dwellers and feed role in parenthood, although only the female ORDER GALLIFORMES

mostly on plants. raises the brood. The precocious offspring can FAMILIES 5

run and feed soon after hatching. SPECIES 300

Although they may be good fliers, the majority Male game birds often have flamboyant plumage,
of game birds, the order Galliformes, take to which they use in courtship displays to attract a D E B AT E
the air only when threatened. None, except the harem of females. Some species are armed with leg W H AT I S A P A R T R I D G E ?
common and Japanese quails, sustains a long spurs to battle with competitors.
migratory flight. Most game birds spend their Traditional categories of birds may not
lives on the ground. Only the members of the d o m e s t i c at e d s p e c i e s reflect biological relationships. Many
The relative ease with which many game bird game birds are called partridges, but
curassow family of tropical America are habitual
the European gray partridge differs
tree-dwellers; these birds make their nests in species, including the turkey, can be kept in subtly from others in that, for example,
trees, too. Most primitive of the game bird group captivity has meant that they are economically males are more distinctively
are the big-footed megapodes of the Indo-Pacific important in many countries. Domesticated patterned than females—perhaps
forests. They bury their eggs in heaps of compost chickens around the world are all descendants making it a closer ally of pheasants.
or in volcanic sand and rely on heat generated of the red jungle fowl of Southeast Asia.

22 in 28 in
55 cm 70 cm

31–36 in
78–92 cm

BRUSH TURKEY
Alectura lathami
f: Megapodiidae
Males of this E. Australian
megapode control the temperature black markings
of their incubation mounds by down the front
of the throat GREAT CURASSOW
adding or removing rotting Crax rubra
vegetation. More females hatch f: Cracidae
at high temperatures.
Ranging from Mexico to Ecuador,
MALEO this large curassow has a curly crest
Macrocephalon maleo like other curassows. Males are
f: Megapodiidae black, while females are brown.
This megapode (large-footed
bird) from Sulawesi island in
Indonesia lays its eggs in sand
and relies on heat from the
sun or underground volcanic
activity for incubation. 231/2 in 33 in
60 cm 84 cm

brown, white,
MALLEEFOWL and black BARE-FACED
Leipoa ocellata wings CURASSOW
f: Megapodiidae Crax fasciolata
Like many megapodes, this southern f: Cracidae
Australian species makes a compost mound This S. American curassow has a
to lay its eggs. It feeds primarily on various sparsely feathered face. Unlike other related
seeds, but is thought to be omnivorous. species, it has no fleshy wattles or bill knob.
 19–21 in
48–53 cm
gray-brown
upperparts
413

olive-brown 18 in 27 in 23–26 in
breast 46 cm 69 cm 59–65 cm

PLAIN GRAY-HEADED BLUE-THROATED

CHACHALACA CHACHALACA PIPING GUAN
Ortalis vetula Ortalis cinereiceps Pipile cumanensis HIGHLAND GUAN
f: Cracidae f: Cracidae f: Cracidae Penelopina nigra
Ranging from Texas Named after their calls, This S. American bird f: Cracidae
to Costa Rica, this chachalacas are brown- has glossy black plumage This C. American bird is
northernmost member colored relatives of typical of tree-dwelling more ground-dwelling than
of the American curassow curassows. This is a species piping guans. They are other guans. It is perhaps the
family is the only one of scrubby habitat from named for their shrill only member of its family
found in the US. Honduras to Colombia. breeding call. 22 in
to nest on the ground.
55 cm

white-tipped tail BEARDED GUAN

26–30 in Penelope barbata 26–30 in
66–76 cm f: Cracidae 67–75 cm
Named for its streaked
neck, the bearded guan,
like related species in
this genus, is a rainforest
SPIX’S GUAN bird. It is found in
Penelope jacquacu Ecuador and Peru.
f: Cracidae
This S. American species
belongs to a group of
brown guans that resemble
chachalacas, but have
shorter legs. It nests in trees
and has a loud mating call. red throat
30 in wattle
76 cm

CAUCA GUAN bare,

Penelope perspicax bluish head 27–30 in
f: Cracidae 68–75 cm
A bronze-sheened
relative of the spix’s guan,
this bird is found only in 24–28 in
the Cauca Valley region in 61–71 cm DUSKY-LEGGED CHESTNUT-BELLIED
the western and northern GUAN GUAN
parts of Colombia. Penelope obscura Penelope ochrogaster
f: Cracidae f: Cracidae
The only brown guan with The throat wattle that
dark rather than reddish legs, is typical of brown guans is
this species occurs in central especially well-developed
S. America, from Brazil to in this species from
northern Argentina. south-central Brazil.

10–12 in 91/2–101/2 in 10 in
26–31 cm 24–27 cm 25 cm

cape of
long feathers

VULTURINE
GUINEA FOWL
Acryllium vulturinum
f: Numididae MOUNTAIN QUAIL CALIFORNIAN QUAIL GAMBEL’S QUAIL
Guinea fowl are African game Oreortyx pictus Callipepla californica Callipepla gambelii
birds that live in groups, f: Odontophoridae f: Odontophoridae f: Odontophoridae
but are monogamous. They all This native of the Rocky Found from Oregon to California, This relative of the Californian quail
have a bare head. This is the Mountains, like other American this species is characterized by its has a longer crest. It occurs in the
largest species of guinea fowl quails, is a ground-nesting, forward-drooping crest, comprised desert in the south of California and
and is found in E. Africa. monogamous species. of a cluster of six feathers. the ranges of the two do not overlap.
 FOW L , GA M E BI R DS, 15 in
414 A N D R E L AT I V E S 38 cm
BIR DS • FOWL , GA ME BIRDS, AND REL ATIVES

121/2–14 in 81/2 in
32–35 cm 21 cm
CHESTNUT-BELLIED
HILL PARTRIDGE
ARABIAN PARTRIDGE Arborophila javanica
Alectoris melanocephala f: Phasianidae
f: Phasianidae Found in S.E. Asian rainforests,
CHUKAR PARTRIDGE This large, red-legged partridge hill partridges are small, cryptically
Alectoris chukar black stripes occurs in semidesert regions of the colored game birds with stumpy
f: Phasianidae on flanks Arabian Peninsula and Yemen. Its tails. Many, such as this Javan species,
With a range extending across central preferred habitat is juniper woodland. have distinctive head markings.
Eurasia, this is the most widespread of the
red-legged partridges—typically birds of
dry country, with striking black markings.

10–15 in 10 in
25–38 cm 26 cm

111/2–121/2 in 11–12 in
29–32 cm 28–30 cm

12 in
31 cm

RED-NECKED FRANCOLIN ROULROUL CHINESE BAMBOO

GRAY PARTRIDGE GRAY FRANCOLIN Francolinus afer Rollulus rouloul PARTRIDGE
Perdix perdix Francolinus pondicerianus f: Phasianidae f: Phasianidae Bambusicola thoracicus
f: Phasianidae f: Phasianidae A ground-nesting bird of forest Related to hill partridges, the f: Phasianidae
This is the most widespread of a As with related jungle fowl, and grassland, this species is S.E. Asian roulroul has a more There are two species of bamboo
small genus of dark-bellied Eurasian males of this S. Asian francolin widespread through Africa, colorful plumage. Males are dark blue partridge, birds of E. Asia that
game birds that are more related to have leg spurs that they from the Democratic Republic and have a reddish crest; females are are related to francolins and
pheasants than other partridges. use in fighting. of Congo to the Cape. green and lack the crest. jungle fowl. This bird is
native to China.

61/2–7 in 151/2–16 in 16–181/2 in

16–18 cm 39–40 cm 41–47 cm

black
throat

16–20 in
COMMON QUAIL SPRUCE GROUSE 40–50 cm
Coturnix coturnix Canachites canadensis
f: Phasianidae f: Phasianidae
Unusually for a game bird, Like other woodland SOOTY GROUSE SHARP-TAILED
this tiny species of W. Eurasian grouse, this N. American Dendragapus fuliginosus GROUSE
grasslands and semidesert is species can digest the needles f: Phasianidae Tympanuchus phasianellus
migratory in the northern of conifer and spruce trees, One of several f: Phasianidae
parts of its range. rejected by most animals. N. American grouse that This N. American grouse
have an inflatable neck sac, is a more widespread, more
this dark-colored species northerly cousin of prairie
inhabits the pine forests chickens. Males have purple
of the Pacific coast. sacs for display.
17 in 15–16 in
43 cm 38–41 cm

GREATER LESSER PRAIRIE

PRAIRIE CHICKEN CHICKEN
Tympanuchus cupido Tympanuchus pallidicinctus
f: Phasianidae f: Phasianidae
This species is from the C. US. Found in southern N. America,
As with other species of prairie this small species of prairie
chicken, males engage in group chicken has a colored neck sac and
courtship displays by inflating drumming call, typical
their colored neck sac. of Tympanuchus grouse.
 231/2–34 in 131/2–14 in RED GROUSE pale gray KALIJ PHEASANT
Lagopus lagopus scotica tail feathers Lophura leucomelanos
60–87 cm 34–36 cm
f: Phasianidae f: Phasianidae 415
Unlike most ptarmigan This pheasant occurs in forests
species, this British race from the Himalayas to Myanmar.
of the widespread There is also an introduced
circumpolar willow population in the
grouse does not turn Hawaiian Islands.
white in winter.
15–16 in
38–41 cm

WESTERN CAPERCAILLIE ROCK PTARMIGAN

Tetrao urogallus Lagopus muta
f: Phasianidae f: Phasianidae
Found in western Eurasian One of three related grouse 231/2–32 in
coniferous forests, this is the that turn white in winter, the 60–80 cm
largest member of the grouse rock ptarmigan is a tundra
family. Males display to females and mountain bird with a SIAMESE FIREBACK
with rattling, popping calls. circumpolar distribution. Lophura diardi
f: Phasianidae 22–30 in
Like many pheasants, this S.E. Asian 55–75 cm
21–35 in bird has bare, red facial skin. As with
SATYR TRAGOPAN 53–89 cm
other species of Lophura, this feature
Tragopan satyra is common to both sexes.
f: Phasianidae
This is a Himalayan species
of Tragopan, a genus of Asian 231/2–47 in
tree-nesting pheasants. Male 60–120 cm
tragopans have an expandable
fleshy lappet and horns that COMMON PHEASANT
are used for display. Phasianus colchicus
f: Phasianidae
Native to the woodlands
of C. and E. Eurasia, this
231/2–28 in species has been introduced
60–70 cm into western Europe, where it white-tipped tail
16–20 in is now common in farmland. fanned in display LADY AMHERST’S PHEASANT
40–50 cm Chrysolophus amherstiae
f: Phasianidae
Like many other pheasants, only males of
this species have spectacular plumage, and
only females care for the young. This
bird is from China and Myanmar.

PALAWAN
PEACOCK-PHEASANT
Polyplectron napoleonis bald head
f: Phasianidae
Male peacock-pheasants
lack the long plumes of their
peafowl relatives, but display
tail feathers marked with
iridescent eyespots.

21/2–71/4 ft
0.8–2.2 m

INDIAN PEAFOWL
Pavo cristatus
f: Phasianidae
This tropical Asian species occurs
in India and Sri Lanka. Males court
females by raising their long plumes,
which are just above the tail.

31/2–4 ft
1.1–1.2 m
16–31 in
41–78 cm
WILD TURKEY
RED JUNGLE FOWL Meleagris gallopavo
Gallus gallus f: Phasianidae
f: Phasianidae Turkeys are large,
Unlike related francolins, males of this wattled game birds native to
Asian ancestor of domestic chickens mate N. America. This species from
with many females. They have a comb the S. US is the ancestor
and wattles, which are absent in females. of domesticated turkeys.
416 WA T E R F O W L
Species in this group are web-footed Unlike geese and swans, male ducks are usually more PHYLUM CHORDATA
birds adapted for surface swimming. brightly plumaged than the females, especially when
BI R D S • WAT ER F OW L

CLASS AVES

The majority feed on plants, although breeding. Both sexes of many duck species have a ORDER ANSERIFORMES

some eat small aquatic animals. vividly colored wing-patch called a speculum. FAMILIES 3

The typical “duckbills” of waterfowl have SPECIES 177

Most waterfowl have short legs set well back on internal plates that have evolved to strain aquatic
the body. These birds, the Anseriformes, swim food drawn in by a muscular tongue. All species
by propelling themselves with webbed feet, and retain this feature, even though some have
all have plumage kept waterproofed with oil adopted different feeding methods. Geese graze on
taken from a gland near the tail. South American grasslands, but swans are more aquatic and use their
screamers and the magpie-goose from Australasia long neck to plunge their head deep in water. Many
have partially webbed feet, spending much of ducks dabble—upending to feed on or just below
their time on land or wading in marshes. These the water surface. Others, such as pochards, dive
birds belong to two ancient families. All other under water to gather food. The saw-billed ducks
waterfowl are classified in a single family, the have a narrow bill with serrated edges to catch
most primitive group being the whistling ducks. fishes. One group of ducks, including eiders, scoters,
and mergansers, includes accomplished sea divers.
s wa n s , g e e s e , a n d d u c k s
In swans and geese, the sexes have similar plumage n esting
patterns. These long-necked, long-winged birds Most waterfowl are monogamous, some pairing for
occur mostly outside the tropics. Many northern life. They nest largely on the ground, but a few nest
species breed close to the Arctic and migrate in trees. The marine ducks come inland to
southward to overwinter. Ducks are breed. All waterfowl produce downy
Flying in formation reduces air drag,
generally smaller and have a shorter neck chicks that can walk and swim soon enabling birds like these snow geese to
than others in the Anatidae family. after hatching. conserve energy on long migrations.

20–43 in 23–28 in 22–28 in

50–110 cm 58–71 cm 56–71 cm

21–22 in
53–56 cm

RED-BREASTED CANADA GOOSE BARNACLE GOOSE HAWAIIAN GOOSE

striking red, Branta canadensis Branta leucopsis Branta sandvicensis
GOOSE black, and
Branta ruficollis white plumage
f: Anatidae f: Anatidae f: Anatidae
f: Anatidae The largest of the Branta This goose breeds on Arctic Confined to the Hawaiian Islands,
This is the most brightly colored geese, this species is native tundra in Greenland and Russia, this goose has reduced foot-webbing
member of the Branta group of to N. America, but has been where it avoids predators by and strong claws, and is adapted to
dark geese. It breeds in introduced to northern Europe. nesting on cliff tops. climbing on to rocky lava flows.
northwestern Siberia, where it
nests close to raptors, possibly
for protection from foxes.
BAR-HEADED GOOSE
Anser indicus 28–30 in
f: Anatidae 71–76 cm
PINK-FOOTED GOOSE This bird is adapted to the thin air
Anser brachyrhynchus of montane C. Asia. Migrating high
f: Anatidae over the Himalayas, it overwinters
This small, gray goose pale gray body
in India and Myanmar. 30–35 in
breeds on rocky outcrops 76–89 cm
of tundra in Greenland and
Iceland, and overwinters in
western Europe. GRAYLAG
GOOSE
Anser anser
f: Anatidae
Widespread across the
grasslands and wetlands
in Eurasia, this typical
gray Anser goose is the
231/2–30 in wild ancestor of
60–75 cm domesticated geese.
 26–35 in 20–231/2 in 28–29 in
66–89 cm 50–60 cm 71–73 cm
417
finely barred, 151/2–171/2 in
gray body 39–44 cm

BI R D S • WAT ER F OW L
EMPEROR GOOSE ASHY-HEADED GOOSE EGYPTIAN GOOSE
Anser canagicus Chloephaga poliocephala Alopochen aegyptiaca PLUMED WHISTLING DUCK
f: Anatidae f: Anatidae f: Anatidae Dendrocygna eytoni
Found in northeastern Siberia S. American Chloephaga geese are perhaps Belonging to a group of southern f: Anatidae
and Alaska, this goose grazes on more related to ducks than to other geese. hemisphere geese that are related Distinct from geese and true ducks,
coastal grass and seaweed. It is less This species occurs in Chile and Argentina. to ducks, the Egyptian goose is whistling ducks are named for their
gregarious than most other geese. Like others in its genus, it may perch in trees. widespread through Africa. unique call. This is an Australian species.

BLUE-WINGED GOOSE COSCOROBA SWAN

28–35 in 30–39 in 231/2–30 in Cyanochen cyanoptera 3–4 ft Coscoroba coscoroba
70–90 cm 75–100 cm 60–75 cm f: Anatidae 0.9–1.2 m f: Anatidae
This goose has a thick plumage Resembling a goose, this is
as an adaptation to the cool the smallest of swans. It
highlands of its native Eritrea is confined to swamps in
and Ethiopia. the southern parts of
Chile and Argentina.

41/4–51/4 ft
1.3–1.6 m

MAGPIE-GOOSE CAPE BARREN GOOSE

Anseranas semipalmata Cereopsis novaehollandiae
f: Anseranatidae f: Anatidae red bill
From Australian This distinctive goose is
wetlands, this species has restricted to southern
long legs and partially Australia and offshore 31/2–41/2 ft
webbed feet. It is not islands, where it gathers 1.1–1.4 m
closely related to any in small flocks and
other waterfowl. grazes on grassland.
BLACK SWAN TRUMPETER SWAN MUTE SWAN
Cygnus atratus Cygnus buccinator Cygnus olor
blackish plumage f: Anatidae f: Anatidae f: Anatidae
This sooty-black swan with Related to other vocal This species breeds in
white wing tips may nest in swans, such as the Eurasian Europe and C. Asia. Like
huge colonies. Native to whooper swan (C. cygnus), other swans, it grazes on
Australia and Tasmania, it has this N. American species underwater vegetation
been introduced to New has a loud, honking call. by submerging its head.
Zealand, Europe, and
N. America.
straight neck 31/2–4 ft
1–1.2 m

all-white body

5–6 ft
1.5–1.8 m
BLACK-NECKED SWAN
Cygnus melancoryphus
f: Anatidae
Found in southern S. America, this species
spends more time on water than other swans
and nests on floating vegetation.

15–16 in 12–13 in 24–26 in 33–37 in

38–40 cm 30–33 cm 61–66 cm 83–95 cm

WHITE-BACKED DUCK AFRICAN PYGMY-GOOSE ORINOCO GOOSE SOUTHERN SCREAMER

Thalassornis leuconotos Nettapus auritus Neochen jubata Chauna torquata
f: Anatidae f: Anatidae f: Anatidae f: Anhimidae
This bird from Africa and Like other pygmy-geese, A S. American relative Screamers are large, bulky
Madagascar is related to this African species nests in of ducks, this goose birds that live in S. American
whistling ducks, but spends tree hollows. It is usually found occurs on tropical wet marshlands. Like other screamers,
more time on water, nesting in wetlands with water lilies, savanna and forest this species has bony wing spurs
on islands of vegetation. on which it feeds. edges along rivers. that may be used in fighting.
 WAT E R F OW L cream, black,
and green
151/2–17 in 17–22 in
39–43 cm 43–56 cm
418 head pattern
18–22 in
45–56 cm
BI R D S • WAT ER F OW L

NORTHERN SHOVELER
Spatula clypeata
AMERICAN WIGEON f: Anatidae
Mareca americana Both sexes of dabbling (surface-feeding)
f: Anatidae BAIKAL TEAL ducks typically have a colored wing-patch
This species feeds by dabbling on Sibirionetta formosa called a speculum. This widespread
the surface of shallow waters and f: Anatidae wetland species of the northern
occasionally upends. Huge flocks This distinctive duck hemisphere has a green speculum.
winter in the Caribbean after breeds in cold, open forests
breeding in N. America. in Siberia on the edge of
the tundra, and migrates
to E. Asia in the winter.

22–26 in
55–65 cm
20–26 in
50–65 cm
MALLARD
20–26 in 15–20 in Anas platyrhynchos
50–65 cm 38–51 cm f: Anatidae
Widespread in the northern hemisphere, the
surface-feeding mallard can interbreed with
related species, perhaps indicating that this
group has only recently evolved.

13–16 in
33–40 cm

orange cheek
plumes

INDIAN RUNNER DOMESTIC DUCK WHITE-CHEEKED PINTAIL BUFFLEHEAD

Anas platyrhynchos Anas platyrhynchos Anas bahamensis Bucephala albeola
f: Anatidae f: Anatidae f: Anatidae f: Anatidae
A domestic descendant Most domesticated ducks have This saltwater dabbling duck The smallest sea duck in N. America,
of the wild mallard, this descended from the mallard. is found in S. American estuaries the bufflehead nests in tree-holes
long-necked breed originated They are raised for their meat, and mangrove swamps. Unlike and sometimes uses those
in the Malay Peninsula and eggs, or down, or kept for temperate northern pintails, both vacated by woodpeckers.
India in the 19th century. ornamental purposes. sexes look alike.
17–20 in
43–51 cm
15–20 in
38–51 cm

MANDARIN DUCK
Aix galericulata
HARLEQUIN DUCK WOOD DUCK f: Anatidae
Histrionicus histrionicus Aix sponsa Wrongly thought to be 16–20 in
f: Anatidae f: Anatidae monogamous, this tree-nesting 41–51 cm
A highly buoyant sea duck, this species rides choppy The newly hatched chicks of this N. American duck is a symbol of love in its
waters and nests beside fast-flowing streams in tree-perching duck jump down from their native northeastern Asia. It has
eastern N. America, Iceland, nests in high tree-holes to reach been introduced in Europe
and western Russia. the water below. and California.

TORRENT DUCK
Merganetta armata
f: Anatidae
A powerful swimmer, this
high-altitude S. American duck
lives in fast-flowing rivers of the
Andes and nests beneath RINGED
riverside rocks. TEAL
Callonetta leucophrys
f: Anatidae
Like other tropical ducks, white flank
17–18 in this S. American teal is not patch
43–46 cm migratory and retains its
14–15 in plumage colors throughout
35–38 cm the year.
 18–22 in 16½–20 in 18–22 in
46–55 cm 42–50 cm 46–54 cm
419
white cheek
patch

BI R D S • WAT ER F OW L
BRONZE-WINGED
DUCK
SURF SCOTER Speculanas specularis
Melanitta perspicillata HOODED MERGANSER f: Anatidae
f: Anatidae Lophodytes cucullatus This duck is found along
The N. American surf scoter, like other f: Anatidae the rivers of S. America.
scoter species, breeds near fresh water Found in N. America, this bird has It is locally called “dog-duck”
and winters at sea. The male’s a saw-edged bill for catching fishes. It after the barking call of
body is entirely black. dives with powerful kicks of the feet. the female.
16–18½ in
40–47 cm
14–18 in 14–17 in 19–24 in
36–45 cm 35–43 cm 48–61 cm

TUFTED DUCK
RUDDY DUCK Aythya fuligula CANVASBACK
Oxyura jamaicensis f: Anatidae Aythya valisineria
f: Anatidae A Eurasian member of the pochard group, f: Anatidae
This N. American duck, now introduced the tufted duck feeds mostly, though not This N. American bird is the largest species
PINK-EARED DUCK in Europe, has a stiff tail which it uses as entirely, on invertebrates in contrast to of pochard, a group of ducks with
Malacorhynchus membranaceus a rudder when diving. its largely vegetarian relatives. typically stocky bodies and large heads.
f: Anatidae
The pink head spot of this widespread
Australian species is less distinctive than SMEW 24–25 in
its zebra pattern. It consumes plankton, 14–17½ in
Mergellus albellus 61–63 cm
strained from water by beak flaps. 35–44 cm
f: Anatidae
15–23 in A member of the
38–58 cm group of cavity-nesting
mergansers, this species
is the only small
white duck found in
northern Eurasia. COMMON
SHELDUCK
LONG-TAILED DUCK Tadorna tadorna
Clangula hyemalis f: Anatidae
f: Anatidae orange
This gooselike duck is largely a
Unlike most other Arctic sea ducks, the long-tailed head lobe
coastal resident in Europe, but in Asia it
duck breeds in saltwater as well as freshwater habitats. migrates southward from inland regions
Males have a distinctive long tail. during winter. It nests in burrows.

20½–23 in 17–25 in 22 in
52–58 cm 43–63 cm 55–56 cm

RED-BREASTED MERGANSER KING EIDER ROSYBILL

Mergus serrator Somateria spectabilis Netta peposaca
f: Anatidae f: Anatidae f: Anatidae
Widespread across the northern hemisphere, the This bird breeds along This S. American duck is related to the diving
red-breasted merganser breeds on coasts and Arctic tundra coastlines. pochards, but spends more time feeding at the
spends more time at sea than Its large body size may help water surface. Only the males have a red bill.
other mergansers. it to dive deeply for
invertebrate prey. 20–24 in 17–19 in
51–61 cm 43–48 cm
rose blush
on breast

CRESTED DUCK STELLER’S EIDER

Lophonetta specularioides Polysticta stelleri
f: Anatidae f: Anatidae
This Andean species Like other related sea ducks
is a possible relic of a of Arctic and subarctic
S. American lineage that regions, this eider
was ancestral to more overwinters farther south
widespread dabbling ducks, in huge flocks, sometimes
such as the mallard. numbering 20,000 birds.
420 PENGUINS
With their two-toned plumage, erect A penguin’s short, densely packed feathers have PHYLUM CHORDATA
posture, and waddling walk, penguins downy bases that trap warm air, and a layer of fat
BIRDS • PENGUINS

CLASS AVES

are instantly recognizable as the classic beneath the skin provides further insulation. The ORDER SPHENISCIFORMES

symbol of the southern oceans. feather tips are greasy and waterproof, lubricated FAMILIES 1

by oil produced from a large gland on the rump. SPECIES 18

Residents of the coastal regions of the southern A complex system of blood-flow through the legs
hemisphere, all penguins are adapted to life in cold and feet ensures that the penguin’s body is not D E B AT E
water. Most species live around islands encircling chilled by standing on snow or ice. The largest F LY I N G A N C E S T O R S ?
Antarctica, although some are found on southern penguin species incubate their eggs on their feet.
coastlines of South America, Africa, and Australasia. All penguins have counter-shaded plumage (dark In the early 20th century, flightless
These flightless birds of the order Sphenisciformes above and pale below), which in the sea provides birds were popularly believed to be
primitive. It was hoped that study of
probably descended from a common ancestor shared camouflage against ocean-going predators, such
their embryos would reveal evidence
with the albatrosses. They may also be distant as leopard seals. of a direct link to a dinosaurian past.
cousins of the divers from the northern hemisphere. Penguin eggs became sought after,
f o r ag i n g a n d n e s t i n g and members of Robert Falcon Scott’s
s p e c i a l a da p tat i o n s Penguins can dive for fishes, shrimp, and krill more last expedition (1910–13) made a
Penguins have legs set far back toward the tail, a than 200 times each day. When they have eggs to courageous journey to an emperor
feature that provides excellent propulsion in water incubate and young to tend, the parents take turns rookery in the bleak Antarctic winter.
and is seen also in birds such as divers and grebes. to forage. In the case of emperor penguins, one of By the time the eggs they collected
On land penguins walk upright, but with webbed the few species to breed on the Antarctic pack ice, received scientific attention, the
embryo theory had been disproved.
feet placed flat on the ground their gait is ungainly. the males incubate the eggs alone throughout the
Modern studies of anatomy, fossils,
In common with other flightless birds, penguins winter while the females remain feeding at sea. and DNA show that the ancestors
have reduced wings, but these are modified in such Most penguins are colonial breeders and often of penguins, and other
a way that they can be used as flippers. In effect, return to the same nest flightless birds, could fly.
penguins “fly” under water. site each season.

18–23 in
45–58 cm
KING PENGUIN EMPEROR
Aptenodytes patagonicus PENGUIN yellow
f: Spheniscidae Aptenodytes forsteri plume
Resembling the emperor f: Spheniscidae
penguin, this sub-Antarctic The largest penguin,
species has an orange-yellow this species breeds in
neck and breast markings, colonies on Antarctic
and incubates a single ice. Males incubate
egg on its feet. eggs during the
bitter polar winter.
3–3¼ ft
90–100 cm

white feathers ROCKHOPPER PENGUIN

contrast with Eudyptes chrysocome
black head f: Spheniscidae
and wings The smallest of the sub-Antarctic crested
penguins, this bird gets its name from its
habit of clambering over rocks and boulders.

3½–4 ft 28 in
1.1–1.2 m 70 cm

14–16 in
35–40 cm
22–23½ in
55–60 cm

LITTLE PENGUIN FIORDLAND PENGUIN MACARONI PENGUIN

Eudyptula minor Eudyptes pachyrhynchus Eudyptes chrysolophus
f: Spheniscidae f: Spheniscidae f: Spheniscidae
A burrow-nester, this Nesting in the cool coastal This bird inhabits islands in
bird is the smallest of all forests of S. New Zealand, the far southern Atlantic and
penguins. It occurs along this species has hairlike crest Indian oceans, but is the only
the coasts of S. Australia feathers and a red bill typical crested penguin to breed on
and New Zealand. of Eudyptes penguins. the Antarctic Peninsula.
 CHINSTRAP
PENGUIN
stubby bill 28–32 in
71–80 cm Pygoscelis antarcticus 421
f: Spheniscidae
The chinstrap penguin dives
white for krill and fishes. It breeds

BIRDS • PENGUINS
eye ring on Antarctic coasts and thin
islands of the South Atlantic. black streak
across face

18–30 in
46–75 cm GENTOO PENGUIN 26–28 in
Pygoscelis papua 67–72 cm
f: Spheniscidae
This penguin breeds on
the Antarctic Peninsula
and Southern Ocean
islands. Its nest is a
simple cluster of sticks,
stones, and feathers.

30 in blue-black
75 cm upperparts

ADELIE PENGUIN YELLOW-EYED

Pygoscelis adeliae PENGUIN
f: Spheniscidae Megadyptes antipodes
One of three “brush-tailed” f: Spheniscidae
Pygoscelis penguins from Antarctica A cousin of Eudyptes crested
and adjacent islands, the Adelie penguins, this rare New Zealand
penguin breeds in colonies of more bird nests in scrub, but not in
than 200,000 pairs. dense colonies like the other
Eudyptes penguins.

GALAPAGOS PENGUIN
Spheniscus mendiculus
f: Spheniscidae
This is the only penguin to black face
breed in tropical waters
cooled by the Humboldt
Current, which flows along
the western coast of
S. America. It nests
in rock crevices.

black
breast band

HUMBOLDT PENGUIN
19–20 in Spheniscus humboldti 27–28 in
48–51 cm f: Spheniscidae 68–70 cm
Found along the Pacific coast of
southern S. America, this species
belongs to a group of burrow-nesters
characterized by bold banded
patterns from flanks to thighs.

26–28 in
65–70 cm

MAGELLANIC JACKASS PENGUIN

PENGUIN Spheniscus demersus
Spheniscus magellanicus f: Spheniscidae
f: Spheniscidae 24–30 in Named for its
Closely related to the Humboldt 61–76 cm donkeylike braying call,
penguin, this banded penguin this bird is the only
lives in colonies around the penguin to breed in
southern tip of S. America Africa, in colonies on
and the Falkland Islands. southwestern coasts.
422 KING PENGUIN black upper
mandible
Apt enody t es pat agon icu s
The king penguin is the second-largest penguin species.
BIRDS • PENGUINS

Only its close cousin, the emperor penguin, is bigger.

Unlike the emperor, the king inhabits subantarctic
islands. It hunts fishes, ignoring the krill taken by
its rivals, and dives to extraordinary depths to get
them—sometimes to more than 660 ft (200 m). It
produces just one egg at a time and takes more
than a year to raise the single young. This means
adults cannot breed annually and the enormous
breeding colonies—containing juveniles of
penguins drink
different ages—maintain a permanent presence sea water and excrete
on favored islands of the southern oceans. excess salt as brine
through the nostrils

size 35–39 in (90–100 cm)

habitat Flat coastal plains and waters
around subantarctic islands
distribution Islands of S. Atlantic
and S. Indian Oceans
diet Mostly lanternfishes, occasionally squid

SPINY TONGUE
A penguin’s tongue is muscular and spiny:
projections on the tongue surface called
papillae have evolved into backward-facing
barbs that help grip fishes caught in dives.
KEEN EYE
Penguins hunt by sight and have good
underwater vision. Bioluminescent
lanternfishes, caught on night dives,
predominate in the diet of this species.

WING PROPULSION
Penguins are flightless birds, but
they are propelled by feet and wings
during dives. Their flipperlike wings
effectively make them “fly” under water.
DENSE FEATHERS
The feathers, arranged in an
outer oily waterproof layer
and inner downy insulating
layers, are adapted for
diving in cold water.
SCALY SKIN
Scales on the legs and feet are
reminiscent of the reptilian
roots of all birds; the dark
skin may help to spread heat
to eggs and young.

EGG PROTECTOR WEBBED FOOT FIRM TAIL

The single egg is incubated above The kicking action of webbed The short tail consists of
the feet and beneath a fold of warm feet helps propel the bird stiffened feathers, used
skin called a brood pouch. Once the forward under water—and as a rudder under water.
chick has hatched, it uses the brood on land, when “tobogganing” In smaller species, it is
pouch for shelter. over snow on its belly. used as a prop on land.
 COUNTERSHADED DIVER
Below its distinctive yellow markings, the 423
king penguin has the tuxedo-style plumage
pattern typical of penguins: white beneath
and dark above. When diving, this pattern
offers camouflage from aquatic predators.

BIRDS • PENGUINS
Seen from below, the pale belly is harder
to see against the sunlit surface, and when
seen from above, the bird’s dark back blends
in with the dark of the water below.

yellow markings are

due to pigments called
carotenoids, which are
absent from some
penguin species

yellow stripe
on lower
mandible

yellow
breast
424 LOONS
These web-footed, fish-eating birds Although their pointed wings are relatively small PHYLUM CHORDATA
of Arctic waters have rear-set legs for their body size, loons are fast fliers. To get
BIRDS • LOONS

CLASS AVES

that limit their movement on land but themselves airborne, the larger species have to ORDER GAVIIFORMES

provide excellent thrust for swimming. patter across open water; only the red-throated FAMILIES 1

loon is able to take off from land. All species fly SPECIES 5

Loons are the only family in the order Gaviiformes. southward to overwinter.
The alternative name for these birds—loon—may
originate from their haunting “lunatic” wailing cries s h a r e d pa r e n t i n g
uttered during the breeding season, or possibly Territorial male loons choose nest sites in
from their ungainly motion when out of water. vegetation on the shores of clear Arctic lakes,
A loon’s legs are set so far back on the body that where both sexes incubate and tend the young.
the bird can only manage an awkward shuffle on Young loon chicks ride on the backs of their
land. In water, loons swim and dive with ease. parents, but they can swim and even dive soon
Their streamlined body and spearlike bill closely after hatching. After breeding, loons lose their
resemble those of penguins, which have similar striking head and neck patterns, and their
Loons fly with their outstretched head
habits—it is possible, though not certain, that duller body plumage makes it more difficult held slightly lower than the body, which
these birds may share a common ancestor. to distinguish the different species. gives them a hunchbacked appearance.

COMMON LOON 27–36 in 30–36 in

Gavia immer 69–91 cm 76–91 cm
f: Gaviidae WHITE-BILLED LOON
One of the largest loons, this bird Gavia adamsii
breeds on lakes in subarctic regions striped f: Gaviidae
of N. America and Iceland. It neck patch black head A large species of Arctic
overwinters on coasts further south, and neck
waters, this white-billed
including those around Britain. bird can be distinguished
from other loons by its
yellowish white bill.

grayish head and neck

21–27 in 23–29 in 23–29 in

53–69 cm 58–74 cm 58–73 cm

BLACK-THROATED
LOON
Gavia arctica
f: Gaviidae
Largely confined to
Eurasia when breeding, this
species sometimes reaches
Alaska, but winters further
RED-THROATED LOON PACIFIC LOON south, including the Pacific
Gavia stellata Gavia pacifica coast of N. America.
f: Gaviidae f: Gaviidae
The smallest loon, this bird breeds on small tundra This species has similar striped plumage to that
pools in circumpolar regions and migrates south to of the black-throated loon. Both species have a
Europe, China, and the southeastern US to overwinter. white throat when not breeding.

white spots
in summer
A L B AT RO S S E S , P E T R E L S , 425
A N D S H E A RWA T E R S

BI R D S • A L BAT ROS SE S , PET R EL S , A N D SH E A RWAT ER S

Long-winged albatrosses and their birds deter predators by regurgitating a noxious PHYLUM CHORDATA
relatives spend much of their lives in oil that they produce in their stomach and CLASS AVES

the air, traveling great distances as they sometimes forcefully eject. This oil is also ORDER PROCELLARIIFORMES

scan the surface of the oceans for fishes. nutritious enough to feed their chicks. FAMILIES 4
SPECIES 147

These birds, the order Procellariiformes, are s l ow r e p ro d u c t i o n

master aeronauts, rarely returning to land except Tubenoses form pair-bonds, sometimes for life,
to breed. Albatrosses, petrels, and shearwaters— which in the larger species can be several decades.
called “tubenoses” because of the tubular nasal Many species breed in colonies on remote islands,
protuberances on their bill—are globally often returning to the same site year after year.
widespread, ocean-going birds, but show the Smaller species nest in cavities and burrows.
greatest diversity in the southern hemisphere. These birds have low breeding rates, but the
Unusually for birds, they locate their sparsely parents invest heavily in rearing their young.
scattered sea prey by smell. All but the smallest Typically, tubenoses make only one breeding
diving species in this group have elongated wings, attempt per season and produce a single egg.
Long-lived wandering albatrosses
and almost all have their webbed feet positioned Despite a lengthy incubation period, chicks are are monogamous.They reinforce their pair-
so far back that some cannot easily walk. These helpless upon hatching and mature very slowly. bonding with an elaborate courtship dance.

WANDERING ALBATROSS
Diomedea exulans
f: Diomedeidae
The largest of the giant Southern Ocean
Diomedea albatrosses, this bird forms
a life-long pair bond and produces a 3½–4½ ft
single young every two years. 1.1–1.4 m
LAYSAN
mostly white body ALBATROSS
Phoebastria immutabilis
dark wings f: Diomedeidae
turn white Albatrosses from the
with age
northern Pacific include those
that breed in the tropics, like
this small species that nests
on islands—including
Hawaii—when breeding.

pale pink bill

30–32 in
77–80 cm

27–29 in BLACK-FOOTED ALBATROSS

68–74 cm Phoebastria nigripes
f: Diomedeidae
Like other northern Pacific albatrosses, this
small, dark species frequently interrupts its
gliding flight with bursts of wing-flapping.
black “brow”

18–20 in
45–50 cm

NORTHERN FULMAR
Fulmarus glacialis
f: Procellariidae
This gull-like petrel is BLACK-BROWED ALBATROSS 32–37 in
common throughout the Thalassarche melanophrys 80–95 cm
northern hemisphere. It f: Diomedeidae
nests on cliffs and can eject This is one of several dark-backed albatrosses called
a foul-smelling stomach mollymawks, from the southern hemisphere. It nests
oil to repel predators. in dense colonies, producing one egg each year.
 A L BAT ROSSE S , PE T R E L S ,
426 A N D S H E A RWAT E R S
sooty
brown head
12 in
BI R D S • A L BAT ROS SE S , PET R EL S , A N D SH E A RWAT ER S

30 cm

AUDUBON’S PINK-FOOTED 19 in 18–18½ in

SHEARWATER SHEARWATER 48 cm 45–47 cm
Puffinus lherminieri Ardenna creatopus
f: Procellariidae f: Procellariidae
This small bird breeds on A variable species existing
tropical oceanic islands. in dark and light forms, this BULLER’S
Different populations are shearwater nests on islands off SHEARWATER
considered by some to be Chile. It migrates to the eastern Ardenna bulleri
separate species. Pacific during summer. f: Procellariidae
This shearwater nests
on islands off N. New 6½–8 in
Zealand but roams across 17–20 cm
the Pacific outside the
18–22 in breeding season.
45–56 cm

CORY’S SHEARWATER ANTARCTIC PRION

Calonectris diomedea Pachyptila desolata
f: Procellariidae f: Procellariidae
A large shearwater that glides with bowed wings, Prions are small, gray
this species breeds on islands in the eastern Southern Ocean petrels
Atlantic and disperses more widely in winter. that skim the sea to strain
plankton with their
12 in flattened bill.
31 cm

14–16 in
36–41 cm

17 in JOUANIN’S
43 cm 16 in
PETREL 41 cm
Bulweria fallax
f: Procellariidae
A tropical petrel of the
northwestern Indian Ocean,
this bird has a weaving flight. SNOW PETREL
It is closely related to the Pagodroma nivea
gliding shearwaters. f: Procellariidae
One of the few birds that
breed on the Antarctic, this
petrel breeds farther south than
ANTARCTIC PETREL any other bird, even wandering BLACK-CAPPED
Thalassoica antarctica to the South Pole. PETREL
f: Procellariidae Pterodroma hasitata
A bird of sub-Antarctic f: Procellariidae
waters, this large species Like many of the small,
dives for fishes and squid. fast species known as gadfly
It breeds on islands petrels, this one has a tropical
around the Antarctic. distribution. It breeds on
islands of the West Indies.
scattered dark feathers
on white body massive
yellowish bill

15½–16 in
39–40 cm

34–39 in 7½–8½ in
CAPE PETREL SOUTHERN GIANT 86–99 cm 19–21 cm
Daption capense PETREL
f: Procellariidae Macronectes giganteus BAND-RUMPED STORM PETREL
One of several members of f: Procellariidae Oceanodroma castro
this family, mostly circumpolar This carrion-eating bird f: Hydrobatidae
in the southern hemisphere, breeds in the South Atlantic. Typical of northern hemisphere storm
the cape petrel breeds on Unlike many other petrels, it petrels, this tiny bird has a white-banded
islands around Antarctica and has strong enough legs to rump and forked tail. It is seen in
winters farther north. walk well on land. both the Atlantic and Pacific.
GREBES 427
Birds of ponds and lakes, grebes the gland produces a secretion that is 50 percent PHYLUM CHORDATA
characteristically swim low in the paraffin. Grebes have small wings and many

BIRDS • GREBES
CLASS AVES

water and dive propelled by their feet. species are often reluctant to fly, although ORDER PODICIPEDIFORMES

They feed on small aquatic animals. northern species are migratory, traveling FAMILIES 1

from inland habitats to the coast to overwinter. SPECIES 23

Like many other diving birds, grebes have legs that Traditionally, grebes, the order Podicipediformes,
are set far back on the body, making them clumsy are classified as close relatives of divers, penguins,
on land but agile in water. The toes of their feet are and albatrosses. However, new research suggests
lobed, providing good power strokes during diving, that they may be related to flamingos.
but minimizing drag in the water between strokes.
The feet may be used for steering, too; in other r i t u a l a n d r e p ro d u c t i o n
diving birds, the tail is used for steering. In the breeding season, some grebes perform
A grebe’s tail is nothing more than a tuft of elaborate courtship rituals. They nest on floating
feathers. This serves more as a social signal than mats of vegetation in freshwater habitats. The
as a rudder, and is often cocked to expose white young are fully mobile upon hatching and can
The courtship ritual of the great crested
feathers beneath. The plumage is dense and well swim, but seek refuge on a parent’s back for grebe culminates with both birds rising up
waterproofed by an oil gland. Uniquely in birds, the first few weeks. from the water clutching a clump of weeds.

9–11½ in 9½–14 in WHITE-TUFTED GREBE 12–15 in

23–29 cm 24–36 cm Rollandia rolland 30–38 cm
f: Podicipedidae
A native of southern
S. America, this is a bird of
open lakes with abundant water
weeds. A related short-winged
Andean species is flightless.

LITTLE GREBE
Tachybaptus ruficollis PIED-BILLED GREBE
f: Podicipedidae Podilymbus podiceps
Found across the Old World, this bird is the most 16–20 in
40–50 cm
11–13½ in
28–34 cm
f: Podicipedidae
widespread of the small, dumpy grebes known as This American bird is more stocky
dabchicks. It has a reddish neck when breeding. and has a stubbier bill than other
grebes. Northern birds overwinter
in the Caribbean, but tropical
populations are sedentary.

black head

RED-NECKED GREBE BLACK-NECKED GREBE

gray flanks Podiceps grisegena Podiceps nigricollis
f: Podicipedidae f: Podicipedidae
This species breeds in Eurasia and Like other Podiceps grebes, this
N. America and overwinters farther sharp-billed diver has a colorful head
south in coastal waters. Like other plumage when breeding. It is found 22–30 in
grebes, it migrates at night. across the northern hemisphere. 55–75 cm

18–20 in
46–51 cm
10–11½ in SILVERY GREBE WESTERN GREBE
25–29 cm Podiceps occipitalis Aechmophorus occidentalis
f: Podicipedidae f: Podicipedidae
This S. American grebe is a colonial One of two similar western
breeder, congregating on alkaline N. American grebes, this
or salty lakes. It is found from the species has a range from
Andes to the Falkland Islands. Canada to Mexico; northern
GREAT CRESTED GREBE populations winter off the
Podiceps cristatus dark gray back Pacific coast.
f: Podicipedidae
This Old World grebe is known for
its courtship display. Like other
grebes, both sexes are colorful. white throat,
breast, and belly
428 FLA MINGOS
These remarkable birds live on salty territory is simply determined by how far a bird’s PHYLUM CHORDATA
lagoons and alkaline lakes. Once neck will stretch from the nest. A few days after
BIRDS • FLAMINGOS

CLASS AVES

classified with storks, flamingos are hatching, flamingo chicks gather together in large ORDER PHOENICOPTERIFORMES

now thought to be related to grebes. crèches. The parent birds feed their young with FAMILIES 1

a liquid food called crop milk. SPECIES 6

Extreme gregariousness defines flamingo life. These

birds of the order Phoenicopteriformes congregate f i lt e r f e e d e r s
in huge flocks, sometimes numbering hundreds of Flamingos have a unique style of filter-feeding using
thousands. These vast groups are so tightly packed a specially adapted bill. Holding its head upside-
that individual birds cannot easily take flight and down, a flamingo strains planktonic algae and
must initially walk or run if disturbed. But the open shrimps from the water through hairlike structures
habitat that flamingos favor, coupled with the lining the bill. Pigments absorbed from the
vigilance of many birds, ensures that predators food give flamingos their characteristic pink
are easily spotted. color. There are few competitors for
The large flocks are necessary to stimulate food, as the organisms are taken from
Some of the biggest flocks of flamingos
breeding, and flamingo courtship involves otherwise barren inland waters are seen in the African Rift Valley, where
group displays. Pairs build mud nests and that are either very salty or caustic. lesser flamingos gather to feed.

CHILEAN FLAMINGO 31/4–41/4 ft CARIBBEAN

Phoenicopterus chilensis 1–1.3 m FLAMINGO extremely
f: Phoenicopteridae black-tipped Phoenicopterus ruber long neck
The most widespread flamingo in bill f: Phoenicopteridae
S. America, the Chilean flamingo This Caribbean species
has a range from Peru to Tierra del differs from the greater
Fuego, and is distinguished by flamingo in being somewhat
smaller and having a pale
gray legs with pink “knees.” pink
pinker plumage. bill
pinkish
white plumage bright red
wing feathers
4–4½ ft 3½–5 ft
1.2–1.4 m 1.1–1.5 m

GREATER FLAMINGO
Phoenicopterus roseus
f: Phoenicopteridae
With a range spanning
Africa, S. Europe, and C. Asia,
this is the largest and most
widespread of the flamingos.

31/4–3½ ft
1–1.1 m

pink “knee”
is the ankle
32–39 in
80–100 cm
slender gray leg

ANDEAN FLAMINGO LESSER FLAMINGO

Phoenicoparrus andinus Phoeniconaias minor
f: Phoenicopteridae f: Phoenicopteridae
One of two flamingo species restricted The smallest flamingo, this species
to the higher altitudes of the Andes, this from Africa and southern Asia
distinctively yellow-legged flamingo may occurs in huge numbers on
wander nomadically between lakes for food. highly alkaline lakes.
STORKS 429
Most species in this group are wetland of dead mammals to scavenging at refuse tips. CLASS CHORDATA
birds with long legs for walking Other storks have slimmer bills, some more or

BIRDS • STORKS
AVES AVES

through marshy or grassy habitats. less tapered to an upswept or downcurved tip. ORDER CICONIIFORMES

They snatch prey with their long bill. All storks have broad wings with “fingered” FAMILIES 1

tips. They soar on rising thermals of warm air to SPECIES 19

Storks, order Ciconiformes, are tall, upright birds, gain height before gliding off over great distances
most often found in open spaces. While some are to a new feeding area or on long migratory
heronlike waterside species, others forage on dry journeys. Large flocks of white storks cross the
ground. They have a long-striding walk but can Mediterranean Sea at its narrower points, in
perch on open branches and many nest in trees. the same way as migratory birds of prey.
Their varied diet includes fishes, amphibians, small
reptiles and mammals, and large insects. In Africa, c ol on i a l n e s t i ng
storks feed on small animals fleeing grassland fires, Many birds of this group are gregarious when
and exploit swarms of locusts and grasshoppers. breeding, and their nesting colonies may include
Storks have a strong and more or less pointed several species. All species produce helpless
Storks typically nest in trees, but in
bill. The marabou uses its heavy, powerful bill for young that must be reared in the nest for western Europe white storks make good
everything from displacing vultures at carcasses some weeks. use of level platforms high on buildings.

4½–5 ft red and 30–36 in WOOLLY-NECKED STORK

1.4–1.5 m black bill 75–91 cm Ciconia episcopus
f: Ciconiidae
The most widespread tropical stork,
this bird occurs in both Africa and
Asia. It prefers wetland habitats
but may stray into pasture.
3–4 ft black and
0.9–1.2 m white plumage

3¼–4 ft black plumage

1–1.2 m

WOOD STORK EUROPEAN

Mycteria americana WHITE STORK
f: Ciconiidae Ciconia ciconia
This N. American bird belongs to f: Ciconiidae
a group of storks with ibislike bills. Three Ciconia species breed
It submerges its open bill in shallow outside the tropics. This
water, snapping it shut on moving prey. European bird uses
thermal updrafts on
migration routes over
land to winter in Africa.

SADDLE-BILL STORK
Ephippiorhynchus senegalensis gap in bill
f: Ciconiidae
Related to the jabiru, this African
stork has a slightly upturned bill
with a yellow “saddle.” It occurs in
long, solitary pairs, even when nesting. 32–37 in
heavy bill 81–94 cm

4–5 ft 4–4½ ft AFRICAN OPENBILL

1.2–1.5 m 1.2–1.4 m Anastomus lamelligerus
f: Ciconiidae
Small, tropical wetland storks,
openbills use their distinctive
bill to catch and manipulate
MARABOU JABIRU mollusks before eating them.
Leptoptilos crumeniferus Jabiru mycteria This species occurs in mainland
f: Ciconiidae f: Ciconiidae Africa and Madagascar.
Like other Leptoptilos A large American
storks, this African species stork, the jabiru is
is bareheaded, which enables the tallest flying bird in
it to scavenge in carrion S. America. It expands
without soiling its feathers. It its featherless neck sac
flies with its head retracted. when excited.
430 I B I S E S , B I T T E R N S , H E RO N S ,
AND PELICANS
BIRDS • IBISES, BITTERNS, HERONS, AND PELICANS

Most species in the order Pelicaniformes fast thrust to snatch, but not spear, prey. Ibises PHYLUM CHORDATA
are wetland birds with long legs for probe for prey with their thin, curved bill, CLASS AVES

walking through marshy or grassy whereas spoonbills sweep their partly opened ORDER PELICANIFORMES

habitats. Their long toes help them bill—with a flat, rounded tip—sideways through FAMILIES 5
SPECIES 118
to stretch forward when hunting or shallow water in search of food. The bill snaps
grasp perches in mangrove swamps shut when prey is detected by touch. Herons
and egrets retract their necks in flight, unlike
or wet thickets. These birds have a spoonbills and ibises.
long bill for snatching prey. Pelicans are massively built, but broad-winged
While most birds in this group eat amphibians and expert fliers. They have a long bill with a
and fishes, they sometimes eat small mammals and deep, flexible pouch. When feeding (mostly when
insects as well. Herons, egrets, and bitterns have swimming, but brown pelicans also plunge-dive)
This pelican chick reaches deep into
modified vertebrae, forming an S-shaped they scoop up a large volume of water, filtering its parent’s throat to feed on regurgitated,
neck that gives them lightning- out food as the pouch empties. partly digested fishes.

gray-green 16–22 in long,

back greenish- 40–55 cm narrow crest
black cap
35–39 in
90–98 cm
GREEN HERON
Butorides virescens
f: Ardeidae 32–39 in
80–100 cm
black tinge to
A small heron of N. American white foreneck
wetlands, the green heron may
sometimes bait fishes with food
at the water’s edge. GREAT EGRET
yellow legs GRAY HERON
and feet Ardea alba Ardea cinerea
f: Ardeidae f: Ardeidae
28–32 in Not a true egret but a large white Familiar across Eurasia
70–80 cm heron, this marsh bird is widespread and Africa, the gray
throughout much of the world. heron, like other larger
herons, breeds in
colonies. It builds
stick nests in trees.
32–39 in
80–100 cm
WHITE-NECKED HERON
LITTLE BITTERN Ardea pacifica
Ixobrychus minutus EURASIAN BITTERN f: Ardeidae
f: Ardeidae Botaurus stellaris This large heron of wet regions in
Among the smallest of the f: Ardeidae Australia and New Guinea hunts
heron-bittern family, this skulking Like its relatives, this for insects and small vertebrates
10½–15 in Old World species clambers bittern is more frequently in marshes and grassland.
27–38 cm
around reeds—sometimes freezing heard than seen. It has a
upright in typical bittern manner. loud, booming call.

LITTLE EGRET
Egretta garzetta
f: Ardeidae
This Old World species has
started to colonize America. It
has a distinct black bill and legs,
and one race has yellow feet.

22–26 in 23½–28 in 22–22½ in 23–25 in

55–65 cm 60–70 cm 55–57 cm 58–63 cm

WHITE-FACED HERON TRICOLORED HERON LITTLE BLUE HERON

Egretta novaehollandiae Egretta tricolor Egretta caerulea
f: Ardeidae f: Ardeidae f: Ardeidae
A member of the egret group from This is an inhabitant of American This American species belongs to
Indonesia, Australia, and New swamps, where, like related species, the egret group. The purplish
Zealand, this species has a varied it uses its daggerlike bill to head and neck turn gray-blue
diet that includes insects and frogs. snatch small animals. outside the breeding season.
 CATTLE EGRET BOAT-BILLED HERON
Bubulcus ibis Cochlearius cochlearius
f: Ardeidae 27–32 in
f: Ardeidae 431
Actually a small white heron, 68–82 cm This heron has a broad
this globally widespread species forages black-tipped bill bill, adapted for scooping
on grassland and often follows livestock animal prey. It inhabits

BIRDS • IBISES, BITTERNS, HERONS, AND PELICANS

to catch disturbed prey. 18–20 in C. and S. American
45–50 cm mangrove swamps.

19–21 in
48–53 cm

16½–18 in BLACK-CROWNED
42–45 cm NIGHT HERON
Nycticorax nycticorax
f: Ardeidae
INDIAN POND long neck plumes
HERON when breeding Night herons have good
Ardeola grayii night vision. Found in
f: Ardeidae 23–26 in
most warm regions except
Common in S. Asia, Australia, this is the most
58–65 cm
this heron stalks aquatic widespread species.
prey, but it may also flush
fishes by flying low
over water. bare head
and neck
26–30 in
black wing 65–75 cm
plumes
23–30 in
59–76 cm

gray body SACRED IBIS

Threskiornis aethiopicus
STRAW-NECKED IBIS f: Threskiornithidae
Threskiornis spinicollis This is a common bird of
f: Threskiornithidae the wetlands and grasslands
So called because of the strawlike of Africa and Madagascar.
feathers at the base of its neck, It has been introduced
this nomadic ibis occurs in New into America
Guinea and Australia. and Europe.

SCARLET IBIS
Eudocimus ruber
27–30 in 22–24 in f: Threskiornithidae
69–76 cm 56–61 cm The national bird of Trinidad,
the scarlet ibis occurs in
tropical America and acquires
its scarlet pigment from
eating crustaceans.
REDDISH EGRET
Egretta rufescens AUSTRALIAN IBIS
f: Ardeidae Threskiornis molucca
This American species has f: Threskiornithidae
white and reddish gray This is a common species throughout
color forms. When fishing, Australia. It often invades urban 30 in 22–26 in
it sometimes holds out its areas, where it is sometimes 75–77 cm 55–65 cm
wings to reduce sun glare. considered a pest.

30–35 in
76–89 cm

HADADA IBIS BLACK-FACED IBIS GLOSSY IBIS

Bostrychia hagedash Theristicus melanopis Plegadis falcinellus
f: Threskiornithidae f: Threskiornithidae f: Threskiornithidae
A common African species found in Found in S. America, this Found throughout warm parts of the
grasslands, forests, parks, and gardens, species occurs in temperate world, this is the most widespread
the hadada ibis is named after its grassland habitat from the ibis species. Its tree-nesting colonies
distinctive flight call. Andes to Patagonia. may be accompanied by herons.
 I BISE S , BI T T E RNS , H E RONS , A N D PE L ICA NS gray
432 plumage
AFRICAN SPOONBILL EURASIAN SPOONBILL
Platalea alba Platalea leucorodia
f: Threskiornithidae f: Threskiornithidae
BIRDS • IBISES, BITTERNS, HERONS, AND PELICANS

The only spoonbill confined This species breeds in Eurasia

to wetlands of Africa, this and overwinters in Africa.
species is characterized Mature birds have a
by its red face broad yellow tip to
and legs. the black bill.
large
bill

35–36 in 32–35 in 4–5 ft

90–92 cm 80–90 cm 1.2–1.5 m

pinkish red ROSEATE SPOONBILL

patch on wing Platalea ajaja
f: Threskiornithidae
pink breast Like other spoonbills, this
tuft when distinctive American species feeds
breeding on small aquatic animals by swinging
SHOEBILL
its bill from side to side in water. Balaeniceps rex
f: Balaenicipitidae
Restricted to swamps from Sudan
to Zambia, this wading bird uses
its huge bill to scoop vertebrate
prey from muddy waters.

22 in
56 cm
28–34 in
71–86 cm
bold head and
neck pattern

orange throat pouch

HAMMERKOP
Scopus umbretta
f: Scopidae
This African wetland bird
all-white plumage builds huge, heavy-walled 3¼–4½ ft
nests using sticks and mud to 1–1.4 m
protect its young, who are
often left for long periods.

4¼–5¼ ft
1.3–1.6 m

4¼–5 ft
1.3–1.5 m

AMERICAN
WHITE PELICAN
Pelecanus erythrorhynchos
f: Pelecanidae BROWN PELICAN
This pelican breeds on inland lakes SPOT-BILLED PELICAN Pelecanus occidentalis
of N. America and overwinters Pelecanus philippensis f: Pelecanidae
on the coasts. In the breeding season, f: Pelecanidae This gray and brown pelican from the
it develops a flat “horn” on its bill. A species from S. Asia, the spot-billed pelican, S. US to S. America is a coastal-
like most other pelicans, fishes by scooping breeding species and, unlike other
prey while swimming on the water surface. pelicans, plunge-dives for fishes.
COR MORANTS, GANNETS, 433

A N D R E L AT I V E S

BIRDS • TROPICBIRDS
The order Suliformes includes marine Frigatebirds cross vast areas of sea but rarely settle PHYLUM CHORDATA

species, though some cormorants and on water, taking flying fishes or stealing food from CLASS AVES
other seabirds in the air. Cormorants and anhingas ORDER SULIFORMES
anhingas also visit freshwater habitats. feed on a variety of aquatic species by diving from FAMILIES 4

The birds in this group have webbing the surface, while gannets and boobies plunge
SPECIES 61

between all four toes. from a height and are more exclusively fish-eaters.

32 in MASKED BOOBY yellow tinge to NORTHERN GANNET

81 cm Sula dactylatra back of head Morus bassanus
f: Sulidae f: Sulidae
This is the largest of the tropical white The three similar species of
BLUE-FOOTED boobies, with a black and white upperparts cold-water gannets breed in large
BOOBY plumage and long, yellow bill. It most colonies on rocky coasts. This one
Sula nebouxii closely resembles the related is a native of the North Atlantic.
f: Sulidae cold-water gannets.
Found on rocky coasts
from California to
Peru and the Galapagos 31–36 in
Islands, this booby flaunts 80–92 cm
its blue feet as part of its
courtship display.
3–3¼ ft
90–100 cm

snakelike
neck spearlike
bill

28 in 20–22 in LITTLE PIED 3¼–3½ ft

71 cm 50–55 cm CORMORANT 1–1.1 m
Microcarbo melanoleucos
f: Phalacrocoracidae
This is an Australasian MAGNIFICENT
30–37 in RED-FACED member of a group of FRIGATEBIRD
75–95 cm SHAG primitive, short-billed Fregata magnificens
Phalacrocorax urile “micro-cormorants” that f: Fregatidae
f: Phalacrocoracidae are largely associated Infrequent feeding
ANHINGA A deep-diving seabird found with freshwater or opportunities and long hours
Anhinga anhinga from Japan to the Bering Sea, estuarine habitats. in flight cause frigatebirds,
f: Anhingidae this species belongs to a group such as this American species,
This American species has a long, cormorantlike of North Pacific cormorants to have a low breeding rate
neck and a straight bill for spearing fishes. that are highly adapted to the and the longest period of
Anhingas are also called snake birds or darters. marine environment. parental care of any bird.

T RO P I C B I R D S black
PHYLUM
CLASS
ORDER
CHORDATA

PHAETHONTIFORMES
AVES

“mask”
Large, ternlike seabirds with a FAMILIES 1
SPECIES 3
long, spikelike tail projection,
tropicbirds nest on tropical
islands and forage widely at sea. WHITE-TAILED RED-BILLED
TROPICBIRD TROPICBIRD
The three species in this order, the Phaethon lepturus Phaethon aethereus mostly white with
f: Phaethontidae f: Phaethontidae black wingtips
Phaethontiformes, have all four toes joined Tropicbirds are streamer-tailed
30–32 in
76–80 cm Found from the E. Pacific
by webs, like gannets and boobies, but are seabirds with feet so feeble to the Atlantic, this typical,
that they use their bellies hole-nesting tropicbird rears a
unable to stand. They locate fishes by sight to shuffle on land. This single, slow-growing chick,
while hovering before plunging from the air. species frequents most an adaptation to scattered 3–3½ ft
0.9–1.1 m
tropical coastlines. ocean food resources.
At their breeding colonies, groups of males
display in swirling, circular flights, swaying long, whiplike
their tail spike to impress watching females. tail projection
434 BIRDS OF PREY
The largest and most important group and flesh but have relatively weak feet. Most have PHYLUM CHORDATA
of day-flying hunters, almost all the a bare head, which prevents an unhygienic, sticky
BIRDS • BIRDS OF PREY

CLASS AVES

birds in the order Accipitriformes mess when probing inside animal carcasses. New ORDER ACCIPITRIFORMES

are exclusively meat-eaters. In some World vultures may be more closely related to FAMILIES 4

storks than to the similar-looking Old World SPECIES 266

habitats they are the top predators. vultures. Eagles, hawks, buzzards, and kites have
Large birds of prey, such as the harpy eagle and stronger feet with long, curved, sharp claws,
Philippine eagle of tropical rainforests, are so used for catching and killing prey. The unique
powerful they can kill large monkeys and small secretary bird of African plains hunts on foot.
deer. The largest species, which include Old Some kites have a forked tail for greater
World vultures and American condors, feed control in flight. Bird-eating hawks have
on dead animals, and many, such as kites and rather short wings and a long tail, for extra
buzzards, eat carrion as well as catch live prey. maneuverability through woodland, while other
Exceptions include the largely vegetarian species have longer, broader wings that allow
palm-nut vulture of Africa. them to soar on rising air currents and glide long
The pale underwings of a turkey
Birds of prey typically have keen vision and distances with minimal effort. In this way, heavy vulture catch the sunlight as it tilts, twists,
hunt by sight. The turkey vulture detects prey vultures can move very long distances to find and turns, searching for food below.
by smell, and other species watch and follow it food each day, and some species perform very
down to hidden carcasses. Birds in this group long annual migrations. Their “fingered”
have a strong, hooked bill for dismembering prey. wingtips reduce turbulence at the wingtip,
Larger vultures are well-equipped to rip hide giving greater efficiency.

25–32 in TURKEY VULTURE

64–81 cm Cathartes aura
f: Cathartidae
Unusually for birds, this widespread
American vulture locates rotting carcasses
by smell. It often nests in dark recesses,
such as under large rocks or stumps.

22–26 in
26–32 in 56–66 cm
67–81 cm

contrasting black
and white wings

white streaks
on massive wings

ANDEAN CONDOR
Vultur gryphus
f: Cathartidae
S. America’s largest flying land bird,
this species soars on updrafts in the
Andes, locating carrion by sight or
by following other scavengers, such
as turkey vultures.
KING VULTURE BLACK VULTURE
Sarcoramphus papa Coragyps atratus
f: Cathartidae f: Cathartidae
This large bird soars high above forests of tropical More gregarious than the related turkey
America looking for carrion. It is distinguished vulture, the black vulture is an opportunistic
by its colorful head and fleshy bill wattle. scavenger found from the C. US to Chile.
 435
201/2–231/2 in 20–221/2 in 20–26 in 121/2–15 in
52–60 cm 51–57 cm 50–65 cm 32–38 cm

BIRDS • BIRDS OF PREY

dark brown to
white coloration

EUROPEAN
HONEY-BUZZARD EURASIAN BUZZARD LONG-LEGGED BUZZARD WHITE-TAILED KITE
Pernis apivorus Buteo buteo Buteo rufinus Elanus leucurus
f: Accipitridae f: Accipitridae f: Accipitridae f: Accipitridae
Belonging to a group of tropical A common raptor, this species has This buzzard of semideserts and Typical of its genus, this sharp-
raptors that eat bee and wasp larvae, light and dark color forms. Its mountains breeds in C. Europe and browed kite habitually hovers when
this species breeds in Eurasia and northern populations overwinter C. Asia, with some populations hunting. It occurs from the US to
overwinters in Africa. in tropical Africa and Asia. migrating to N. Africa in winter. S. America, outside the Amazon basin.

WHITE-EYED BUZZARD
Butastur teesa 15–17 in 20–25 in
f: Accipitridae 38–43 cm 50–64 cm
This small buzzard-hawk from
S. Asia is more terrestrial than related
species, hunting small animals and
insects on the ground.

SWALLOW-TAILED KITE
Elanoides forficatus
f: Accipitridae
This insect-eating bird of prey is
a graceful and agile flier. It breeds
in the S.E. US and C. America,
and overwinters in S. America.

BRAHMINY KITE
Haliastur indus
f: Accipitridae
With a range from India to
Australasia, this riverside and coastal
scavenger also hunts for live prey
such as fishes and small mammals.

white neck ruff sharply hooked bill

OSPREY
31/4–41/2 ft 201/2–26 in Pandion haliaetus head plumes
1–1.4 m 52–66 cm f: Accipitridae 17–20 in
Found almost worldwide, the red-brown 43–51 cm
fish-eating osprey plunges for tail
its prey and has a reversible
outer toe for a better grip
of its slippery catch.

41/4–5 ft
1.3–1.5 m

elongated central
tail feathers

PALM-NUT VULTURE SECRETARY BIRD long legs

Gypohierax angolensis Sagittarius serpentarius
f: Accipitridae f: Accipitridae
Unusually for a vulture, this One of the few raptors
African species has a largely 231/2 in to hunt on the ground, this
vegetarian diet, consisting of oil 60 cm long-legged bird of African
palm fruit. However, it also savanna chases small animals,
feeds on fishes and carrion. often stamping to disable them.
 BIRDS OF PREY WHITE-BELLIED SEA EAGLE
Haliaeetus leucogaster
436 f: Accipitridae
With a distribution from India
to Australasia along lakes and 28–34 in
28–38 in 72–85 cm
71–96 cm rivers, like other large eagles,
this fishing raptor builds
BALD EAGLE huge stick nests.
Haliaeetus leucocephalus
f: Accipitridae
This N. American sea eagle, the
US’s national emblem, captures
or scavenges fishes, sometimes
hunting cooperatively. It breeds
in woodland near water.

28–35 in
70–90 cm
22–28 in
55–72 cm

WHITE-HEADED VULTURE
Trigonoceps occipitalis
f: Accipitridae
Found in N., E., and S. Africa,
this vulture is often seen in pairs,
and is usually outnumbered by
BONELLI’S EAGLE other vulture species when
Aquila fasciatus feeding on carcasses.
f: Accipitridae
A woodland- and mountain-dwelling species,
this long-winged buzzardlike eagle has a range
that extends from S. Eurasia to N. Africa. EGYPTIAN VULTURE
Neophron percnopterus
f: Accipitridae
28–33 in A relative of the palm-nut vulture, this
70–83 cm
bird of S. Eurasia and Africa routinely
uses rocks to crack open ostrich eggs.

EASTERN IMPERIAL EAGLE pale head

Aquila heliaca
22–26 in f: Accipitridae
55–65 cm True eagles, including those of
the Aquila genus, such as this
Eurasian species, are described 23½–28 in
as “booted” because of their 60–70 cm
fully feathered legs.

AFRICAN HAWK-EAGLE
Aquila spilogaster
f: Accipitridae 24–30 in
A small raptor from 61–75 cm
sub-Saharan Africa,
this hawk eagle hunts in
wooded savanna and
hilly country.

23½–39 in dark brown

60–100 cm flight feathers

feathered legs

GOLDEN EAGLE CHANGEABLE HAWK-EAGLE

Aquila chrysaetos Spizaetus cirrhatus
f: Accipitridae f: Accipitridae
A graceful soaring bird, this large, Often crested, Asian hawk-eagles are
long-tailed eagle occurs in open country forest raptors. This variable species
across the northern hemisphere. It has dark and pale forms and is found
frequents forests in some regions. from the Himalayas to Indonesia.
 AFRICAN WHITE-BACKED VULTURE 3¼–4 ft
Gyps africanus 1–1.2 m
f: Accipitridae 437
One of the commonest vultures of the
sub-Saharan savanna, this species gathers
in large numbers near carcasses. These
LAPPET-FACED

BIRDS • BIRDS OF PREY

birds are seen in towns and villages.
VULTURE
Torgos tracheliotus
f: Accipitridae
Like the related Gyps
vultures, this carrion-eater
of arid Africa has a long
neck and a bare head to
prevent carcasses from
soiling its plumage.

34–38 in 3¼–4¼ ft
85–97 cm 1–1.3 m
35–39 in
90–98 cm

BEARDED VULTURE
Gypaetus barbatus
RÜPPELL’S VULTURE f: Accipitridae
EURASIAN GRIFFON Gyps rueppelli Native to the mountains
Gyps fulvus f: Accipitridae of Africa and Eurasia, this
f: Accipitridae An African relative of diamond-tailed, solitary
This vulture occurs in mountainous the Eurasian griffon, this vulture subsists largely
regions of S.W. Eurasia and darker species inhabits arid on marrow obtained by
N.E. Africa. It breeds areas. It has been recorded dropping bones on
and roosts among rocks as flying at higher altitudes rocks to split
and on ledges. than any other bird. them open.

neck ruff turns bulbous

white with age bill

18–20 in 18–22 in
46–51 cm 46–56 cm

BLACK-COLLARED
HAWK
Busarellus nigricollis WHITE HAWK
f: Accipitridae Pseudastur albicollis
A relative of the snail kite, f: Accipitridae
this species occurs in C. and This forest hawk of C. and S.
S. American wetlands, where America hunts for reptiles, especially
it fishes by dropping feet first snakes. It has been described as
into floating vegetation. lethargic and easily approached.

14–16 in 23½–26 in
36–40 cm 60–66 cm

DARK CHANTING
GOSHAWK
Melierax metabates
f: Accipitridae
An African hawk of dry,
open country, this bird
resembles a harrier
when in flight. It has a
musical piping call.

3–3½ ft 17–22 in
0.9–1.1 m 43–56 cm SNAIL KITE
Rostrhamus sociabilis AFRICAN HARRIER-HAWK
f: Accipitridae Polyboroides typus
This bird from Florida and f: Accipitridae
C. and S. American marshes has This raptor from sub-Saharan Africa eats oil-palm
a strongly curved bill, adapted fruit and hunts small vertebrates. Its flexible
for feeding on aquatic snails. “double-jointed” legs help it take prey from tree-holes.
438 RÜ P P E L L’ S
V U LT U R E
Gy ps r ueppelli
BIRDS • BIRDS OF PREY

One of the iconic scavengers of the African plains,

from Senegal east to Sudan and Tanzania, Rüppell’s
vulture flies so high in roaming for food that its blood
is specially adapted to capture oxygen in the thin air.
This vulture patrols dry mountainous terrain, leaving its
cliff-top roost sites in early morning to rise on updrafts
instead of thermals. It uses acute vision to find carcasses and
will wait patiently—days, if necessary—for predators to leave
a kill. Like most vultures, it eats soft, rotting flesh and offal. size 34–38 in (85–97 cm)
However, its longer neck means it can reach deeper into habitat Gorges in dry, open country
corpses than many of its competitors—gorging its fill distribution North and East Africa
diet Carrion
until, with some difficulty, it returns to the skies.

THIRD EYELID SCALLOPING nostril

A typical feature of birds, this This vulture’s dark wing
membrane cleans the surface feathers are broadly tipped
of the eye, and may help with lighter marking, giving
protect against flying debris the bird a scalloped
during frenzied feeding. appearance from
a distance.
WHITE COLLAR
White fluffy feathers
encircling the base of the
neck form a ruff. This may
be discolored with dust and
blood from carcasses.
PLUMAGE
Beneath the
patterned feathers
that mark the
contours of the body
are fluffy down
feathers that trap
body heat—vital
at high altitude.

hooked bill

rows of covert LEG A HEAD FOR GORE

feathers smooth Although the bird’s Although the sparse down of the
passage of longer, stiffer upper legs are vulture’s head and neck is frequently
air flow flight feathers feathered, its lower blood-stained, a fully feathered head
provide thrust legs are bare and so would get clogged with sticky debris
and lift for remain relatively as the bird reached deep inside the
flying clean when the bird is carcasses of large mammals to feed.
feeding on carcasses. The vulture’s hooked bill is used to
tear semi-rotten flesh, and is long
enough to probe carcasses.

FOOT
Because vultures use
their feet for walking
rather than killing, they
lack the big talons that
are typical of predatory
WING birds of prey.
Long, broad wings help the
vulture soar or glide, saving
energy. Takeoff can be a
struggle after a heavy meal.
pinkish gray skin
of head and neck
has a light coating 439
of down
 BIRDS OF PREY 19–22 in
48–56 cm
25–27 in
63–68 cm
440

17–18½ in
43–47 cm
17½–20½ in 12–14½ in
44–52 cm 30–37 cm

NORTHERN HARRIER BLACK-BREASTED

WESTERN MARSH HARRIER Circus hudsonius LIZARD BUZZARD SNAKE EAGLE
MONTAGU’S HARRIER Circus aeruginosus f: Accipitridae Kaupifalco monogrammicus Circaetus pectoralis
Circus pygargus f: Accipitridae Harriers are characterized by a f: Accipitridae f: Accipitridae
f: Accipitridae The males of this species are narrow tail; narrow, pointed A native of the African savanna, A bird of African grasslands,
Eurasian harriers, such as the Montagu’s brown, like the females, with gray wings; and long legs. This this bird mostly preys on large this eagle feeds on lizards
harrier, migrate to Africa and S. Asia. This on their wings and tail. Males of species is widespread insects, such as grasshoppers, and small mammals as
species inhabits grassland and reed beds. other harrier species are all gray. in N. America. but also eats small vertebrates. well as snakes.

red facial skin

SHIKRA
Accipiter badius
f: Accipitridae
19–24 in 11–16 in A typical Accipiter hawk
48–62 cm 28–40 cm with a long tail and short
wings, the Old World
shikra has a dashing flight
for catching small
animals, including birds.
NORTHERN GOSHAWK EURASIAN SPARROWHAWK
Accipiter gentilis Accipiter nisus
f: Accipitridae f: Accipitridae
This large hawk from N. One of nearly 50 species of Accipiter 10–14 in
America and Eurasia is capable of hawks, this hunter of small birds 25–35 cm
maneuvering through tall trees is found in woodland habitats
to catch squirrels and grouse. from Europe to Japan.

long, broad
23½–26 in 22–23½ in wings
60–66 cm 55–60 cm

RED KITE BLACK KITE

Milvus milvus Milvus migrans
f: Accipitridae f: Accipitridae
Like other Milvus kites, Found in open country across
this species from Europe Eurasia, Africa, and Australasia,
and the Middle East is the black kite has a varied diet
slightly weak-footed, but that includes live fish and small
accomplished at soaring. It mammals, carrion, and
frequently feeds on carrion. sometimes human refuse.
22–28 in
22–30 in 55–70 cm
55–75 cm

CRESTED SERPENT
EAGLE
Spilornis cheela BATELEUR
f: Accipitridae Terathopius ecaudatus red feet
Belonging to a group of Asian f: Accipitridae
snake eagles, this species This African savanna bird is the only snake
inhabits a range from India to eagle that regularly eats carrion. Bateleur
the Philippines, and is often means “acrobat” in French, a reference
found near fresh water. to this bird’s acrobatic flight.
FA L C O N S A N D C A R AC A R A S 441
Falcons are no longer considered to fliers. Some falcons hunt by catching large insects PHYLUM CHORDATA
be closely related to the main order and birds in the air, while others dive onto prey

BIRDS • FALCONS AND CARACARAS

CLASS AVES

of birds of prey, the Accipitriformes, on the ground. Some even take on larger birds ORDER FALCONIFORMES

but instead have their own order, in dramatic dives, stooping to strike at speed. FAMILIES 1

the Falconiformes. Several “hover” at a fixed point above the ground, SPECIES 67

to search for prey below, often detecting them

Falcons have a hooked bill, similar to that of the by the ultraviolet light emitted by their urine. 9½–13 in
Accipitriformes, but with an added “notch” used Caracaras also hunt and scavenge on the ground. 24–33 cm

to kill prey that is caught and held in the feet. Unlike most true birds of prey, falcons make no
They also have strong feet with sharp, curved nest but lay eggs on a ledge or in an old nest of
claws. Most have long wings and are marvelous some other species.

slate-gray
upperparts
dark
“moustache”
13½–23 in 10–12 in 8–12 in
34–58 cm 26–30 cm 20–31 cm

PEREGRINE
FALCON AMUR FALCON AMERICAN KESTREL
Falco peregrinus Falco amurensis Falco sparverius
f: Falconidae f: Falconidae f: Falconidae
As the fastest bird of prey, this Unusually for falcons, this raptor Kestrels are small falcons that
yellow falcon dives steeply on prey in habitually gathers in flocks; it breeds habitually hover when hunting.
feet midair. It is found throughout the in marshy woodland across Siberia This species is distributed
world in open country, including and China and migrates to southern throughout the Americas,
tundra and semidesert. Africa to overwinter. including the Caribbean islands.

COMMON KESTREL
Falco tinnunculus 12½–15½ in
f: Falconidae 32–39 cm
Like other kestrels, this species
BAT FALCON of open country across Eurasia
Falco rufigularis 9–12 in and Asia uses updrafts to MERLIN
f: Falconidae 23–30 cm
maintain its hovering, as it Falco columbarius
This swift-flying American scans the ground for prey. f: Falconidae
bird hunts birds, bats, and large A nimble predator with
insects at twilight. It is found a dashing flight, this
from Mexico to Argentina. falcon captures birds in
midair over hills and
moorland across the
northern hemisphere.
7–8½ in
18–21 cm yellowish-red
facial skin
black crown
CRESTED CARACARA and crest
Caracara cheriway
AFRICAN
16–18 in
40–46 cm
19–21 in
48–53 cm
f: Falconidae
PYGMY FALCON This common species of
Polihierax semitorquatus caracara is found in open
f: Falconidae country, from the S. US to
This African bird dives for northern S. America. It nests
insects and lizards on the in trees or on the ground.
ground. It breeds in
the nests of weaver birds
and may raise a brood
cooperatively with others.

STRIATED 21–24 in YELLOW-HEADED

CARACARA 53–62 cm CARACARA
Phalcoboenus australis Milvago chimachima MOUNTAIN CARACARA
f: Falconidae f: Falconidae Phalcoboenus megalopterus 19½–23 in
The tendency of this A hawklike scavenger f: Falconidae 49–58 cm
fearless caracara to that also eats oil palm Relatives of falcons, caracaras
attack newborn lambs fruit, this bird of are more sluggish and longer legged.
has led to its southern S. America Like other caracaras, this species of
persecution in its native frequents savanna and the high Andes is a scavenger, but
Falkland Islands. forest edges. also hunts small animals.
442 B U S TA R D S
Bustards are medium to very large in semideserts, open grasslands, and crops. PHYLUM CHORDATA
ground-living birds (including some In flight, they show large white areas on their
BIR DS • BUSTAR DS

CLASS AVES

of the heaviest flying birds) with long, long, broad wings. Larger species are powerful, ORDER OTIDIFORMES
FAMILIES 1
stout legs but a short bill. steady fliers, while smaller ones have rapid
SPECIES 26
wingbeats, looking more like small wildfowl
Birds in this order, the Otidiformes, have a steady, or gamebirds with square-tipped wings. Social
long-striding walk and pick insects, reptiles, small birds, they form flocks where numbers remain
mammals, and vegetable matter from the ground sufficiently high.

KORI BUSTARD AUSTRALIAN BUSTARD

Ardeotis kori Ardeotis australis 2½–5 ft 22–26 in
f: Otidae f: Otidae 0.8–1.5 m 55–65 cm
Among the heaviest flying birds, weighing This bustard inhabits grasslands and open
up to 42 lb (19 kg), this bustard occurs woodland of Australia and southern New
across eastern and southern Africa. It feeds Guinea. Males have a throat sac, which
on small vertebrates, carrion, and seeds. they inflate when displaying.

orange-brown
wings

28–43 in
3¼–4½ ft 70–110 cm
1–1.4 m
HOUBARA BUSTARD
Chlamydotis undulata
f: Otidae
A native of arid country, this bird
is found in open plains and barren
rufous desert areas of N. Africa and the
breast band 16–18 in Canary Islands.
40–45 cm
GREAT BUSTARD
Otis tarda
f: Otidae
Male bustards are bigger than females, LITTLE BUSTARD
especially so in this species of the Tetrax tetrax
Eurasian steppes. Adult birds take six f: Otidae
years to mature and develop plumes.
This diminutive bustard
breeds in open country across
Eurasia and migrates
southward in winter. In flight,
it resembles a shelduck.

21 in
53 cm

RED-CRESTED BUSTARD
Lophotis ruficrista
f: Otidae
Like other bustards, this
southern African species has a
spectacular courtship display.
Males perform tumbling aerial
flights, and pairs call in duet.
C R A N E S , R A I L S , A N D R E L AT I V E S 443
From graceful dancing cranes to small and have short toes with the hind toe reduced PHYLUM CHORDATA
skulking rails, this order contains a or absent. The finfoots and coots have lobed,

BIR DS • CR ANES, R AILS, AND REL ATIVES

CLASS AVES

wide variety of ground-dwelling birds not webbed, feet. Rails and crakes have longer ORDER GRUIFORMES

of both dry and wetland habitats. toes and a body that is deep but very slim, FAMILIES 6
SPECIES 188
helping movement through dense vegetation,
Mostly long-legged and long-billed, birds in the including reedbeds, in marshes, or
order Gruiformes are behaviorally very diverse. drier areas. Some on remote
Cranes walk on the ground, do not perch in trees, islands are flightless.

26–28 in
65–70 cm
YELLOW-LEGGED
BUTTONQUAIL
olive-brown Turnix tanki
wings and body f: Turnicidae
Like other buttonquails, this east
Asian species inhabits tropical
grasslands. The females are more
colorful, and compete for the drab
males, who take care of the young.
LIMPKIN
Aramus guarauna
f: Aramidae
Mostly nocturnal, this small cousin of cranes
inhabits tropical American wetlands and uses its 6 in
tweezerlike bill to extract snails from shells. 15 cm
9–12 in
22–30 cm
7½–9 in
19–23 cm

CORNCRAKE
Crex crex
f: Rallidae
This shy grassland crake is best
identified by its rasping “crex
crex” call. It breeds in Eurasia
BLACK CRAKE and overwinters in Africa.
Amaurornis flavirostra
f: Rallidae
This African crake is widespread
south of the Sahara. Unlike
many of its secretive relatives, it
long, gray legs is often seen out in the open.

BUFF-BANDED RAIL
Gallirallus philippensis
f: Rallidae
Unlike some of the other Indo-Pacific
Gallirallus species that are flightless or
nearly so, this rail has managed to
disperse to many oceanic islands,
from the Philippines to New Zealand.
11–13 in 15–19 in
28–33 cm 38–48 cm

5–7 in
13–18 cm

YELLOW RAIL KING RAIL

Coturnicops noveboracensis Rallus elegans
f: Rallidae f: Rallidae
One of several barred-backed rails, This species is found in eastern
this tiny, secretive N. American N. America, Mexico, and Cuba. It
species is most easily detected by forages in concealed locations for
its nocturnal clicking call. insects, spiders, shrimp, and snails.
 C R A N E S , R A I L S , A N D R E L AT I V E S grayish-brown
upperparts
444
9–11 in 8–10½ in
23–28 cm 20–27 cm

12½–16 in
32–41 cm

WATER RAIL VIRGINIA RAIL

Rallus aquaticus Rallus limicola
f: Rallidae f: Rallidae CLAPPER RAIL
Rallus rails are long-billed This long-distance Rallus longirostris
marsh birds with a narrow body migrant has a range dull flank f: Rallidae
for passing through reedbeds. Like from N. America to stripes Unlike many other rails,
other members of the genus, this northern S. America. It is this tropical American species
Eurasian bird is rarely seen away a secretive bird that is prefers salt marshes and mangrove
from dense cover. difficult to spot. swamps. It has a distinctive
clacking call.

10–13 in
26–33 cm
red bill with
yellow tip

SUNGREBE
Heliornis fulica 8½–10½ in
15½–16 in
39–40 cm
f: Heliornithidae 21–27 cm
This species is a tropical American finfoot. Like
all finfoots, it is a secretive bird of slow-moving
water, where it feeds on small animals.

RUDDY-BREASTED CRAKE
Porzana fusca
AMERICAN COOT 12–14 in f: Rallidae
Fulica americana 30–36 cm This crake occurs in wetlands of eastern Asia, but
f: Rallidae is also seen in mangroves or drier habitats. It has
Found from N. America to distinctive chestnut-colored underparts.
northern S. America, this bird,
unlike true rails, lives on water.
It forages in shallow water and
feeds on land.

8–10 in
20–25 cm

purplish-blue
underparts SORA RAIL
Porzana carolina
f: Rallidae
This crake is the most common N. American
green wings member of the rail family. It breeds in shallow
wetland and overwinters in the Caribbean.

7–9 in
WHITE-BROWED CRAKE 18–22 cm
Porzana cinerea
f: Rallidae
Shorter bills and distinctive calls distinguish
crakes from rails. This gray-fronted bird is a
typical Porzana crake. Its range extends
from the Malay Peninsula to Polynesia.

yellow legs

PURPLE
GALLINULE COMMON MOORHEN
Porphyrio martinica Gallinula chloropus
f: Rallidae f: Rallidae
Occurring in tropical American Moorhens are noisy birds with
marshes, this species belongs to a dark plumage and jerky 12½–14 in
group of rails called swamphens. movements. One of several 32–35 cm
It is characterized by blue-purple Gallinula species, this one has
plumage and a blue forehead shield. an almost global distribution.
 gray-white plumage bare red head
445

19–22 in 3½–4 ft 3 1/4–3½ ft

48–56 cm 1–1.2 m 1–1.1 m

GRAY-WINGED TRUMPETER BROLGA

Psophia crepitans Grus rubicunda
f: Psophiidae f: Gruidae BLUE CRANE
Named for their loud calls, This Australian crane Anthropoides paradiseus
trumpeters are gregarious Amazonian has a bare red head and f: Gruidae
ground birds that are weak fliers. black dewlap (flap under The long wing plumes of this African
This is a black-bodied species, the chin). It performs a crane resemble a tail. When not
which—like other trumpeters— spectacular prancing breeding, it is nomadic, frequenting
has a “hunchback” appearance. courtship display. lakeside, grassland, and farmland.

slate-gray
plumage

GRAY CROWNED
4½–5 ft
CRANE
1.4–1.5 m Balearica regulorum
f: Gruidae
African crowned cranes are red
the only cranes that can grip wattle
branches, enabling them to roost
in trees. This is the most
southerly species.
3½–4 ft
1.1–1.2 m
3½–4 ft
RED-CROWNED 1.1–1.2 m
CRANE
Grus japonensis 3½ ft COMMON CRANE
f: Gruidae 1.1 m Grus grus
Declining in numbers, this SANDHILL CRANE f: Gruidae
bird breeds in Siberia and Grus canadensis Frequenting marsh,
overwinters in Korea and f: Gruidae heathland, and tundra,
China. The heaviest of all A N. American crane, this this bird breeds in Eurasia
cranes, like related species, species also reaches westward and migrates to N. Africa
it has a bare, red into Siberia. It migrates south in and S. Asia, often in a
crown patch. family groups, as far as Mexico. V-formation.

K AG U A N D S U N BI T T E R N long crest

Recently separated from the cranes PHYLUM CHORDATA long, slender bill

and rails, the tropical kagu and CLASS

ORDER EURYPYGIFORMES
AVES

sunbittern are unique birds of damp FAMILIES 2

forest, limited to very restricted SPECIES 2

geographical ranges.
closely barred
upperparts
A handful of species worldwide are difficult to
off-white
place in a relationship with others: these two plumage
appear to be closely related, but look very 22 in
different. Their position in the evolutionary 55 cm

tree is still controversial. The kagu reveals

broad gray and white bands across its wings, KAGU
when they are spread in courtship displays. Rhynochetos jubatus
SUNBITTERN f: Rhynochetidae
The sunbittern likewise shows unexpected patches Eurypyga helias 17–19 in Confined to the forests of New
f: Eurypygidae 43–48 cm Caledonia in the southwestern
of color on its upper wings and tail, which are This heronlike predator inhabits the Pacific, the kagu lacks strong
fanned horizontally to create a single, broad humid forests of C. and S. America.
It flashes its colored wing spots for
muscles necessary for flight.
Instead, it uses its wings to
pattern when it is displaying. display or to startle intruders. glide and for display.
 juveniles lose their buff
face feathers as they
mature, leaving a patch
of white cheek skin

red throat sac

CROWN AND COLOR

This species sports a bristly golden
crown, a neat black forehead, and bare
white cheeks, edged with a patch of red,
neck feathers lack
which is broader in East African birds. Both
interlocking barbs, males and females have a red throat sac
giving a loose, that can be inflated with air, then rapidly
“hairy” appearance deflated to give a booming call.
 G R AY C ROW N E D 447
CRANE
B alear ica regu lor um
Gray crowned cranes belong to the Gruidae family of birds,
famous for their choreography; for these birds, dance is an
important part of life. On the open savanna they display by
jumping, flapping their wings, and bowing—sometimes, it
seems, just to relieve aggression or reinforce pair-bonding—
but mainly to court their mate by flaunting their elaborate
head ornamentation. They lack the long, coiled windpipe of
the longer-billed crane species, so instead of calling like a
bugle they honk like a goose. They also boom during courtship
by puffing air from an inflated red throat sac. During the
breeding season pairs retire to wetter habitats, where thicker
vegetation conceals their nest—a circular platform made size 31/2 ft (1.1 m)
habitat Open country
from grasses and sedges. Here, the chicks are hidden from distribution E. and S. Africa
predators, allowing the parents to roost, uniquely for diet Grasses, seeds, invertebrates,
cranes, perched high in trees. small vertebrates

RUFFLED NECK
black head feathers Long, tapered contour feathers
give the appearance give the crowned crane a shaggy
of a bulging
forehead appearance around the upper
THIRD EYELID part of the body and the lower
Also known as a part of the neck. Most of
nictitating membrane, the bird’s plumage is gray.
from the Latin nictare,
meaning “to blink,” this
nostril translucent eyelid—the
same as in other birds—
moves across the eye to
clean its surface.

GOLDEN PLUMES
When the wings are folded,
long, golden feathers on the
upper wings—just above
the flight feathers—hang
down over the side
of the body.

white wing
covert feathers
bill is shorter and
stouter than in other
crane species

black primary
FEET AND CLAWS flight feathers brown secondary
Crowned cranes have long flight feathers
hind toes, setting them apart LONG LEGS WING
from other crane species. This Although long legs help in dancing When crowned cranes fly overhead, their white
enables them to perch in displays, and are good for wading, underwing patches are clearly visible. Although
trees—perhaps a vestige of crowned cranes have shorter legs strong-winged, these tropical birds—unlike other
tree-dwelling ancestors. than those of other crane species. cranes—do not sustain long-distance migrations.
448 WA D E R S , G U L L S , A N D A U K S
Largely birds of coastal areas, waders of sandpipers, snipe, and allies, the birds use their PHYLUM CHORDATA
are diverse in form and habit. Many are long bill to probe deep mud. Some of their relatives,
BI R D S • WA DER S , GU L L S , A N D AU K S

CLASS AVES

adapted for feeding in mud and water, such as shanks and curlews, have long legs, too, and ORDER CHARADRIIFORMES

with long legs and a probing bill. so can wade into deeper water to feed. FAMILIES 19
SPECIES 383

The birds collectively known as waders, or ocean ic bi r ds

Charadriiformes, fall into three main groups. Of The final group in this order is made up of gulls, D E B AT E
these, two groups are made up of shorebirds. terns, jaegers, and auks—web-footed birds that ORIGIN OF A GULL
Plovers and allied species are mostly short-legged, are the most ocean-going of the order. They
short-billed birds that feed on small invertebrates may spend much of their lives at sea and some New species appear when populations
found close to the surface of the ground. Some travel immense distances on migration. Gulls diverge too much to interbreed with
are opportunistic predators that are also seen others. In this way it was thought that
birds in this group, such as lapwings, favor drier
European herring gulls diverged from
habitats inland. Other species in the plover group foraging far inland. Auks, distributed around Asian ancestors that spread eastward
are more adapted to wetland. Stilts and avocets the Arctic, are specialized for diving for ocean- to encircle the Arctic. Recent evidence
sweep shallow water with their needlelike bill, swimming prey. Their black-and-white coloration suggests herring gulls descended from
while oystercatchers use their long, stout bill for gives them a superficial resemblance to penguins, gulls isolated in the North Atlantic.
opening mollusk shells. In the group consisting but they are not close relatives.

SNOWY SHEATHBILL 8–9 in

Chionis albus 20–22 cm
f: Chionidae
This is one of two white sheathbills,
Antarctic cousins of plovers, which
scavenge and consume carrion
and the food and young
of other birds.

16–17½ in 19½–22 in
40–44 cm 49–55 cm
13½–16 in
34–41 cm

EURASIAN STONE CURLEW GREAT STONE CURLEW MAGELLANIC PLOVER

Burhinus oedicnemus Esacus recurvirostris Pluvianellus socialis
f: Burhinidae f: Burhinidae f: Chionidae
Named after their shrill, curlewlike Large Esacus stone curlews This unusual S. American
calls, the mostly nocturnal stone have a chisellike bill for hunting wader—the only one to
15–16 in regurgitate food for chicks—
curlews are related to plovers. This prey, such as crabs, near water. 38–41 cm
widespread Eurasian species This is a S. Asian species. is more closely related to
occurs in dry mudflats inland. sheathbills than other plovers.
IBISBILL
Ibidorhyncha struthersii
f: Ibidorhynchidae
The only plover relative with
a long, down-curved bill, this
dark brown to bird plucks invertebrates
black body from stony riverbeds in
mountains of C. Asia.
long, red bill

16½–18½ in 16–18 in 13–16 in

42–47 cm 40–45 cm 33–40 cm

AMERICAN BLACK pink legs

OYSTERCATCHER
Haematopus bachmani EURASIAN OYSTERCATCHER CRAB PLOVER
f: Haematopodidae Haematopus ostralegus Dromas ardeola
Black oystercatchers generally have f: Haematopodidae f: Dromadidae
more restricted ranges than pied The most widespread oystercatcher, The crab plover may be more closely
(having two or more colors) this pied species breeds in N. Eurasia. related to gulls than to true plovers.
species. This species is confined Like others, it uses its long bill Found around Indian Ocean coastlines,
to the west coast of N. America. to open bivalve mollusks. it uses its stout bill for eating crabs.
 RED-NECKED PIED AVOCET
AVOCET Recurvirostra avosetta
Recurvirostra f: Recurvirostridae 449
novaehollandiae This Eurasian wader is
f: Recurvirostridae typical of avocets: relatives
needlelike bill A nomadic bird of of stilts that take small
Australian wetlands, this aquatic animals by sweeping
distinctively marked their upturned bill
BANDED STILT 16–18 in wader is a sociable species
Cladorhynchus leucocephalus through water.
40–46 cm that feeds in large flocks.
f: Recurvirostridae
14–16 in Stilts hunt small swimming 16½–18 in
36–45 cm 42–45 cm
invertebrates. This Australian species
gathers in large flocks on salt lakes
to feed on brine shrimp.

black and upturned bill

white body

8 in
BLACK-WINGED
WRYBILL STILT
20 cm Anarhynchus frontalis Himantopus himantopus
f: Charadriidae f: Recurvirostridae
Related to lapwings, this wader This highly variable stilt is
from New Zealand is the only bird nearly global in distribution. 13–14 in
with a bill bent sideways, which it uses There are white-necked and 33–36 cm
to reach invertebrates under stones. black-necked forms that
may be separate species.

INLAND SPUR-WINGED PLOVER

DOTTEREL Vanellus spinosus
Peltohyas australis f: Charadriidae
f: Charadriidae Occurring in wetlands across Africa
This sand-colored relative and the Middle East, this bird is one of
of lapwings is found in several lapwings, such as masked and
arid parts of Australia, southern, with a spur on each wing.
white often far from water.
crown 7½–9 in 10–10½ in
19–23 cm 25–27 cm

10–11½ in
26–29 cm

10–12 in
25–30 cm
14–15 in
35–38 cm

11–12 in
black underparts 28–31 cm
when breeding

BLACK-BELLIED EURASIAN GOLDEN PLOVER MASKED LAPWING NORTHERN LAPWING

PLOVER Pluvialis apricaria Vanellus miles Vanellus vanellus
Pluvialis squatarola f: Charadriidae f: Charadriidae f: Charadriidae
f: Charadriidae Golden plovers are more closely related Many lapwings have yellow, fleshy Lapwings are named for the lapping
The only member of the golden to stilts and oystercatchers than they are wattles. These facial adornments sound of their flight. This Eurasian
plover group with gray, not to other plovers. This species develops blue-gray are especially prominent on species has a distinctive, pointed crest. It
yellow, speckled upperparts, a black belly when breeding. feet and legs this bird from Australasia. is also called a peewit for its unique call.
this species breeds around
Arctic coastal tundra.
7–8 in 9–10½ in 8–9 in
18–20 cm 23–27 cm 20–22 cm

EURASIAN
DOTTEREL
COMMON RINGED PLOVER KILLDEER Charadrius morinellus
Charadrius hiaticula Charadrius vociferus f: Charadriidae
f: Charadriidae f: Charadriidae Females of this tundra-
One of the most widespread This long-tailed, grassland breeding bird are more
ring-necked plovers, this species breeds plover migrates between N. and brightly colored than males,
around the Arctic and overwinters S. America, but there are resident but like many other plovers
in Africa and S.W. Asia. populations in Peru and Chile. both become duller in winter.
 WA D E R S , G U L L S , long tail golden
450 A N D AU K S when breeding neck patch

11–12 in
BI R D S • WA DER S , GU L L S , A N D AU K S

28–31 cm AFRICAN
JACANA
Actophilornis africanus
f: Jacanidae
Typical of the jacana
family, this African
9–12 in wetland bird has enlarged 8–10½ in
23–31 cm feet for walking on 20–27 cm
floating vegetation.
12–23 in 6½–9 in
31–58 cm 17–23 cm

BRONZE-WINGED JACANA COMB-CRESTED JACANA

Metopidius indicus Irediparra gallinacea
f: Jacanidae f: Jacanidae
This species is widespread across Found from Asia to Australia, this species
India and S.E. Asia. Males have a is named for its fleshy head wattle. As
flattened “forearm” bone that helps with other jacanas, males incubate the
them lift chicks with their wings. eggs and care for the young. PHEASANT-TAILED JACANA WATTLED JACANA
Hydrophasianus chirurgus Jacana jacana
f: Jacanidae f: Jacanidae
This southern Asian spur-winged The dominant female of this
6–7½ in 9–10 in jacana is the only species S. American species breeds
15–19 cm 23–25 cm that changes plumage—losing with many males, perhaps to
its long tail—outside the compensate for crocodile
breeding season. predation of eggs.

PLAINS WANDERER
9–10 in Pedionomus torquatus
23–26 cm
f: Pedionomidae
This quaillike bird of
Australian grassland—
the only member of its 6½–8 in
GREATER PAINTED-SNIPE 9–10 in 16–20 cm
Rostratula benghalensis family—is actually related 23–25 cm
f: Rostratulidae to the wetland jacanas.
SHORT-BILLED DOWITCHER
Related to the jacanas, the short-legged Limnodromus griseus
painted-snipes share their female- f: Scolopacidae RED KNOT DUNLIN
dominant mating system. This bird is Dowitchers are related to snipes and Calidris canutus Calidris alpina
from Old World tropical wetlands. are reddish colored in their breeding f: Scolopacidae f: Scolopacidae
plumage. The short-billed dowitcher Like many migratory waders, the Globally common, the dunlin
is confined to the Americas. Arctic-breeding red knot molts is typical of waders that breed
its distinctive summer plumage around the Arctic and flock to
to more drab colors in winter. the warmer south in the winter.
6½–7½ in
17–19 cm
10–10½ in JACK SNIPE long, slightly 8–8½ in
25–27 cm Lymnocryptes minimus upturned bill 20–21 cm
f: Scolopacidae
Birds of the snipe-woodcock group
have a long bill, short legs, and are
COMMON SNIPE camouflaged by their plumage. This SANDERLING
Gallinago gallinago Old World bird is the smallest Calidris alba
f: Scolopacidae in this group.
f: Scolopacidae
Typical of waders, chicks of This sandpiper breeds within
this globally widespread species rufous (reddish) the Arctic Circle, farther
are active soon after hatching. underparts north than other waders. In
Unlike other waders, snipes during breeding winter, it flocks on sandy
feed their chicks. shores farther south.

black and white 16–17½ in 7–7½ in

upperparts 40–44 cm 18–19 cm

HUDSONIAN GODWIT 14½–16½ in BLACK-TAILED GODWIT RED-NECKED PHALAROPE

Limosa haemastica 37–42 cm Limosa limosa Phalaropus lobatus
f: Scolopacidae f: Scolopacidae f: Scolopacidae
This is one of two American Typical of godwits, this Old World This species has a breeding range that includes
godwits, and is named after its species has a slightly upturned bill. much of the Arctic region. Females are brightly
breeding habitat, which includes It ranges far inland to forage on colored and attract males with displays during
the shores of Hudson Bay. grassland, heath, and pasture. breeding; males care for offspring.
 16–16½ in 7–8 in 451
40–42 cm 18–20 cm
18–26 in
45–66 cm

LONG-BILLED CURLEW WHIMBREL BUFF-BREASTED SANDPIPER

Numenius americanus Numenius phaeopus Tryngites subruficollis
f: Scolopacidae f: Scolopacidae f: Scolopacidae
A typical curlew, this American A mid-sized curlew, the whimbrel has This wader breeds in the tundra
species is a large wader with a long, a distinctive trilling call and breeds in of N. America and extreme
down-curved bill for probing circumpolar regions. Some individuals E. Siberia. It winters in the
deeply in mud for invertebrate prey. winter as far away as Australia. grasslands of S. America.

10½–11½ in 9–10 in large neck ruff in

27–29 cm 23–25 cm breeding male

COMMON LESSER
REDSHANK YELLOWLEGS
Tringa totanus Tringa flavipes
f: Scolopacidae f: Scolopacidae
Most waders breed near fresh Yellowlegs belong to the
water, but the Old World shank group. This species
common redshank—named breeds in forests in
after its colored legs—may Alaska and Canada,
sometimes breed on and overwinters
salt marshes. in the Caribbean.

10–11 in
26–28 cm

8–12 in
20–30 cm
10–12 in
26–30 cm
RUFF
Calidris pugnax
AMERICAN WOODCOCK f: Scolopacidae
Scolopax minor Males of this Old World marsh and
WANDERING TATTLER f: Scolopacidae meadow species transform during
Tringa incana Like other woodcocks and snipes, breeding: they lose their gray winter
f: Scolopacidae this American bird is superbly plumage and take on a rufous (reddish
This shank species breeds camouflaged and its eyes are brown) and black coloration,
in Alaska, during which its placed high, enabling all-around with a showy neck ruff.
breast is barred. It winters vision for predators.
farther south along the
Pacific coastline of America.

7–8 in
18–20 cm

10½–11½ in
27–29 cm

black-spotted 9–9½ in 5½–6½ in

chest and belly 22–24 cm 14–16 cm
in summer

SPOTTED SANDPIPER RUDDY TURNSTONE SPOON-BILLED SANDPIPER WHITE-BELLIED SEEDSNIPE

Actitis macularius Arenaria interpres Calidris pygmaea Attagis malouinus
f: Scolopacidae f: Scolopacidae f: Scolopacidae f: Thinocoridae
Like the related common sandpiper This bird, which breeds across Like spoonbills, this wader from Four species of short-billed,
reddish (A. hypoleucos) from Eurasia, this the northern hemisphere, is E. Asia uses its distinctive spoonlike herbivorous seedsnipe live in the
legs American sandpiper has a short bill named for the way it flips bill to sweep shallow water open habitats of S. America. This
for pecking food from dry ground. objects to search for prey. for invertebrate prey. species is confined to the southern tip.
 WA D E R S , G U L L S , A N D AU K S black-bordered
creamy throat
452
7½–8½ in 7½–9½ in 9½–11 in
19–21 cm 19–24 cm 24–28 cm
BI R D S • WA DER S , GU L L S , A N D AU K S

BLACK-WINGED PRATINCOLE
Glareola nordmanni
f: Glareolidae
Like other pratincoles, this species
is migratory. It breeds in E. Europe and
C. Asia and overwinters in Africa.

CREAM-COLORED AUSTRALIAN
COURSER PRATINCOLE
Cursorius cursor Stiltia isabella forked tail
f: Glareolidae f: Glareolidae 9–10 in
Often nocturnal, coursers are This pratincole from Australia 23–26 cm
inconspicuous, long-legged and Indonesia usually stays
ground birds, similar to close to fresh water, but has 10½–11 in 6½–7½ in
plovers. This is a migratory special glands that enable it 27–28 cm 17–19 cm
species from Asia and Africa. to drink salt water, too.
HEUGLIN’S
COURSER LITTLE PRATINCOLE COLLARED PRATINCOLE
Rhinoptilus cinctus Glareola lactea Glareola pratincola
f: Glareolidae f: Glareolidae f: Glareolidae
Most coursers are This small species is from Like most pratincoles,
restricted to arid deserts S. Asia. Like its relatives, this species from S. Europe
20–23½ in and scrub, but this African the little pratincole and Africa gathers in
50–60 cm species also ventures into feeds on insects in flight. large, noisy flocks on
woodland habitats. It has a forked tail. open wetlands.

18–18½ in
45–47 cm

SWALLOW-TAILED GULL GRAY GULL

Creagrus furcatus Leucophaeus modestus
f: Laridae 16½–17½ in 14–16 in f: Laridae
42–44 cm 36–41 cm
The only nocturnal gull, this Confined to Peru and Chile,
species breeds on the Galapagos this species breeds in the
Islands and overwinters Atacama Desert, one of the
in S. America. It feeds DOLPHIN GULL LAUGHING GULL driest regions of the world.
on fishes and squid. Leucophaeus scoresbii Leucophaeus atricilla
f: Laridae f: Laridae
10½–12½ in SABINE’S GULL This gull is famously An American hooded gull,
27–32 cm Xema sabini aggressive to other birds. It this species has a laughing
f: Laridae is confined to the southern call. It is a colonial breeder
A relative of the ivory gull, tip of S. America and on coastal estuaries and
this bird is an Arctic breeder, the Falkland Islands. salt marshes.
but it migrates huge
distances to overwinter in S.
America and Africa.
slender,
black bill
narrow
black collar
wedge-shaped
tail 11–12 in 13½–14½ in 16–18 in
28–30 cm 34–37 cm 40–45 cm

15–16 in
38–40 cm
COMMON
ROSS’S GULL BONAPARTE’S GULL BLACK-HEADED GULL SILVER GULL
Rhodostethia rosea Chroicocephalus philadelphia Chroicocephalus ridibundus Chroicocephalus novaehollandiae
f: Laridae f: Laridae f: Laridae f: Laridae
A distinctive pink gull, Ross’s gull A N. American masked gull, this Like other members of the masked gull Despite its different appearance,
breeds in swampy and wooded species breeds in wet coniferous group, this dark-headed gull becomes this Australian gull is related to
Arctic tundra, and winters forests in Canada and winters white- headed in winter. It is abundant the common black-headed gull. It
along coasts and at sea. along Caribbean coasts. in the northern hemisphere. visits human refuse sites to feed.
 large
16–16½ in 20–26 in 22–26 in white head
40–42 cm 50–67 cm 55–66 cm 453
slaty black back
and wings

BI R D S • WA DER S , GU L L S , A N D AU K S
MEW GULL PACIFIC GULL WESTERN GULL 25–31 in
Larus canus Larus pacificus Larus occidentalis 64–78 cm
f: Laridae f: Laridae f: Laridae
This northern hemisphere Belonging to a group of This large gull is found on GREAT
species, whose name derives from southern gulls with banded the Pacific coast of BLACK-BACKED GULL
its distinctive call, is also called tails, this large-billed Australian N. America. It nests in Larus marinus pink feet
the common gull. It breeds inland species cracks open shellfish by colonies, usually offshore f: Laridae
on moors, as well as on the coast. dropping them on rocks. on islands or rocks. The world’s largest gull, this
North Atlantic coastal bird is an
aggressive predator and preys on
24–27 in other seabirds and their chicks.
GREAT 62–68 cm
BLACK-HEADED GULL 18–20 in
Ichthyaetus ichthyaetus GLAUCOUS GULL 46–51 cm
f: Laridae Larus hyperboreus
One member of an Asian group f: Laridae
of black-headed gulls, this species A large, Arctic-breeding
breeds in Russia and overwinters on species, this coastal gull is much
Mediterranean and Indian ocean paler than most related
coasts. It is also known as Pallas’s gull. white-headed gulls found in the RING-BILLED GULL
northern hemisphere. Larus delawarensis
f: Laridae
22½–24 in Breeding in N. America
57–61 cm tricolored bill and overwintering in the
20½–23½ in Caribbean, this species has a dark
52–60 cm bill ring. It often forages on
agricultural land.

18–21 in
46–53 cm
white-edged eye
HERRING GULL
Larus argentatus
f: Laridae
Related to the lesser
black-backed gull (L. fuscus),
this species is often seen in
coastal towns in Europe and gray body
eastern N. America. 16½–18 in 16–17 in
42–45 cm 40–43 cm

HEERMANN’S GULL
SOOTY GULL Larus heermanni
Ichthyaetus hemprichii IVORY GULL f: Laridae
f: Laridae Pagophila eburnea This dark gull from
As with many warm-region gulls, f: Laridae N. America is actually related
this species from Asia and Africa This Arctic gull is rarely to northern white-headed gulls.
has dark plumage, which may found far from pack ice. It often forages with brown
have evolved as an adaptation It follows polar bears to pelicans and steals their food.
to strong sunshine. scavenge at their kills.

14–16 in 15–16 in
yellow legs 35–40 cm 38–40 cm

15½–17 in
39–43 cm
WHITE-EYED GULL RED-LEGGED KITTIWAKE BLACK-LEGGED KITTIWAKE
Ichthyaetus leucophthalmus Rissa brevirostris Rissa tridactyla
f: Laridae f: Laridae f: Laridae
This relative of the sooty gull is This kittiwake breeds only on islands in The world’s most abundant species of gull,
confined to the Red Sea region, the Bering Sea in the North Pacific and this colonial cliff-nesting bird is found
where it is threatened by oil spills. spends winter at sea further south. in the North Atlantic and Pacific oceans.
 WA D E R S , G U L L S , A N D AU K S 16–16½ in
white cheek stripe 9–9½ in
22–24 cm
454 40–42 cm

11–13 in INCA TERN

28–33 cm Larosterna inca
BI R D S • WA DER S , GU L L S , A N D AU K S

f: Laridae
A distinctively patterned BLACK TERN
bird, the Inca tern is Chlidonias niger
found in Peru and Chile, f: Laridae
where it breeds on This small freshwater tern breeds
rocky coastlines. in the wetlands of the northern
bright red hemisphere and winters in
WHITE TERN legs S. America and Africa.
Gygis alba
f: Laridae
This small, all-white tern from the islands of the
tropical Atlantic and Indian oceans is notable
for laying its eggs on bare branches in trees. black cap
GREATER CRESTED TERN
Thalasseus bergii
f: Laridae
18½–21½ in
This Old World tern belongs 47–54 cm
to a group of related crested
terns, characterized by a
LESSER black nape tuft.
CRESTED TERN
Thalasseus bengalensis 18–19½ in
f: Laridae 46–49 cm
In this relative of the greater
crested tern, the bill changes
from yellow to orange
during breeding. 14–14½ in
35–37 cm

9–9½ in
22–24 cm

LITTLE TERN black markings

on tips of CASPIAN TERN
Sternula albifrons underwings Hydroprogne caspia
f: Laridae f: Laridae
The Old World little tern 13–14 in The largest tern species,
belongs to a group of 33–36 cm this bird is found in most
coastal species with a white continents. It is a colonial
“blaze” above the eye. ground-nester, like
most sea terns.

SOOTY TERN
Onychoprion fuscatus
f: Laridae
This white-blazed sea tern breeds on tropical
SAUNDERS’S TERN 9–9½ in islands. It is also known as the wideawake long, black legs
Sternula saundersi 23–24 cm tern because of its noisy colonies. 13–15 in
f: Laridae 33–38 cm
A small tern from the Red Sea
and Indian Ocean, Saunders’s 12–12½ in
tern used to be considered 30–32 cm
a race of the little tern. BRIDLED TERN ROSEATE TERN
Onychoprion anaethetus Sterna dougallii
f: Laridae f: Laridae
This tern of tropical and subtropical As with related sea terns, the roseate
12½–13½ in regions has a white eye-blaze, similar tern’s black cap fades in winter. This
32–34 cm to little and sooty terns. It spends bird is found mostly in the southern
much of its time at sea. hemisphere and is migratory.

13–14 in
12–12½ in 33–35 cm
30–32 cm
WHITE-CHEEKED TERN BLACK-NAPED TERN ARCTIC TERN
Sterna repressa Sterna sumatrana Sterna paradisaea
f: Laridae f: Laridae f: Laridae
This species from the Red Sea and Found on the Indian and This tern migrates the greatest
Indian Ocean can be identified by Pacific oceans, the black-naped distance of any animal: from its Arctic
its plumage, which is darker than tern nests in small colonies, breeding grounds to Antarctica.
that of other gray terns. usually apart from other terns. It feeds on fishes and crustaceans.
 16–20 in stout, 18–20 in
40–50 cm hooked bill 46–51 cm
16–18 in
40–45 cm
455
BROWN NODDY
Anous stolidus

BI R D S • WA DER S , GU L L S , A N D AU K S
f: Laridae
Noddies are dark or white 20½–21 in
tropical members of the tern 52–54 cm
family. The largest species, the POMARINE JAEGER
brown noddy is widespread Stercorarius pomarinus
around the world. f: Stercorariidae
BLACK SKIMMER brownish- Jaegers (also called skuas in Europe)
Rynchops niger gray body are aggressive gull-like birds. This
f: Laridae Arctic species will kill and eat other
Skimmers are the only birds with seabirds and even attack humans
projecting lower bill mandibles, that threaten its nest.
used to skim water for fishes. 19–21 in
This species is found in 48–53 cm
PARASITIC JAEGER
N. and S. America. Stercorarius parasiticus
SOUTH POLAR SKUA f: Stercorariidae
Stercorarius maccormicki This is the most common
f: Stercorariidae Arctic species of jaeger.
This large bird, with a Like most of its relatives,
reputation for attacking other it mobs other seabirds
seabirds, is one of the few 16–18 in to steal their prey.
41–46 cm
waders to breed on the
Antarctic coast.
LONG-TAILED JAEGER 9½–10 in
Stercorarius longicaudus 24–25 cm
f: Stercorariidae
14½–15½ in Like other species, this
37–39 cm smallest member of the jaeger
family is migratory. It breeds
9½–10½ in
in circumpolar regions and 24–27 cm
winters farther south.
6½–7½ in
17–19 cm MARBLED MURRELET
Brachyramphus marmoratus
DOVEKIE f: Alcidae
Alle alle This small American auk nests in
f: Alcidae trees in coniferous woods. Fledged
RAZORBILL This tiny auk breeds chicks leave the nest at night
Alca torda on Arctic islands and and head for the sea.
f: Alcidae winters at sea farther
The North Atlantic razorbill CRESTED AUKLET south. It feeds on tiny
has a flattened, white-barred Aethia cristatella fishes and crustaceans. dark brown to
bill. As with other auks, its f: Alcidae black head
pointed eggs cannot roll off Like other auklets, this North
cliff-top breeding sites. Pacific species feeds on planktonic
crustaceans. Pairs anoint themselves
with secretions from their backs 12–12½ in 11–11½ in
during courtship. 30–32 cm 28–29 cm

12–14 in
30–36 cm RHINOCEROS AUKLET
Cerorhinca monocerata
f: Alcidae
PIGEON GUILLEMOT BLACK GUILLEMOT This North Pacific relative of puffins
Cepphus columba Cepphus grylle shares their burrow-nesting habits,
f: Alcidae f: Alcidae and is named for the hornlike bill
This North Pacific auk, fully adapted This species of northern projection in breeding adults.
to cold conditions, cannot migrate N. American and Eurasian
south through warmer waters, just coasts breeds in sparser
as southern hemisphere penguins colonies than other auks and
cannot move northward. winters mainly inshore. 15–16 in
38–41 cm

white wing patch 10–11½ in 13½–14 in

26–29 cm 34–36 cm

ATLANTIC PUFFIN
Fratercula arctica TUFTED PUFFIN
red feet f: Alcidae Fratercula cirrhata COMMON MURRE
A small, North Atlantic f: Alcidae Uria aalge
member of the auk family, Like its Atlantic relative, f: Alcidae
this species—like other this larger Pacific puffin A typical diving member of the
puffins—nests in colonies, catches small fishes and auk family, this bird breeds on
in burrows, usually can hold many at a time, North Atlantic and Pacific coasts,
below turf. crosswise, in its bill. and winters at sea.
456 SA N D G RO U S E
Sand-colored plumage camouflages sociable birds that gather in flocks to make early PHYLUM CHORDATA
these birds in their desert habitats, morning, or sometimes evening, forays to drinking
BIRDS • SANDGROUSE

CLASS AVES

where they are well adapted for living holes, often traveling considerable distances. They ORDER PTEROCLIFORMES

in an extremely dry environment. feed exclusively on seeds. FAMILIES 1

SPECIES 16

With their rounded body and short legs, sandgrouse wat e r c a r r i e r s

might be mistaken for partridges until they take Sandgrouse breed during the rainy season to
flight, darting quickly and acrobatically in the take advantage of seed harvests. Their nest is
air. These birds, the Pterocliformes, are found nothing more than a shallow depression on the
in arid areas of Asia, Africa, Madagascar, and ground. Both parents incubate and care for their
southern Europe. They are not related to the young. Remarkably, the males supply water to
subarctic grouse species but are more closely their chicks, which are normally raised far from
related to pigeons. water sources. The male’s belly feathers soak
Sandgrouse have long pointed wings and all species up and retain moisture when the bird visits a
have cryptic plumage—mottled on the back, but drinking hole. Back at the nest, the chicks drink
Namaqua sandgrouse drink at a
often boldly marked with brown or white stripes from the parent’s watering hole. Like all sandgrouse, they
or blotches on the head and underparts. They are soaked feathers. gather in large flocks to confuse predators.

PALLAS’S 12–16 in 10–11 in black and white bands on

SANDGROUSE 30–41 cm 25–28 cm forecrown (male only)
Syrrhaptes paradoxus
f: Pteroclidae barred buff
One of two C. Asian plumage
sandgrouses, this large species
has feathered toes, a long tail,
and a long flight feather
on each wing.

DOUBLE-BANDED
SANDGROUSE
Pterocles bicinctus
f: Pteroclidae
long, pointed tail A bird of southern African
savanna and open woodland,
this is one of several sandgrouse
species in which the males have
distinctive belly stripes.
CHESTNUT-BELLIED SANDGROUSE
Pterocles exustus 12–13 in
f: Pteroclidae 31–33 cm
white-barred
With a range that extends from Senegal to wings
Kenya, and east as far as India, this bird of
open desert gathers in huge flocks.

10½–12 in
27–30 cm

black
breast band

9½–10 in
24–26 cm
CROWNED SANDGROUSE LICHTENSTEIN’S SANDGROUSE
Pterocles coronatus Pterocles lichtensteinii
f: Pteroclidae f: Pteroclidae
This yellow-throated sandgrouse is This small bird is less gregarious
found from the stony desert of the than other species of sandgrouse. It inhabits
Sahara to Pakistan. It can tolerate high bushy scrubland, and semidesert regions
temperatures and even brackish water. from N. and E. Africa to Pakistan.
P I G E O N S A N D D OV E S 457
This is a highly successful group of without tilting back their head, because of a PHYLUM CHORDATA
seed-eating and fruit-eating birds pumping action in their food-pipe. This allows

BIRDS • PIGEONS AND DOVES

CLASS AVES

that are distributed almost worldwide them to drink continuously, which is a particular ORDER COLUMBIFORMES

outside the coldest regions. benefit in arid environments. They can store FAMILIES 1

food in their bill, and they feed their chicks on SPECIES 344

After the parrots, pigeons and doves form the a secretion from the bill. This secretion is similar
largest group of vegetarian tree-perching birds: to mammalian milk. D E B AT E
the order Columbiformes. But, whereas parrots A MODIFIED PIGEON?
have a heavy, hooked bill for cracking large nuts, t h r e at s a n d e x t i n c t i o n s
pigeons have a less robust bill for feeding on smaller The success of many pigeon species is due to high Mid-19th-century scientists placed
seeds and grain. Some tropical groups, such as rates of reproduction. However, some species are the extinct flightless dodo from
Mauritius in the pigeon order, and
the Indo-Pacific fruit doves, are specialized fruit- threatened by humans and others have already
recent analysis indicates that it was
eaters of the rainforest canopy. been lost. The 17th-century demise of the dodo, related to the Nicobar pigeon. So the
Most birds in this order are short-legged and a a vulnerable flightless species, was the result of dodo was really a modified pigeon,
few spend almost all of their time on the ground. human impact. And the passenger pigeon, once with Indo-Pacific ancestors that
Pigeons and doves are unusual among birds one of North America’s commonest birds, might already have been flightless.
in being able to suck up water when drinking was hunted to extinction in the 1900s.

EUROPEAN TURTLE DOVE black and white 10–11 in 10–10½ in red patch
Streptopelia turtur striped neck patch 26–28 cm 25–27 cm around eyes
f: Columbidae
With a range from Africa to
Eurasia, turtle doves are small,
slim relatives of Columba pigeons. white-speckled
Many, such as this western wings
species, have distinctive
neck markings.

LAUGHING DOVE
Streptopelia senegalensis
f: Columbidae
Common in villages and oases in Africa
and southern Asia, this turtle dove is
named after its distinctive chuckling song.
15–17 in
38–43 cm

10–11 in 13–15 in
BROWN CUCKOO-DOVE 26–28 cm 33–38 cm
Macropygia amboinensis
f: Columbidae NAMAQUA DOVE SPECKLED PIGEON
Long-tailed Macropygia doves are Oena capensis Columba guinea
cuckoolike birds of Indo-Pacific f: Columbidae f: Columbidae
rainforests. Like others, this This long-tailed, ground-feeding This large African pigeon is common in open
species from the Moluccas species belongs to a group of small country south of the Sahara, and it often
(Indonesia), New Guinea, and African wood doves. Its range extends gathers around towns and villages.
Australia has many races. into Madagascar and Saudi Arabia.

12½ in 15–17 in 12–14 in

32 cm 38–43 cm 31–35 cm

12–14 in
31–35 cm

PINK PIGEON
Nesoenas mayeri ROCK PIGEON DOMESTIC PIGEON
f: Columbidae WOODPIGEON Columba livia Columba livia
Perhaps related to turtle doves, Columba palumbus f: Columbidae f: Columbidae
this rare bird is restricted to f: Columbidae True wild pigeons inhabit cliffs and Domesticated and feral descendants of
Mauritius. Its once threatened This large W. Eurasian pigeon is hedges. In its natural environment, this the rock pigeon are found in urban areas
population recovered due to a successful commonly found in woods and farmland, species lives in mountainous regions worldwide. These birds have variable
captive-breeding program. and often enters parks and gardens. of Europe and Asia. plumage patterns.
 PIGEONS A N D DOV E S WOMPOO FRUIT DOVE
Ptilinopus magnificus
458 f: Columbidae
Fruit doves are multicolored relatives of
13–16 in 6½–9 in imperial pigeons (Ducula sp.).
33–40 cm 17–23 cm This large species from New Guinea
BIRDS • PIGEONS AND DOVES

and Australia lives in the

rainforest canopy.

INCA DOVE
Columbina inca
f: Columbidae
From arid regions of the yellow markings
S. US and C. America, this on wings
species belongs to a group of
mostly drab-colored tropical
American ground-doves.
8 in
20 cm

NICOBAR PIGEON DIAMOND DOVE

Caloenas nicobarica Geopelia cuneata
f: Columbidae f: Columbidae
Possibly related to the extinct A small, nomadic bird from
Mauritian dodo, this pigeon the dry interiors of Australia, 11½–22 in
29–55 cm
inhabits coastal regions and this dove occurs in groups,
island forests from Malaysia flocking in large numbers
to New Guinea. at watering holes.

10–12 in WHITE-TIPPED
25–31 cm DOVE
Leptotila verreauxi dark green tail
f: Columbidae 14 in 9–13½ in
35 cm 23–34 cm
A tropical relative of
the mourning dove, this
species is widespread
in C. and S. America,
and reaches as far
north as Texas.
12 in
30 cm

dark pink head

and breast
MINDANAO SULAWESI MOURNING
BLEEDING-HEART GROUND DOVE DOVE
PIGEON Gallicolumba tristigmata Zenaida macroura
Gallicolumba criniger f: Columbidae f: Columbidae
f: Columbidae From the forests of Sulawesi, So called because of its
The five species of Philippine Indonesia, this terrestrial bird mournful call, this long-tailed
bleeding-heart pigeons are named is related to Philippine bleeding- dove occurs in open country
emerald- for their blood-red breast patch. heart pigeons, and perhaps to throughout N. and C. America,
green wings This bird inhabits the southern various doves of arid Australia. and the Caribbean.
and back islands of the archipelago.

18 in 15½–17½ in
45 cm 39–44 cm

16–18 in
40–46 cm PIED
IMPERIAL PIGEON
Ducula bicolor
f: Columbidae
9–11 in Imperial pigeons are large,
23–28 cm fruit-eating rainforest birds.
This species from S.E. Asia and
Australasia has a white plumage
that is often stained due
to its diet.

EMERALD DOVE TOPKNOT PIGEON GREEN IMPERIAL PIGEON

Chalcophaps indica Lopholaimus antarcticus Ducula aenea
f: Columbidae f:Columbidae f: Columbidae
An iridescent green bird, this A large, hawklike Found in rainforest canopy,
ground-feeding species occurs in pigeon from E. Australia, from India to S.E. Asia, this large
rainforests from India to islands this species is named for the bird has a deep, booming call
of the southwest Pacific. It has a double crest of feathers on and has a diet consisting
diet of fruits and seeds. its forecrown and crown. mainly of fruits.
 VICTORIA
CROWNED PIGEON side-flattened
Goura victoria tail 459
f: Columbidae
Crowned pigeons are the largest
of the pigeons. This bird from N. New

BIRDS • PIGEONS AND DOVES

Guinea is distinguished from the southern
29–30 in crowned pigeon by its white-tipped crest.
74–75 cm 14–15 in
36–38 cm

WONGA PIGEON 18–20 in

Leucosarcia melanoleuca 45–50 cm
f: Columbidae
Confined to eastern
Australia, this distinctively
patterned pigeon inhabits
woodland and scrub
from S. Queensland
to Victoria.

PHEASANT PIGEON
Otidiphaps nobilis
f: Columbidae
Recent research suggests this
New Guinean ground pigeon belongs
to a group that includes crowned pigeons
and perhaps even the extinct dodo.
fanlike crown
10½–12 in
KEY WEST 27–31 cm
SOUTHERN QUAIL-DOVE
CROWNED PIGEON Geotrygon chrysia
Goura scheepmakeri f: Columbidae
f: Columbidae Quail-doves are birds
This species inhabits the forests of of tropical American forests.
the southern part of New Guinea. This iridescent species
It has bluish-gray upperparts, 30 in occurs in the Caribbean,
a maroon chest, and a lacy crest. 75 cm including the Bahamas.
10–11 in
25–28 cm

bluish-gray maroon chest

plumage
AFRICAN
GREEN PIGEON
Treron calvus
f: Columbidae
More than 20 species of Treron
pigeons occur in the tropical regions
of Africa and Asia. This species is
13–14 in widespread south of the Sahara.
33–36 cm

COMMON BRONZEWING
Phaps chalcoptera grayish body
f: Columbidae
Bronzewings are fast-flying
Australian pigeons that feed on the
ground. They have iridescent wing
patches, which are particularly
extensive in this widespread
woodland species.

8–9 in
20–22 cm

12–14 in
31–35 cm

white wing iridescent

patch wing patch
SPINIFEX PIGEON CRESTED PIGEON
Geophaps plumifera Ocyphaps lophotes
f: Columbidae f: Columbidae
This Australian bronzewing pigeon is found One of several species of Australian bronzewing
in arid, rocky habitats with an abundance pigeons, this species is widespread in open
of spinifex grass, in which it nests. country throughout the continent.
460 PA R RO T S A N D C O C K AT O O S
Most parrots are birds of tropical scatters light on yellow pigment. The Australasian PHYLUM CHORDATA
forests, although a few favor open cockatoos—parrots with a distinctive erectile
BIR DS • PAR ROTS A ND COCK ATOOS

CLASS AVES

habitats. This order, the Psittaciformes, crest—lack this feather texture and so lack ORDER PSITTACIFORMES

is diverse and often colorful. green and blue colors. FAMILIES 4

Australasia’s high parrot diversity, which SPECIES 398

The most recognizable feature of a parrot is includes the brush-tongued, nectar-feeding

its decurved bill. Both mandibles of the bill lorikeets, suggests that the group originated
articulate with its skull: the upper mandible can in this part of the world. The most primitive
move up, just as the lower can move down. This parrots, the kea and the nocturnal, flightless
allows the bird not only to crack open hard seeds kakapo, are both found in New Zealand.
and nuts, but also to use its bill for gripping when
clambering through trees. The legs are strong soci a l speci es
and the feet, with two forward-facing and two Parrots are sociable, often living in large flocks,
backward-facing toes, can grasp and manipulate and almost all species form strong pair-bonds.
foodstuffs. Bright colors are common, often Their engaging personalities make them popular
Green-winged macaws socialize at
in both sexes, with green plumage predominating. as pets, but the international cagebird trade has a clay-lick, which provides sodium in an
This color is the result of feather structure, which driven many species to the edge of extinction. area where the mineral is very limited.

KEA barred green

Nestor notabilis plumage
f: Psittacidae
An opportunistic alpine 23½ in 14 in
omnivore, the kea eats 60 cm 36 cm
carrion and live petrel
chicks. It belongs to an
ancient group of parrots
in New Zealand that
includes the kakapo.
19 in
48 cm

KAKAPO GALAH
short tail Strigops habroptila Eolophus roseicapilla
f: Psittacidae f: Psittacidae
The only flightless parrot, The only cockatoo with a deep
this large, nocturnal bird pink neck and underparts,
occurs in the small islands this small relative of the
off New Zealand. Males white cockatoos is widespread
5–6 in 43/4–6 in make booming calls throughout Australia in areas
13–15 cm 12–15 cm to attract females. with scattered trees.

19½ in 20–24 in
49 cm 50–61 cm

VERNAL HANGING
PARROT BLUE-CROWNED SULPHUR-CRESTED
Loriculus vernalis HANGING PARROT COCKATOO
f: Psittacidae Loriculus galgulus Cacatua galerita RED-TAILED BLACK COCKATOO
Hanging parrots are Indo-Pacific f: Psittacidae f: Psittacidae Calyptorhynchus banksii
birds, even though recent genetic Hanging parrots, unusually among birds, White cockatoos are screeching f: Psittacidae
evidence suggests affinities with sleep upside down. Like other such parrots, parrots with a range that extends from One of several Australian black cockatoos with
African lovebirds. This species is this small forest bird from S.E. Asia has a Indonesia to the Pacific Rim. This species colored tail panels, this glossy species has a
found from India to Thailand. short tail and mainly green plumage. occurs in New Guinea and Australia. typical wailing call and ponderous wing beats.
 10 in 12 in
25 cm 30 cm
461
BLACK-WINGED
LORY
12½ in 12 in Eos cyanogenia
32 cm 30 cm f: Psittacidae
The Eos lories are vividly red
and violet Indonesian birds.
This is the easternmost
species, restricted to the
area around Geelvink
CHATTERING LORY DUSKY LORY Bay, New Guinea.
Lorius garrulus Pseudeos fuscata
f: Psittacidae f: Psittacidae
COCKATIEL One of a group of green-winged The patchy brown coloration
Nymphicus hollandicus red lories from New Guinea and of this species is unique among bright
f: Psittacidae surrounding islands, this species lories. This bird occurs in New scarlet
A bird of Australia’s dry interior, is from the Moluccas. Guinea and nearby islands. head
this species resembles a parakeet,
but its DNA shows it is a
miniature cockatoo.

9½ in
24 cm
blue band
7 in across back
18 cm and chest
OLIVE-HEADED
LORIKEET VARIED LORIKEET
Trichoglossus euteles Psitteuteles versicolor
f: Psittacidae f: Psittacidae
Found on the island of Timor, A small lorikeet from northern
this long-tailed relative of the Australian woodlands, this species
rainbow lorikeet has bright green nests in eucalyptus tree cavities, like
plumage, typical of many parrots. many other parrots in its habitat.

RAINBOW LORIKEET
Trichoglossus haematodus
f: Psittacidae
A variable species,
this bird is found in
pale green many habitats with
streak on nectar-giving blossom.
green back Its range includes most of
Australasia and the islands
of the southwest Pacific.

17 in 10–12 in
43 cm 25–30 cm

mainly
purple rump green body
13–15½ in
33–39 cm

AUSTRALIAN
KING PARROT
Alisterus scapularis
f: Psittacidae
King parrots are rainforest
birds from tropical
Australasia and represent an ECLECTUS PARROT
evolutionary link to Asian Eclectus roratus
parakeets. This species f: Psittacidae
occurs in E. Australia. Males and females of this parrot
are so different they were originally
classified as separate species. They are
7 in 10½ in found in tropical Australasian rainforests.
18 cm 27 cm

RED-FRONTED AUSTRALIAN 13½–15 in

PARAKEET RINGNECK 34–38 cm
Cyanoramphus novaezelandiae Barnardius zonarius
BUDGERIGAR f: Psittacidae f: Psittacidae
Melopsittacus undulatus Small parakeets with colored Named for its yellow
f: Psittacidae foreheads have diversified in neck ring, this Australian
A nomadic Australian bird, this small the southwest Pacific, and bird has black- and
parrot from arid regions flocks to include the only parakeets— green-headed forms.
watering holes. Despite being a seed-eater, like this species—from It is widespread
it is related to nectar-feeding lorikeets. New Zealand. in woodland.
 PA R ROT S A N D C O C K ATO O S black mask
462

14 in 8 in 18½ in 12 in
BIR DS • PAR ROTS A ND COCK ATOOS

36 cm 20 cm 47 cm 30 cm

yellow belly

TURQUOISE PARROT EASTERN ROSELLA

Neophema pulchella Platycercus eximius
f: Psittacidae f: Psittacidae
A member of a group of small, Rosellas are broad-tailed parrots
green grass parrots from with a range from Australia to
Australia, the turquoise parrot nearby Pacific islands. A typical
is an open woodland bird from rosella, this variable E. Australian
the southeast of the continent. bird has white cheeks.

RED-FAN PARROT ROSE-RINGED MASKED SHINING PARROT

Deroptyus accipitrinus PARAKEET Prosopeia personata blue plumage
f: Psittacidae Psittacula krameri f: Psittacidae and upperparts
Possibly related more closely f: Psittacidae One of the three species of
to macaws than to other This is the most widespread Asian shining parrots restricted to Fiji,
short-tailed parrots, this parakeet that reaches westward this parrot is declining in numbers
S. American parrot raises a into N. Africa. This species has also due to loss of its native forest.
red neck ruff when excited. been introduced in Europe.

15–16½ in 16–18½ in
38–42 cm 40–47 cm
gray-black
undertail

16 in
white face 40 cm

13 in
33 cm
SUPERB PARROT
Polytelis swainsonii
f: Psittacidae
PRINCESS PARROT This S.E. Australian bird
Polytelis alexandrae belongs to a group of long-tailed
f: Psittacidae Australian parrots and breeds in
A nomad from central red gum forests. It is rapidly
Australia, this bird follows declining in number.
watercourses and gathers
near spinifex grass. It often
breeds in small colonies yellow
in eucalyptus trees. undertail

pale gray
underparts

6 in 6½–7 in
15 cm 17–18 cm

YELLOW-COLLARED ROSY-FACED 14–14½ in

LOVEBIRD LOVEBIRD 35–37 cm
GRAY PARROT Agapornis personatus Agapornis roseicollis
Psittacus erithacus f: Psittacidae f: Psittacidae BROWN-NECKED PARROT
f: Psittacidae Lovebirds are small African parrots This social lovebird from dry Poicephalus robustus
A native of African rainforests, that live in small flocks. Unlike woodland and semidesert f: Psittacidae
this highly intelligent species is an most parrots, this Tanzanian habitats of S.W. Africa The largest of a group of mainly green-gray
red tail accomplished mimic. This has led to species builds a nest—a domed habitually gathers near parrots from Africa, this bird occurs in forests
its exploitation in the bird trade. structure in a tree cavity. watering holes. from Gambia to the Cape of Good Hope.
 white facial 463
patch with lines
of black feathers

BIR DS • PAR ROTS A ND COCK ATOOS

15 in 16 in
38 cm 40 cm

BLUE-FRONTED PARROT ST. VINCENT PARROT

Amazona aestiva Amazona guildingii
34 in f: Psittacidae f: Psittacidae
85 cm A member of a large group Several Caribbean Amazona parrots
of mostly green parrots, this are faced with extinction. This
BLUE-AND-YELLOW powerful bill species occurs in open forest large species is being conserved by
MACAW habitats of central-eastern an ongoing breeding program
Ara ararauna S. America. in its native island of St. Vincent.
f: Psittacidae
Macaws are large, long-tailed
parrots with sparsely feathered 43/4–5½ in
12–14 cm
facial patches. One of two blue and
yellow species, this bird comes
from northern S. America.

31–35 in 9½–11 in
79–89 cm 24–28 cm

22–23½ in
55–60 cm

BLUE-HEADED PARROT PACIFIC PARROTLET

bright yellow Pionus menstruus Forpus coelestis
feathers on f: Psittacidae f: Psittacidae
blue wings A relative of the Amazona Parrotlets are tiny green American
parrots, this small parrot is parrots; only New Guinean pygmy
RED-FRONTED SCARLET MACAW common in lowland forests parrots are smaller. This species
MACAW Ara macao from Costa Rica to Bolivia. occurs in western Ecuador and Peru.
Ara rubrogenys f: Psittacidae
f: Psittacidae Like other macaws, this
This small macaw is restricted noisy, flocking bird has a
to the arid scrub of central heavy bill for breaking nuts 12 in white eye-ring
Bolivia. Its small population and palm fruit. Its range 30 cm
is threatened by habitat loss extends from S. Mexico
and wildlife trafficking. to C. Brazil.
JANDAYA PARAKEET
31/4 ft long, red tail Aratinga jandaya
1m f: Psittacidae
Aratinga is a genus
HYACINTH MACAW of predominantly green
Anodorhynchus hyacinthinus 8–10 in
20–25 cm “mini-macaws” found in
f: Psittacidae northeastern Brazil. Some
Unlike other macaws, this giant Brazilian have extensive splashes
parrot has its facial patch reduced to an of golden coloration.
eye-ring—similar to that of the related
Aratinga parakeets.
10 in
25 cm

17½–18 in 11½ in red belly

44–46 cm 29 cm

MAROON-BELLIED
YELLOW-CHEVRONED PARAKEET
PARAKEET MONK PARAKEET Pyrrhura frontalis
BURROWING PARAKEET Brotogeris chiriri Myiopsitta monachus f: Psittacidae
Cyanoliseus patagonus f: Psittacidae f: Psittacidae Relatives of Aratinga, most
f: Psittacidae Native to central S. America, This colonial breeder from Pyrrhura parakeets have
A Patagonian relative of macaws, escaped cagebirds of this temperate S. America is prominent splashes of
this colonial breeder nests in species have established the only parrot to build a maroon or red, such
tunnels in earth banks. It forms feral populations in stick nest, often a large as this species from
faithful pair-bonds, unlike most birds. warmer parts of the US. communal structure. eastern S. America.
464 T U R AC O S
Confined to Africa, this group includes than indicative of any relationship. They live PHYLUM CHORDATA
the strikingly colorful turacos, which in forests, subsisting largely on seasonal fruit.
BIRDS • TURACOS

CLASS AVES
are largely green, blue or purple, the Although their bill is remarkably short, a long ORDER MUSOPHAGIFORMES

dull, grayish go-away birds (named tail helps them maintain balance when reaching FAMILIES 1

for their distinctive nasal calls), and for food in the tree canopy. They move heavily SPECIES 23

through foliage and fly short distances across

the plantain-eaters. forest clearings. Turacos have vividly colored red crest
In the order Musophagiformes, these birds plumage that derives from green and red pigments,
share certain features with cuckoos, such as foot which are uniquely copper-based. Most species in 16–17 in
structure, which appears to be coincidental rather this group have a short, thick, upstanding crest. 40–43 cm

HARTLAUB’S RED-CRESTED TURACO

TURACO Tauraco erythrolophus white face
Tauraco hartlaubi f: Musophagidae
f: Musophagidae One of few red-crested turacos, this
A blue-crested member of species has a less bright body plumage KNYSNA TURACO
the green turaco group, this than others in the green group. It is Tauraco corythaix
species is found in highland commonly found in the evergreen f: Musophagidae
forest in E. Africa. forests and woodlands of Angola. A South African relative of the
green turaco, this species shares
the white eye-stripes, but differs
17 in long tail in having a white-tipped crest.
43 cm

18–18½ in
45–47 cm

bright crimson
flight feathers

16 in 20–21½ in 18–20 in
40 cm GREEN TURACO 51–54 cm 45–50 cm
Tauraco persa
f: Musophagidae
With a range from Senegal
RUSPOLI’S TURACO to Angola, this is the most
Tauraco ruspolii widespread of a group of
f: Musophagidae green turacos with VIOLET TURACO
A distinctively white- crimson wing-feathers
Musophaga violacea
crested green turaco, this visible in flight.
f: Musophagidae
species has one of the One of two glossy,
most restricted ranges of violet-colored turacos, this
16–17 in
any turaco, being confined 40–43 cm species ranges through
to forests in S. Ethiopia. forests of W. and C. Africa.
ROSS’S TURACO
Musophaga rossae fanlike crest
f: Musophagidae
The second-largest turaco, this yellow bill
species is distinguished by its with red tip
18½–20 in upright, red crest. It is an E. African
47–50 cm counterpart of the violet turaco.
20 in
50 cm

19 in blue body
48 cm

BARE-FACED
GO-AWAY BIRD
Corythaixoides personatus
f: Musophagidae
Like other go-away birds, 28–30 in
this E. African species is
70–75 cm EASTERN GRAY
GRAY GO-AWAY BIRD PLANTAIN-EATER
Corythaixoides concolor found in savanna GREAT BLUE TURACO Crinifer zonurus
f: Musophagidae woodland. It raises its Corythaeola cristata f: Musophagidae
crest when excited. f: Musophagidae
Named after its call, “kay- Like the related go-away birds,
waaay,” this southern African long, This fan-crested turaco from plantain-eaters are drably colored.
species is a typical go-away bird broad tail W. and C. Africa is the largest and most Despite their name, they favor figs, not
with a pointed, shaggy crest. distinctive member of the turaco family. plantains. This species is from E. Africa.
H OAT Z I N
HOATZIN
Opisthocomus hoazin
f: Opisthocomidae spiky crest 465
Of uncertain taxonomic affiliation,
the hoatzin is a plant-eater of
The hoatzin is an enigmatic South PHYLUM CHORDATA S. American riverine forests. Its
American bird that has an entirely

BIRDS • CUCKOOS
CLASS AVES chicks have claws on their wings
for clambering through branches.
vegetarian diet of leaves. ORDER OPISTHOCOMIFORMES
FAMILIES 1
24–26 in
Some features of the hoatzin, such as strong SPECIES 1
61–66 cm
feet and a short, curved bill, are similar to
those of gamebirds. However, the hoatzin also shares features with cuckoos, a
relationship that appeared to be confirmed by early DNA (deoxyribonucleic acid)
studies. Later research has thrown doubt on this—leaving the hoatzin in a long neck
group of its own, the only representative of the order Opisthocomiformes.

C U C KO O S
Cuckoos are rather short-legged, Almost all these ground cuckoos build nests PHYLUM CHORDATA
long-winged, mostly softly feathered and raise their own young, as do their Old CLASS AVES

birds that include dull, grayish species World counterparts, such as the tropical coucals ORDER CUCULIFORMES

as well as a small group with patches and couas. Cuckoos that lay eggs in nests of FAMILIES 1
SPECIES 149
of vivid emerald green. other bird species are called “brood parasites.”
Remarkably, this habit has evolved independently
All members of the Cuculiformes have feet with at least twice in Old World cuckoos. The chicks
two toes pointing forward and two backward. of crested cuckoos (Clamator sp.) outgrow their
thick, black
The most primitive cuckoos are heavy-bodied host nest so quickly that host chicks die from bars on belly
American birds that forage on or near the starvation. Another line of cuckoos, which
ground—a lifestyle taken to the extreme by the includes the Eurasian common cuckoo, evolved
sprinting roadrunner. Inspite of their reputation, chicks with more murderous intent: they actively
not all cuckoos lay their eggs in other birds’ nests. eject host eggs and chicks. long
wings

14–16 in GREAT SPOTTED JACOBIN CUCKOO 12–13½ in

35–40 cm CUCKOO Clamator jacobinus 32–34 cm
Clamator glandarius f: Cuculidae
f: Cuculidae Clamator cuckoos are 11–13½ in
With a range from large crested Old World 28–34 cm
Europe to Africa, this cuckoos. Like others,
cuckoo lays eggs in this tropical species HIMALAYAN
magpie nests, but, like from Africa and Asia CUCKOO
other Clamator species, 13½ in eats hairy caterpillars Cuculus saturatus
does not evict the 34 cm that are avoided by f: Cuculidae
host’s young. competitors. COMMON CUCKOO This species from Asia and
Cuculus canorus Australia is typical of a
f: Cuculidae group of Old World barred
PALLID This widespread Eurasian cuckoos that resemble
FAN-TAILED CUCKOO CUCKOO species overwinters in Africa sparrowhawks—a fact
Cacomantis flabelliformis Cacomantis pallidus and S. Asia. It has the that perhaps helps them to
f: Cuculidae f: Cuculidae well-known “cooc-coo” call flush hosts from nests.
This brown-breasted Many Old World cuckoos that gives cuckoos their name.
cuckoo of Australasian have barred plumage,
woodlands is one of but in some, such as this
very few cuckoos that Australian species, the DIDERIC CUCKOO
12–13 in barring occurs only in
are found in the 30–33 cm Chrysococcyx caprius
Pacific—and the only immature birds. f: Cuculidae
one in Fiji. An African member of a
9½–11 in
9½ in 24–28 cm group of small bronze-sheened
24 cm tropical cuckoos, this species
9 in lays its eggs in the nest of the
23 cm weever bird.

GRAY-BELLIED CUCKOO BRUSH CUCKOO

Cacomantis passerinus Cacomantis variolosus KLAAS’S CUCKOO
f: Cuculidae f: Cuculidae 6½–7½ in Chrysococcyx klaas
Found from the Malay Peninsula Related to Asian brown-breasted 17–19 cm f: Cuculidae
to Australia, this bird is one of cuckoos, this plain-colored This small African cuckoo is 6½–7 in
many Indo-Pacific, brown- species breeds in southern Asia. a close relative of the dideric 16–18 cm
breasted species that are closely The montane populations migrate cuckoo, differing in being greener
related to Cuculus cuckoos. to warm lowlands in winter. and lacking white wing spots.
 C UC KOOS 24 in 13½ in
62 cm 34 cm
466
10–12½ in 15 in GUIRA CUCKOO
26–32 cm 38 cm Guira guira
BIRDS • CUCKOOS

f: Cuculidae
This shaggy-looking
bird belongs to the ani
GIANT COUA group of S. American
Coua gigas cuckoos. It occurs in
f: Cuculidae noisy flocks and nests
Couas are Madagascan ground-cuckoos communally in trees.
with blue facial skin and long eyelashes.
This bird lives in dry coastal forests.

YELLOW-BILLED BLACK-BELLIED 19–20½ in GREATER

CUCKOO CUCKOO 48–52 cm COUCAL
Coccyzus americanus Piaya melanogaster Centropus sinensis
f: Cuculidae f: Cuculidae f: Cuculidae
One of several American tree Like other American tree Like other coucals,
cuckoos with brown plumage cuckoos, this species builds this S. Asian species is
and white-spotted tails, this its own nest and cares for a strong-legged bird
species migrates between its young. It is found from with spurred feet and
N. and S. America. Colombia to Bolivia. long hind claws.

rufous
head crest

11 in 14 in streaked, reddish- 23½–32 in

28 cm 36 cm brown wings 60–80 cm

PHEASANT-COUCAL
PHEASANT-CUCKOO Centropus phasianinus
Dromococcyx phasianellus f: Cuculidae
f: Cuculidae Parental care in coucals is provided mostly
This American ground by the males. This Australasian species has
cuckoo inhabits the a black body when breeding, and builds
forest floor in tropical a cup-shaped nest in grass.
S. America, where it lays
its eggs in the nests of
smaller passerines. GROOVE-BILLED ANI
Crotophaga sulcirostris white
f: Cuculidae cheek
A heavy-billed American cuckoo stripe
occurring from California to
long tail Argentina, the communal-
nesting ani is a clumsy flier
PAVONINE but runs well.
CUCKOO
Dromococcyx pavoninus
f: Cuculidae gray-brown
A smaller relative of upperparts with
the pheasant-cuckoo, this white spots
tropical S. American bird 13 in
is a ground predator. 33 cm
It feeds on invertebrates.

long, graduated tail

22 in 15½–18 in
56 cm 39–46 cm

CORAL-BILLED
GROUND CUCKOO GREATER
Carpococcyx renauldi ROADRUNNER COMMON KOEL
f: Cuculidae Geococcyx californianus Eudynamys scolopaceus
One of three species 26–27 in f: Cuculidae f: Cuculidae
of Asian ground cuckoos 65–68 cm This fast-running predator is an Unusually for cuckoos, this tropical
related to Madagascan American ground cuckoo. The roadrunner brood parasite, found from Asia to
couas, this bird inhabits inhabits deserts and nests in cacti. and Australasia, is a fruit-eater. Males are
rainforests of S.E. Asia. occurs in a range from the US to Mexico. black and females are gray-brown.
OW L S 467
With acute senses, powerful weaponry, hearing, helped by the facial disk, which serves PHYLUM CHORDATA
and noiseless flight, owls are superbly to guide sounds to the large ear openings. A

BIRDS • OWLS
CLASS AVES

adapted as nighttime hunters. Only downward-pointing bill minimizes interference ORDER STRIGIFORMES

a few species are active by day. with sound. An owl can determine the direction FAMILIES 2

of prey with great accuracy from sounds. Most of SPECIES 243

Although not related to birds of prey, owls, which the nocturnal species have one ear slightly higher
belong to the order Strigiformes, are similarly than the other, an arrangement that helps them to D E B AT E
equipped with a hooked bill and strong talons. pick up sounds in a vertical direction, too. SCREECH OWLS
Most species have cryptic plumage for camouflage
when roosting during the day. Barn owls are silent hunters There are more than 60 species of owls
distinct in having a heart-shaped facial disk; owls Hunting owls usually swoop on their prey and in the genus Otus. In their native forests,
these small, cryptically patterned birds
with a round facial disk are more typical, although extend their feet to make the grab. The approach
are most reliably identified by their call.
they are a family of great diversity, ranging from is almost silent on large rounded wings that Although individual species’ calls differ
hawklike owlets to massive eagle owls. require minimal flapping. A soft fringe of serrated in frequency and duration of notes, Otus
flight feathers breaks up air turbulence, muffling owls fall into two well-defined groups:
sight an d h ea r i ng the sound of wing-beats. Some day-flying species scops owls, from the Old World, which
All owls have large, forward-facing eyes that are lack these fringed feathers. Most owls prey on call with slow notes; and American
efficient at letting in light and allow the birds to small mammals such as mice and voles, but some screech owls, which have faster, piercing
see well in dim conditions. Their binocular vision of the smaller species take large insects, and a trills. The distinction was enough for
is good for judging distance during attacks, but the few owls are specialized fish-eaters. An owl tears some ornithologists to advocate placing
the screech owls in their own separate
eyes are fixed in their sockets. To track moving larger carcasses apart with its hooked bill, but
genus—Megascops. Recent research
prey, an owl must turn its whole head. The bird swallows smaller prey whole. The indigestible shows that this division is underpinned
can do this through a wide range of movement by parts of a meal, such as bones and fur, are by differences
means of a long, flexible neck concealed beneath compressed in one part of the stomach and in DNA.
the plumage. Owls also have highly sensitive later regurgitated in the form of a pellet.

BARN OWL heart-shaped 10–17 in 7½–8 in rufous body

Tyto alba facial disk 26–43 cm 19–20 cm
f: Tytonidae
Found virtually worldwide
outside deserts and polar
regions, this is the most
widespread species of
owl. It is famous for its
blood-curdling screech.

10–18 in
25–45 cm

golden
upperparts

ASHY-FACED OWL EURASIAN SCOPS OWL

Tyto glaucops Otus scops
f: Tytonidae f: Strigidae
Confined to the Caribbean island of Hispaniola, This small, agile owl from 9–9½ in
this owl of dry forest is threatened by W. Eurasia is a typical tufted scops 22–24 cm
competition from the stronger barn owl. owl of the Old World. It nests in
cavities of trees or buildings.
RAINFOREST SCOPS OWL
Otus rutilus
f: Strigidae
7½–10 in 6½–10 in Restricted to Madagascar,
19–25 cm 16–25 cm this owl is common in wooded
areas. Most are gray; the rarer
reddish-brown birds are
confined to rainforests.
WESTERN
SCREECH OWL
Megascops kennicottii EASTERN SCREECH OWL
f: Strigidae Megascops asio
A common owl of western f: Strigidae
N. America, this species Widespread throughout eastern N.
favors wooded areas near America, this owl occurs in gray or rufous
rivers but will also enter (reddish brown) color varieties, with more
parkland and towns. rufous birds farther east in its range.
 OW L S 181/2–21 in 231/2–24 in 141/2–151/2 in
47–53 cm 60–62 cm 37–39 cm
468
dark brown
eyes
concentric
BIRDS • OWLS

dark rings on pale gray

facial disc plumage with
brown markings

26–28 in
65–70 cm
TAWNY OWL
BROWN WOOD OWL Strix aluco
Strix leptogrammica f: Strigidae
f: Strigidae Occurring throughout Eurasia,
This owl occurs in lowland the tawny owl frequents farmland,
tropical forests throughout India towns, and gardens. It belongs to
and S.E. Asia. Hard to see, this a group of related wood owls.
wood owl can be recognized by
its distinctive call.

17–20 in 181/2–19 in
43–50 cm 47–48 cm

BARRED OWL URAL OWL

Strix varia Strix uralensis
f: Strigidae f: Strigidae
A large wood owl native A relative of the great
to eastern N. America, gray owl, this N. Eurasian
this aggressive species is species frequents SPOTTED
GREAT spreading westward and coniferous and broadleaf OWL
GRAY OWL displacing the smaller forests, and may also Strix occidentalis
Strix nebulosa spotted owl. enter towns. f: Strigidae
f: Strigidae Native to western
A large owl of the circumpolar N. America, this wood
regions, this species occurs in 18–20 in owl occurs in mature
coniferous forest. It sometimes 45–50 cm coniferous forest, where
hunts during the day and preys it hunts flying squirrels
on large rodents and birds. and similar-sized prey.

81/2–10 in 231/2–30 in
21–25 cm 60–75 cm

10–11 in
25–28 cm

DESERT EAGLE-OWL
Bubo ascalaphus
f: Strigidae
A bird of the Sahara Desert,
RUFESCENT this species is a smaller, paler,
SCREECH OWL and longer-legged relative of
Megascops ingens the Eurasian eagle-owl.
f: Strigidae
This little-known bird from heavily barred
wet montane forests in northern underparts
S. America is larger than many other TROPICAL SCREECH OWL EURASIAN
screech owls, and has smaller ears. Megascops choliba 26–30 in EAGLE-OWL
f: Strigidae 66–75 cm Bubo bubo
The most widespread tropical f: Strigidae
American screech owl, this One of the largest species
species occurs in a range from of owl, this widespread
Costa Rica to Argentina. It exists eagle-owl occurs throughout
in brown and gray forms. Eurasia, and hunts animals
81/2–9 in 18–27 in as large as deer.
22–23 cm 46–68 cm

BLACK-CAPPED
SCREECH OWL VERREAUX’S EAGLE-OWL
Megascops atricapilla GREAT HORNED OWL Bubo lacteus
f: Strigidae Bubo virginianus f: Strigidae
Screech owls have diversified in forests f: Strigidae This is the largest African owl.
throughout the Americas. Many have The most widespread American owl, this It is widespread south of the
conspicuous ear tufts. This species eagle-owl is found from Alaska to Argentina, Sahara Desert, where it feeds
is confined to C. and S. Brazil. in habitats as diverse as forests and deserts. on small game animals.
 14 in 14–18 in ELF OWL
36 cm 36–45 cm Micrathene whitneyi
ear tuft
f: Strigidae 469
Like other insect-eaters,
white facial this tiny Mexican desert owl
disc does not need to fly silently,

BIRDS • OWLS
so it lacks the fringed,
sound-muffling wing
feathers of larger species.
5–6 in
13–15 cm

brilliant
yellow eyes
NORTHERN HAWK-OWL
9–91/2 in Surnia ulula
22–24 cm f: Strigidae
This relative of pygmy owls
is found in subarctic forests.
With its small head, long tail,
and daytime activity, it is the
most hawklike owl.
SOUTHERN STRIPED OWL broad
WHITE-FACED OWL Pseudoscops clamator ear tufts
Ptilopsis granti f: Strigidae
f: Strigidae A S. American
Like many other owl eared owl, this species
species, this bird of lives in open, marshy
sub-Saharan Africa habitats and nests in
breeds in the stick nests ground vegetation
of other birds. or low tree-holes.

18 in 201/2–28 in
46 cm SNOWY OWL 52–71 cm
Nyctea scandiaca
f: Strigidae
An Arctic species of eagle-owl,
this large, ground-nesting predator
breeds on open tundra, where it
feeds on lemmings and ptarmigan.

LEAST
5–6 in PYGMY OWL
13–15 cm Glaucidium
SPECTACLED OWL minutissimum
Pulsatrix perspicillata f: Strigidae
f: Strigidae Like other related small
This is a distinctive species owls, this woodland
from C. and S. American forests. bird of Paraguay and
Its fledglings are white with S.E. Brazil is active
a contrasting black face. during the day and night.

NORTHERN
PYGMY OWL
Glaucidium gnoma
f: Strigidae
Pygmy owls often fearlessly
tackle prey bigger than
themselves. This little
predator from western
N. America has been
seen attacking grouse.

6–61/2 in 61/2–7 in
15–17 cm 17–18 cm
6–7 in
15–18 cm
FERRUGINOUS
PYGMY OWL CUBAN PYGMY OWL 18–181/2 in
Glaucidium brasilianum Glaucidium siju 46–47 cm BUFFY FISH OWL
f: Strigidae f: Strigidae Ketupa ketupu
Typical of pygmy owls, Many owls nest in decayed f: Strigidae
this American species has tree cavities, but pygmy long claws This S.E. Asian relative of
eyespots on the back of the owls habitually choose old eagle-owls catches fishes and
head to confuse predators. It woodpecker holes. This other aquatic prey with the
flicks its tail when excited. species is restricted to Cuba. help of its long-clawed feet.
 OW L S LONG-EARED OWL SHORT-EARED OWL yellow
Asio flammeus
Asio otus eye
470 f: Strigidae f: Strigidae
Found in forests and This owl is found in open
8½–11 in heathland through country in a range that
21–28 cm much of the northern includes the Americas,
BIRDS • OWLS

hemisphere, this owl can Eurasia, and N. Africa,

flatten its long ear tufts, and is even found on
rendering them invisible. many Pacific islands.

7½–10 in 12–14½ in 13½–17 in

19–25 cm 31–37 cm 34–43 cm

TENGMALM’S OWL BURROWING OWL

Aegolius funereus Athene cunicularia
f: Strigidae f: Strigidae pale brown,
This owl of northern A long-legged owl, this partly diurnal heart-shaped face
circumpolar forests is adept at species inhabits grassland and deserts
locating small mammals under across the Americas, nesting in burrows
snow. It hunts during daylight. of prairie dogs or other animals.
yellow
eye
PEL’S FISHING OWL 25–26 in
Scotopelia peli 63–65 cm
f: Strigidae
The largest African fishing
owl, this bird frequents
riverside forests, where it heavily streaked
hunts from low perches in upperparts
slow-running waters. It
eats mainly fishes, but also
crabs and frogs.

barred
underparts
15–17 in
38–43 cm

12–14 in 15 in
30–35 cm 38 cm

BARKING OWL
Ninox connivens MOREPORK BLACK-AND-WHITE
f: Strigidae Ninox novaeseelandiae OWL
Found from the f: Strigidae Ciccaba nigrolineata
Moluccas, Indonesia, Typical of Australasian f: Strigidae
to Australia, this hawk-owls, this bird has large, This barred owl of dense
woodland hawk-owl is yellow eyes. Unusually for owls, tropical forests from Mexico
named for its doglike the male of the species to Ecuador is a close relative
barking call. is larger than the female. of the Strix wood owls.

M ESITES
WHITE-BREASTED
MESITE
Mesitornis variegatus
f: Mesitornithidae
This bird is found in three
These small, insectivorous birds of PHYLUM CHORDATA isolated areas of undisturbed
forest and scrub in Madagascar are CLASS AVES deciduous forest where
small groups forage among
flightless, or almost so. ORDER MESITORNITHIFORMES
leaf litter on the ground
FAMILIES 1 for insects, spiders,
In the family Mesitornithidae, mesites resemble SPECIES 3 and seeds.
passerines in their use of song for territorial
defense, but they are social birds. Small groups often feed, rest, and preen together. 12 in
Two species of the family are monogamous and their sexes look alike, while one is 31 cm
polygamous, with marked differences between the male and the female bird.
NIGHTJARS 471
All nightjars and their relatives are is particularly wide in the aptly named frogmouths PHYLUM CHORDATA
nocturnal. Most are insect-eaters, from Australasia, which are predators of small

BIRDS • NIGHTJARS
CLASS AVES

snatching their prey on the wing using vertebrates. Of the group, only the oilbird is ORDER CAPRIMULGIFORMES

an exceptionally wide bill gape. vegetarian: a fruit-eater that roosts by day in caves. FAMILIES 4

All these birds are masters of camouflage. SPECIES 122

Voracious predators by night, concealed by cryptic Nightjars blend perfectly with forest litter when
plumage during the day, nightjars have similarities on the ground; in trees they perch lengthwise
to owls, and some ornithologists have thought the on a branch to minimize detection. Frogmouths
two groups to be related. Recent studies, however, and potoos freeze when disturbed to mimic tree
link nightjars with swifts and hummingbirds. stumps, closing their eyes to complete the effect.
The connection is supported by features such
as the weakness of their feet and the ability of minimal nests
some species to enter a torpid state. All in this group are minimalist nesters. Nightjars
Nightjars belong to the order Caprimulgiformes. simply lay their eggs in leaf litter on the ground.
Typically, they have a big head and long wings for For oilbirds, a nest is a pile of droppings on a
A great potoo opens its enormous gape,
quick aerial maneuvers. Their short bill opens cave ledge, while potoos use nothing more than displaying a key feature of a group of birds
enormously wide to catch flying insects. The gape a depression on a branch. that can scoop up flying insects on the wing.

orange-yellow long bristles

16–19 in eye at base of bill 9–9½ in
41–48 cm 22–24 cm

OILBIRD
Steatornis caripensis
f: Steatornithidae
A fruit-eating relative of nightjars, 12½–18 in
this cave-nesting species from 32–46 cm
northern S. America is the
only nocturnal bird that TAWNY
navigates by echolocation. FROGMOUTH
Podargus strigoides
f: Podargidae COMMON NIGHTHAWK
This nocturnal Australian Chordeiles minor
bird hunts from perches. f: Caprimulgidae
Like other frogmouths, it Nighthawks lack the bill bristles of nightjars,
erect, roosts motionless during and this absence perhaps helps in their aerial
stumplike 14–16 in the day, resembling pursuit of insects. This insect-eater migrates
posture 36–41 cm a broken branch. between N. and S. America.

8 in
20 cm
gray-brown, OCELLATED POORWILL
barklike Nyctiphrynus ocellatus
plumage f: Caprimulgidae
Poorwills are small American nightjars,
so called because of their mournful,
monotonous calls. This dark-colored
species occurs in tropical forests.

gray mottled
plumage

7½–8½ in 9½–11 in
19–21 cm 24–28 cm
COMMON
POTOO
Nyctibius griseus COMMON POORWILL
f: Nyctibiidae Phalaenoptilus nuttallii
Potoos are nocturnal f: Caprimulgidae COMMON PAURAQUE
insectivores of C. and From arid parts of the US and Nyctidromus albicollis
S. America. This species Mexico, this small nightjar is f: Caprimulgidae
has a haunting call. Its one of the few birds that This C. and S. American nighthawk
plumage provides excellent enters a hibernationlike frequents scrubby habitats. It is often
camouflage against tree bark. torpor in winter. seen resting on dirt roads at night.
 NIGHTJARS
reddish brown collar
472

10–11 in
26–28 cm

EUROPEAN NIGHTJAR SPOT-TAILED NIGHTJAR

Caprimulgus europaeus Caprimulgus maculicaudus
f: Caprimulgidae f: Caprimulgidae
Like many nightjars of northern temperate Found from Mexico to Paraguay, this 8 in
regions, this species is migratory. Found species may be more closely related to 20 cm
in W. Eurasian heath and woodland, tropical American nighthawks than
it overwinters in Africa. Old World nightjars.

10–10½ in
25–27 cm

9 in 9 in
22 cm 23 cm DUSKY NIGHTJAR
Caprimulgus saturatus
f: Caprimulgidae
LARGE-TAILED NIGHTJAR PLAIN NIGHTJAR Perhaps genetically closer
Caprimulgus macrurus Caprimulgus inornatus to American poorwills
f: Caprimulgidae f: Caprimulgidae than Old World nightjars,
Among the most widespread of the Old This African bird occurs from Mauritania this species occurs in
World nightjars, this species ranges from to Saudi Arabia, and migrates south to montane forests of
Pakistan to Australia and New Guinea. Liberia, Congo, and Tanzania. Costa Rica and Panama.

LONG-TAILED NIGHTJAR LADDER-TAILED

Caprimulgus climacurus NIGHTJAR
f: Caprimulgidae 9–11 in Hydropsalis climacocerca
The long tail of some nightjars 23–28 cm f: Caprimulgidae
may be used in a courtship This Amazonian species
10–14 in display. This African species has belongs to a group of
25–35 cm a range from Senegal to Ethiopia. S. American nightjars, in
which males have a long, forked
tail with white markings. These
give the bird its name and are
MADAGASCAR NIGHTJAR possibly used to attract females.
Caprimulgus madagascariensis
f: Caprimulgidae 8½ in
This nightjar from Madagascar 21 cm
and Aldabra Island (Seychelles) has 9 in
a call that resembles the sound of 22 cm
a marble bouncing on
a hard floor.

SHORT-TAILED NIGHTHAWK
Lurocalis semitorquatus
f: Caprimulgidae
One of several tropical American LONG-TRAINED
nighthawks, this bird has a short NIGHTJAR
tail and long wings, which give Macropsalis creagra
mottled plumage it a batlike appearance on its f: Caprimulgidae
for camouflage
insect-chasing flights. Males of many nightjars
have an elaborate tail.
This species evolved
elongated tail from tropical American
feather nighthawks and is limited
to eastern S. America.
13½–30 in
34–76 cm
STANDARD-WINGED NIGHTJAR
Macrodipteryx longipennis
f: Caprimulgidae flaglike
This African species eats insects, including moths and 8½–9 in flight feathers
beetles. Breeding males have flaglike flight feathers, longer 21–23 cm
than the body, which they raise in courtship displays.
HU M MINGBIRDS AND SWIFTS 473
The dashing flight of swifts—fastest Hummingbirds have the wing control to fly PHYLUM CHORDATA
of all birds—and the audible hovering backward, some of them whirring at more than

BIRDS • HUMMINGBIRDS AND SWIFTS

CLASS AVES

of hummingbirds characterize the 70 wing-beats per second—sustained by a high ORDER APODIFORMES

master aeronauts united in this order. metabolism fueled by energy-rich nectar. Insects and FAMILIES 4

spiders provide hummingbirds with protein to feed SPECIES 405

Both swifts and hummingbirds, members of the to their chicks. Hummingbirds use cobwebs to glue
order Apodiformes, have tiny feet that are useful together their thimble-sized nests, while swifts
only for perching, but in compensation the birds use saliva. The swifts’ range is almost worldwide,
have exceptional maneuverability on the wing. reaching oceanic islands. Hummingbirds are
Swifts skim through the skies to catch flying insects. found only in the Americas.

COMMON SWIFT CHIMNEY SWIFT

Apus apus Chaetura pelagica
f: Apodidae f: Apodidae 43/4–6 in 8–9 in
This Eurasian swift Originally nesting in caves 12–15 cm 20–22 cm
nests in holes in cliffs and tree-holes, this eastern
or buildings, making it a N. American species now
common bird in towns. It nests primarily in chimneys
overwinters in Africa. in urban centers. It
overwinters in S. America.
6½ in 6–7 in
16–17 cm 15–18 cm ALPINE SWIFT
Tachymarptis melba
f: Apodidae
Distinctively white-
bellied, with a dark
undertail, the Alpine swift
occurs in southern
WHITE-THROATED SWIFT Eurasia, Africa, and
Aeronautes saxatalis Madagascar. It feeds
f: Apodidae on large insects.
Found in canyons and mountains of western
N. America and C. America, this bird gathers
in flocks in the evenings to roost. SCALE-THROATED
HERMIT
Phaethornis eurynome
f: Trochilidae
Hermits are dull-colored
hummingbirds with a
43/4 in long, curving bill for
deeply 12 cm probing Heliconia flowers. decurved bill
forked This species is from
tail eastern S. America.

5 in 41/4 in
13 cm 11 cm
WHITE-TIPPED SICKLEBILL
Eutoxeres aquila metallic violet-blue
f: Trochilidae violet “ear” blue underparts
This bird has a range that extends from Costa patch throat
Rica to Peru. It uses its decurved bill to feed
from similarly curved Heliconia flowers.

4 in 5½–6 in
10 cm 14–15 cm

GREEN-FRONTED
LANCEBILL WHITE-VENTED VIOLET SABERWING
Doryfera ludovicae VIOLET-EAR Campylopterus hemileucurus
f: Trochilidae Colibri serrirostris f: Trochilidae
This inhabitant of the forests f: Trochilidae Saberwings are named for
of the Andes takes nectar From a group of the thickened shafts of
from five species of violet-eared hummingbirds, their flight feathers. This
epiphytes (plants that grow most of which occur in C. American bird is the
on other plants), one of highland forests, this is a largest hummingbird
which is a mistletoe. species of S. American savanna. outside S. America.
 HUMMINGBIRDS AND SWIFTS 2¾ in 2¾–3½ in
7 cm
474 7–9 cm

3½ in
9 cm

HOODED BRAZILIAN RUBY RUBY-THROATED

VISORBEARER Clytolaema rubricauda HUMMINGBIRD
Augastes lumachella f: Trochilidae Archilochus colubris
f: Trochilidae Named for the male’s f: Trochilidae
brown-black Visorbearers, with their brilliant ruby-red throat, This tiny bee hummingbird
plumage green facial mask, are this hummingbird is found is the eastern US’s only
hummingbirds of arid in S.E. Brazil, but may be breeding hummingbird. It
habitats, such as highland closely related to species flies nonstop to the Gulf
savanna. This species found in the Andes. of Mexico to overwinter.
occurs in E. Brazil.
3¼ in
8 cm 4–4¼ in
10–11 cm

RUBY TOPAZ 4–7 in 4¼ in

Chrysolampis mosquitus 10–18 cm 11 cm
f: Trochilidae
This S. American, open-country
species has a ruby-red crown and
yellow throat, but in poor light the
bird can appear all black.

LONG-TAILED BUFF-TAILED BUFF-BELLIED

bright orange SYLPH CORONET HUMMINGBIRD
tail Aglaiocercus kingi Boissonneaua flavescens Amazilia yucatanensis
f: Trochilidae f: Trochilidae f: Trochilidae
Belonging to a diverse group Found in Colombia, Venezuela, Belonging to a large
of Andean hummingbirds and Ecuador, this relative of group of emerald
called coquettes, this bird Andean hummingbirds forages hummingbirds found
inhabits woodland borders near flowering trees from in C. America, this bird
3½ in and gardens. Males have a mid-level to forest canopy, is a Mexican species
9 cm long, deeply forked tail. often with other birds. of open woodland.
dark
BLUE-CHINNED ear patch
SAPPHIRE BLUE-THROATED
Chlorostilbon notatus HUMMINGBIRD
f: Trochilidae Lampornis clemenciae
One of many emerald short, f: Trochilidae
hummingbirds with metallic straight bill This large Mexican species belongs to a
green plumage, this is a group of hummingbirds called mountain
forest and farmland bird gems that originated in C. America. The
from northern S. America. northerly populations are migratory.

4¾ in 4 in
12 cm 10 cm

4¼ in 3½ in
11 cm 9 cm

COLLARED INCA
Coeligena torquata
f: Trochilidae SPECKLED HUMMINGBIRD ANNA’S HUMMINGBIRD
Incas form a group of forest Adelomyia melanogenys Calypte anna
hummingbirds native to the f: Trochilidae f: Trochilidae
Andes. The collared inca is A common hummingbird from the A bee hummingbird of western N. America,
one of the most widespread Andes, this species has duller plumage this species overwinters farther north than
of the group, found from than many of its relatives in the other hummingbirds. The bird feeds on
Colombia to Bolivia. coquette group. Both sexes are similar. nectar from a wide range of flowers.
 WHITE-EARED LUCIFER
HUMMINGBIRD HUMMINGBIRD
Basilinna leucotis 5 in 4 in 3½ in Calothorax lucifer 475
f: Trochilidae 13 cm 10 cm 9 cm f: Trochilidae
This emerald hummingbird This bee hummingbird is
frequents pine and oak montane found in a range from the

BIRDS • HUMMINGBIRDS AND SWIFTS

forests, often near streams, from southern US to Mexico. It
S. Arizona to Nicaragua. lives in semideserts, especially
those with Agave plants.

ANDEAN HILLSTAR
Oreotrochilus estella
f: Trochilidae purple
3½–4 in An Andean coquette throat
9–10 cm hummingbird, this species patch
lives higher than other species
of the group, and feeds in air
so thin that it cannot hover.

straight, black bill

FORK-TAILED BROAD-BILLED
WOODNYMPH HUMMINGBIRD
Thalurania furcata Cynanthus latirostris
f: Trochilidae f: Trochilidae
A relative of the C. American emerald Males of this Mexican scrub
hummingbirds, this S. American species hummingbird fly back and
occurs in lowland forests as far forth in a pendulum display
south as N. Argentina. to attract females. The
species belongs to the 2–2¼ in 4¾ in
“emerald” group. 5–6 cm 12 cm
iridescent
green throat

4 in 9–10 in
10 cm 23–26 cm

SWORD-BILLED BEE HUMMINGBIRD RACKET-TAILED

HUMMINGBIRD Mellisuga helenae PUFFLEG
Ensifera ensifera f: Trochilidae Ocreatus underwoodii
f: Trochilidae This tiny hummingbird f: Trochilidae
The only bird with a bill is confined to Cuba, but Belonging to the inca group,
longer than its body, this is related to migratory this bee-sized hummingbird
member of the Andean inca N. American hummingbirds. of Andean forests is attracted
group favors the trumpet- Males of the species are the to brush blossoms, such as
shaped flowers of passion vines. smallest of all birds. those of legumes.

white spot
near the eye
RUFOUS
2¾–3½ in HUMMINGBIRD 3½ in
7–9 cm Selasphorus rufus 9 cm
4 in rufous f: Trochilidae
10 cm (reddish brown) Sustaining the longest
upperparts migration of comparably
STRIPE-BREASTED small birds, this aggressively
STARTHROAT territorial bee
Heliomaster squamosus hummingbird takes a route
f: Trochilidae from Alaska to Mexico.
Male starthroats have a vividly
colored band around their
throat. Although starthroats
are related to C. American
mountain gems, this species 3½ in
is from eastern Brazil. 9 cm

4¾ in 3½ in
12 cm 9 cm

WHITE-NECKED GORGETED SUNANGEL CALLIOPE PLOVERCREST

JACOBIN Heliangelus strophianus HUMMINGBIRD Stephanoxis lalandi
Florisuga mellivora f: Trochilidae Stellula calliope f: Trochilidae
f: Trochilidae Sunangels belong to the f: Trochilidae This distinctive bird
Genetic research suggests that coquette group of Andean One of the migratory bee may be related to larger
this large, canopy-dwelling bird of hummingbirds. This species hummingbirds, this bird breeds saberwings of C. America,
tropical American lowlands belongs is found from Colombia in open forests of western but is found in montane
to a small lineage independent of to Ecuador and lives in N. America and overwinters in forests of eastern
other groups in its family. wet forest thickets. the Mexican semidesert. S. America.
476 T RO G O N S
Gaudily colored birds of tropical though reluctant to break cover. The first and PHYLUM CHORDATA
forests, the fruit-eating trogons have second toes of a trogon’s foot face backward, CLASS AVES

a wide bill, delicate plumage, and a whereas the third and fourth toes face forward. ORDER TROGONIFORMES

foot structure unique to this group. This arrangement is seen in no other birds. FAMILIES 1

These unusual feet are so weak that a trogon SPECIES 43

Crow-sized birds belonging to the can barely manage more than a shuffle dark
order Trogoniformes are found in when perched. violet-blue
crown
tropical America, Africa, and Asia. Trogons use their wide bill for tackling large
The males have brighter plumage than fruit and taking invertebrates such as caterpillars.
the females. Long-tailed and short- They also use their bill for excavating nesting
winged, trogons are capable fliers, holes in rotten wood and termite mounds.

iridescent 12–14 in 10½–12½ in

green-blue 31–36 cm 27–32 cm
plumage

10–11 in
RED-HEADED 26–28 cm
14–39 in
35–100 cm TROGON ORANGE-BREASTED
Harpactes erythrocephalus TROGON
f: Trogonidae Harpactes oreskios
An Asian trogon found from f: Trogonidae
the Himalayas to Sumatra, The color pattern of this bird, dull
this elusive bird, like many upperparts, and bright underparts, is
of its cousins, perches typical of many Asian trogons. This
motionless for long periods. species is a native of S.E. Asia.

long tail (only

in male)
CUBAN TROGON
Priotelus temnurus
f: Trogonidae
RESPLENDENT
QUETZAL One of two species of
Pharomachrus mocinno Priotelus trogons confined to
f: Trogonidae 11–12 in 10–10½ in the Caribbean, this is the
Quetzals are iridescent 28–30 cm 25–27 cm national bird of Cuba.
American trogons. The
male’s long tail in this
C. American species
is almost half its ELEGANT TROGON MASKED TROGON
overall length. Trogon elegans Trogon personatus
f: Trogonidae f: Trogonidae
The only trogon with a range Males of this species
that extends to the US, this from S. American montane jagged-tipped tail
bird occurs in montane forests have plumage
forests from southern that resembles that of the
Arizona to C. America. elegant trogon.

MOUSEBIRDS SPECKLED MOUSEBIRD

Colius striatus
f: Coliidae
The largest and one of the most
Named for their scurrying, rodentlike PHYLUM CHORDATA
widespread of mousebirds, this
species occurs from Nigeria to South
behavior, the mousebirds are a small CLASS AVES Africa in savanna and open woodland.
group of drab-colored, long-tailed birds ORDER COLIIFORMES
12–14 in
FAMILIES 1
confined to sub-Saharan Africa. SPECIES 6
30–35 cm
13–14 in
33–35 cm
Mousebirds, the sole family of the order Coliiformes,
are softly feathered in shades of brown or gray and have an erectile crest and long tail. BLUE-NAPED
MOUSEBIRD
Gregarious and agile, they resemble parakeets in behavior. These tree-dwelling birds build Urocolius macrourus
twiggy, cup-shaped nests. The chicks hatch at an advanced stage of development and f: Coliidae
Urocolius mousebirds are
quickly master flight. Ornithologists think that mousebirds are the remnant stronger fliers and less
of a more diverse group that, in prehistoric times, ranged beyond Africa. Fossil “mouselike” than Colius
mousebirds. This scrubland
mousebirds have been found in Europe. Their relationship to other birds is dweller is found from
uncertain: mousebirds could be allied with trogons, kingfishers, or woodpeckers. Senegal to Tanzania.
K I N G F I S H E R S A N D R E L AT I V E S 477
This order, Coraciiformes, has a efficiently, and their sharp-edged, long bill helps PHYLUM CHORDATA
worldwide distribution. All members grip slippery prey. Fish-eaters have a slender

BIR DS • KINGFISHERS AND REL ATIVES

CLASS AVES

perch with three toes pointing bill, while a broader, scooplike bill is a feature ORDER CORACIIFORMES

forward, as do the Passerines, of woodland kingfishers that catch their prey FAMILIES 6

directly from dry ground. Their short legs are SPECIES 177
but their two outer toes are fused ideal for perching, but kingfishers cannot walk.
at the base. Their feet are, however, also used for excavating
Kingfishers often have bright green or blue elongated nesting cavities in earth or sand banks.
colors, created not by pigments but by the
scattering of light from the microstructure of o t h e r r e l at i v e s
their feathers. They follow diverse lifestyles, Other members of this order, the American
not all reflected in the popular English name. motmots and the Old World rollers, are all
Fish-eating species locate prey by sight and land hunters. All kingfisher relatives display a
dive onto it, either from a fixed perch or after a wide variety of bill shapes. In the aerial-hunting
hover in midair. Others also hunt land animals, bee-eaters, the long bill holds stinging insects
Despite their daggerlike bill,
including lizards, rodents, and insects. A large away from the head, and is used like forceps kingfishers grasp prey, rather than stab it,
head and a strong neck helps these birds to dive to squeeze out insect venom. like almost all other fish-eating birds.

LILAC-BREASTED 11–12 in 11½–12½ in EUROPEAN ROLLER

ROLLER 28–30 cm 29–32 cm Coracias garrulus
Coracias caudatus f: Coraciidae
f: Coraciidae The most widespread of
Like other Coracias rollers, this 14–15 in the Coracias rollers, named
African bird is a predator of 36–38 cm for their somersaulting
ground animals, swooping to courtship displays, this
take lizards, rodents, and western Eurasian species
large invertebrates. overwinters in Africa.
12½–14 in
32–36 cm

BLUE-BELLIED
ROLLER black stripe
Coracias cyanogaster through eye
RUFOUS-CROWNED 14–16 in f: Coraciidae
ROLLER 36–41 cm This C. African roller, white throat
Coracias naevius like many of its relatives,
f: Coraciidae nests in tree cavities.
Somewhat dull in color RACQUET-TAILED PITTA-LIKE
It is common throughout ROLLER GROUND ROLLER
compared to other species, its extensive range.
this large roller inhabits Coracias spatulatus Atelornis pittoides
arid areas across sub- f: Coraciidae f: Brachypteraciidae
Saharan Africa. This distinctive As the name suggests,
fork-tailed roller ground rollers are
from E.Africa has mostly terrestrial.
outer tail feathers This rainforest species
with projecting is the most brightly
“flag-tipped” vanes. colored of the family.

DOLLARBIRD
Eurystomus orientalis
f: Coraciidae 10 in
Named for its pale, 26 cm
dollar-shaped underwing long,brown tail
markings, this roller occurs with dark barring
in woodlands from the
Himalayas to Australia. 18½–20½ in
47–52 cm

10½–12 in
27–30 cm yellow bill

BROAD-BILLED
ROLLER LONG-TAILED
Eurystomus glaucurus GROUND ROLLER
f: Coraciidae Uratelornis chimaera
Typical of its genus, this f: Brachypteraciidae
African species is longer Like other ground rollers,
winged and more agile in this species from the dry forests of
10½–12 in pursuit of flying prey than S.W. Madagascar nests in holes
27–30 cm Coracias rollers. excavated in the ground.
 KINGFISHERS
478 A N D R E L AT I V E S
8 in
20 cm
BIR DS • KINGFISHERS AND REL ATIVES

41/2 in
11 cm GRAY-HEADED
WHITE-THROATED KINGFISHER
KINGFISHER Halcyon leucocephala
Halcyon smyrnensis f: Alcedinidae
f: Alcedinidae Tree kingfishers are a group
Like many other tree kingfishers, of mainly insect-eating forest
this southern Asian species is birds. The gray-headed
a noisy bird. It has whinnying kingfisher is an African species
and laughterlike calls. that may also eat fishes.

JAMAICAN TODY
Todus todus white chest
f: Todidae and throat
Todies are tiny, green
kingfisherlike birds from the
Caribbean, with a flattened bill
for catching insects in flight. dark eye patch
This species is found only in contrasts with the
the woodlands of Jamaica. white head
heavy bill
111/2 in 43/4–5 in
29 cm 12–13 cm

bright blue back 16–181/2 in

41–47 cm
and wings

AFRICAN PYGMY KINGFISHER

Ispidina picta
f: Alcedinidae
This African bird belongs to a group of river
kingfishers, but mainly feeds on insects so occurs
away from water in woodland and savanna.

7–81/2 in
18–21 cm

7–8 in
18–20 cm YELLOW-BILLED
LAUGHING KINGFISHER
GREEN KINGFISHER KOOKABURRA Syma torotoro
Chloroceryle americana Dacelo novaeguineae f: Alcedinidae
f: Alcedinidae f: Alcedinidae This tree kingfisher occurs
A tropical American bird, this Widespread in Australian in rainforests of New
water kingfisher plunge-dives woodland, this large tree Guinea and N. Australia.
for fishes and aquatic insects kingfisher preys on reptiles It feeds mainly on insects,
and often perches on rocks as well as invertebrates, and but also eats worms
in streambeds. has a loud, chuckling call. and small lizards.

61/2–71/2 in 43/4–5 in 61/2 in

17–19 cm 12–13 cm 16–17 cm

azure-blue
upperparts

orange
underparts 11–111/2 in
28–29 cm

AZURE KINGFISHER LITTLE KINGFISHER COMMON KINGFISHER PIED

Ceyx azureus Ceyx pusillus Alcedo atthis KINGFISHER
f: Alcedinidae f: Alcedinidae f: Alcedinidae Ceryle rudis
Like other river kingfishers, This tiny Australasian Found in Eurasia and f: Alcedinidae
the azure kingfisher nests in species of mangroves is a N. Africa, this bird is typical From Africa and S. Asia, this water
holes along riverbanks. It is river kingfisher. It eats tiny of a group of diminutive, kingfisher of rivers and lakes often gathers
found in creeks and mangroves fishes, insect larvae, small short-tailed river kingfishers in small groups. It can hover in one
in Australia and New Guinea. crustaceans, and shrimps. from the Old World. spot for long periods of time.
 479
11–14 in
28–35 cm

BIRDS • CUCKOO-ROLLER
12–14 in 141/2 in 141/2–16 in
30–35 cm 37 cm 37–41 cm
single blue
chest band

BELTED KINGFISHER BUFF-BREASTED BROWN-WINGED

Megaceryle alcyon PARADISE KINGFISHER KINGFISHER
f: Alcedinidae Tanysiptera sylvia Pelargopsis amauroptera STORK-BILLED KINGFISHER
A water kingfisher from f: Alcedinidae f: Alcedinidae Pelargopsis capensis
N. America, the belted Many tree kingfishers, such as this A tree kingfisher of mangrove f: Alcedinidae
kingfisher is a fishing specialist species from New Guinea and forests, this species occupies Occurring in southern Asia, this bird is found
that often hovers over water N. Australia, nest in holes in a range from India to the near water in forests, especially outside the
before diving for prey. treetop termite mounds. Malay Peninsula. range of the related brown-winged kingfisher.

turquoise-blue
white central 18 in crown
10–11 in tail feathers 46 cm
25–28 cm
BLUE-CROWNED
MOTMOT
COLLARED RUFOUS MOTMOT Momotus momota
KINGFISHER Baryphthengus martii f: Momotidae
Todiramphus chloris f: Momotidae Like the related bee-eaters
f: Alcedinidae Like other motmots, and many kingfishers, this
Favoring mangroves, this stout-billed bird from typical brightly colored
this tree kingfisher is widespread C. and northern S. America motmot from tropical
in coastal areas across southern sallies for large insects America nests in
Asia to the Pacific. and small vertebrates.
16 in
tunnels in riverbanks. TURQUOISE-
41 cm BROWED MOTMOT
Eumomota superciliosa
f: Momotidae
13 in As with other motmots,
33 cm the racket-tipped tail
black of this C. American
eye-stripe
long, slightly motmot is formed by
curved bill weakened feather barbs
9 in 81/2–91/2 in that break away.
23 cm 22–24 cm

10–111/2 in
25–29 cm
81/2–10 in
22–25 cm
EUROPEAN BEE-EATER RAINBOW WHITE-FRONTED
Merops apiaster BEE-EATER BEE-EATER GREEN BEE-EATER
f: Meropidae Merops ornatus Merops bullockoides Merops orientalis
Like other bee-eaters, this f: Meropidae f: Meropidae f: Meropidae
bird from southwestern The only bee-eater in Australia, This African bee-eater breeds Found in dry, open habitats
Eurasia and Africa catches southernmost populations of in large colonies, which have in Africa and S. Asia, the
insects in flight. Before eating this species migrate northward a complex social system with green bee-eater nests in
bees, it removes the sting to overwinter in northern non-breeding birds assisting looser colonies than other
from its prey. Australia and Indonesia. in raising the young. species of bee-eater.

C U C KO O - RO L L E R Small eye in
bulky head

Comprising a single rare species from PHYLUM CHORDATA

Male has
iridescent
the Madagascar forest, the order CLASS AVES upperparts
Leptosomiformes is placed close to the ORDER LEPTOSOMIFORMES
FAMILIES 1
quetzals of South America, though its SPECIES 1
Short,
affinity to other orders is unclear. thick bill
The cuckoo-roller, which survives in fragmented
remnant forests, seems less endangered than many
other Madagascan species. It feeds like a flycatcher CUCKOO-ROLLER 151/2–191/2 in
Leptosomus discolor 40–50 cm
on large insects, but also eats small reptiles. Similar f: Leptosomatidae
to cuckoos, it is able to digest hairy caterpillars, Superficially resembling a cuckoo,
this Madagascan predator of small
which most species avoid. Again similar to cuckoos, animals is not closely related to true
its two toes face forward and two backward. rollers. It nests in tree cavities.
 SHOVEL-BILLED ASSASSIN spiky crown feathers
480 A bold, dry-land, woodland-edge kingfisher, form short crest
the laughing kookaburra sits on an exposed
perch looking for prey. It uses its shovel-
shaped bill and muscular neck to grab small
snakes and rodents, hitting larger prey
BIR DS • KINGFISHERS AND REL ATIVES

against the ground to subdue them.

broad, dark
cheek-patch
below large eye

lower mandible
used to scoop up
small prey

white underside helps hide

the bird against the sky
from prey below
L AU G H I N G KO O K A B U R R A 481
Dacelo novaeguineae
Laughing kookaburras are famously vocal. A first

BIR DS • KINGFISHERS AND REL ATIVES

loud, chuckling laugh spreads into a loud chorus from
several birds, the staccato notes echoing through the
Australian bush. “Laughing jackass” was a popular
nickname for this large-headed, short-legged bird; another,
“bushman’s clock,” reflects the regularity of calls at dawn and
dusk. Its nest is an unlined hollow in a tree stump or in a hole excavated with
its bill in a termite mound. Like other kingfishers, its three to five eggs are pure
white. If food is short, the final egg is small and the resultant weak chick is likely
to be killed by its stronger siblings. Adult laughing kookaburras form lifelong pair- size 16–181⁄2in (41–47 cm)
bonds, but are helped at the nest by up to five immature birds from previous years, habitat Open, bushy woodland and eucalypt forest
distribution Eastern Australia, introduced in other parts of
and all may incubate the eggs and then brood and feed the chicks. This species is Australia and New Zealand
widespread and relatively common, but recent wildfires in Australian woodland diet Lizards, snakes, small mammals, insects, worms, snails, and
have seriously affected many populations. other invertebrates

EYE
A sensitive area of the retina in
each eye allows exceptionally sharp
vision. The laughing kookaburra
moves its head to align a second,
sensitive area in each eye to give
binocular vision when searching
for prey below.
upper mandible
tapers to a
slender point TONGUE
The broad, pointed tongue sits neatly
inside the deep lower mandible. Sticky
with mucus, it has a small “hook”
at the rear, and helps to manipulate
and swallow wriggling or muscular
prey items.

FOOT
As with other kingfishers, the long outer toe
and middle toe are fused together. The very NOSTRIL
short leg gives a secure, upright stance on An open nostril high at
a perch, but means the laughing kookaburra the base of the bill
can only shuffle when on the ground. minimizes the risk of
clogging by sand or mud.
The long, bristly feathers
behind it help protect
the eye.

TAIL
The laughing
kookaburra’s tail is long
for a kingfisher. In flight,
the tail is spread wide,
revealing all 12 feathers.

WING
Short and broad, the wing is curved into
outer tail feathers have an airfoil: the thick, muscular inner
more white bands than “arm” tapers to flat, bladelike flight
inner feathers feathers behind and at the tip.
482 HORNBILLS, HOOPOES, AND
WO O D H O O P O E S
BIRDS • HORNBILLS, HOOPOES, AND WOOD HOOPOES

Hornbills (including some giants of the curved bill, while wood hoopoes hop on trees and PHYLUM CHORDATA
bird world), hoopoes, wood hoopoes, probe inside loose bark. Hornbills have a larger, CLASS AVES

and the similar scimitarbills form an longer bill, some with a large, horny adornment on ORDER BUCEROTIFORMES

Old World order, the Bucerotiformes, top. They range from medium-sized to very large FAMILIES 4
SPECIES 74
some boldly colored and all strikingly forest species. Ground hornbills are extremely
large, heavy birds that feed exclusively on the
patterned and distinctive in form.
ground. Some hornbills nest in tree holes and males
Hoopoes walk on the ground and probe for seal the females inside with mud, feeding them
invertebrates and small reptiles with their slender, through a narrow slit until the young hatch.

reddish bill

16½–18 in RED-BILLED TRUMPETER 23–26 in 30–32 in

42–45 cm HORNBILL HORNBILL 58–65 cm 75–80 cm
Tockus erythrorhynchus Bycanistes bucinator
f: Bucerotidae f: Bucerotidae
Tockus hornbills are small This hornbill inhabits
gray, white, and African predators, most the forests of eastern to
black plumage with a red or yellow bill. southern Africa. It is
This species inhabits similar to the related
savanna and open woodland silvery-cheeked hornbill, SILVERY-CHEEKED
from Senegal to Namibia. but has red facial skin. HORNBILL
Bycanistes brevis
f: Bucerotidae
This fruit-eating pied hornbill
is a forest species from
E. Africa. As with other
hornbills, the male has a larger
bill than the female, with a
white more prominent casque.
underparts ORIENTAL PIED
HORNBILL
Anthracoceros albirostris
f: Bucerotidae
The most widespread of the
Asian pied hornbills, this bird
occurs from the Himalayas to black-tipped
Bali (Indonesia), in cultivated crest, raised
land as well as forest. when landing

28 in 3¼ in
70 cm 1m

28 in 10–12½ in
70 cm 25–32 cm

black and
white bars 12–14 in
on wings 30–36 cm

NORTHERN GROUND
MALABAR PIED HORNBILL GREEN
HORNBILL Bucorvus abyssinicus WOOD HOOPOE
Anthracoceros coronatus f: Bucorvidae Phoeniculus purpureus
f: Bucerotidae One of two predatory HOOPOE f: Phoeniculidae
Like most other Asian ground hornbills of Upupa epops This African wood hoopoe
hornbills, this pied African grasslands, this f: Upupidae climbs trees like a woodpecker,
species from India and Sri species from Senegal to Kenya This species nests in tree-holes in Africa and but has a hoopoelike bill
Lanka is omnivorous, and tolerates drier conditions Eurasia. It has a flickering, butterflylike flight and to probe rotting wood
eats a great deal of fruit. than its southern counterpart. a strong bill to dig for ground-living invertebrates. for invertebrates.
WO O D PE C K E R S A N D TOU C A N S 483
Mostly tree-dwelling cavity nesters, a long barbed tongue. Other birds in the order PHYLUM CHORDATA
birds of this order, the Piciformes, have are largely confined to the tropics. Honeyguides

BIRDS • WOODPECKERS AND TOUCANS

CLASS AVES

a common foot structure. More than feed on the wax from raided bee nests. Jacamars ORDER PICIFORMES

half of the species are woodpeckers. hunt large insects, while puffbirds are flycatchers. FAMILIES 9
SPECIES 445
Barbets have a serrated bill for eating fruits; the
The woodpeckers are almost worldwide in tropical American barbets are cousins of the
distribution. These birds cling to trunks with toucans. Honeyguides are brood parasites,
zygodactyl feet typical of the order (two toes laying eggs in the nests of other birds, including
pointing forward, two backward) and, propped woodpeckers. Others in this group nest in
by their stiffened tail, drum through the wood tree-holes, or may dig burrows in the
with their strong bill. They extract prey with ground or in termite mounds.

22–23½ in 19 in
55–60 cm 48 cm

CHANNEL-BILLED
21–23½ in 17 in CUVIER’S TOUCAN TOUCAN
53–60 cm 43 cm Ramphastos tucanus Ramphastos vitellinus
cuvieri f: Ramphastidae
f: Ramphastidae Ramphastos toucans are
This Amazonian bird is often black with a white or
regarded as a subspecies of yellow chest. In this
WHITE-THROATED RED-BREASTED the white-throated toucan, pale orange variable S. American
TOUCAN TOUCAN from which it differs by around eye species, the chest color
Ramphastos tucanus Ramphastos dicolorus having a darker bill. depends on race.
f: Ramphastidae f: Ramphastidae
Like other toucans, this Among the smallest
species from northern of the Ramphastos toucans, 22–26 in
S. America breeds in tree this species from eastern 55–65 cm
cavities, often making use of S. America is the only one
abandoned woodpecker nests. with extensive red underparts.

12–14 in 14 in
30–35 cm 35 cm
TOCO TOUCAN
Ramphastos toco
f: Ramphastidae
EMERALD This bird from northern
TOUCANET S. America is the largest of the
Aulacorhynchus prasinus toucans. Unlike other forest
f: Ramphastidae species, it also inhabits more
The most widespread of open woodland habitats.
a group of small green
toucans, this species massive,
occurs from Mexico black-tipped
to Bolivia. The species orange bill
consists of several races. SPOT-BILLED
TOUCANET
Selenidera maculirostris blue eye-skin
f: Ramphastidae
This S. Brazilian species is
one of several dichromatic
toucanets, which are the only
toucans with different-
colored sexes. The females
SAFFRON are marked with brown.
TOUCANET
Pteroglossus bailloni
f: Ramphastidae 16 in 14½ in
41 cm 37 cm
Found in S.E. Brazil, this
toucan has a yellow-olive ornate black
plumage. It is related bill with
to the more brightly COLLARED
ARACARI yellow bands
colored aracaris.
Pteroglossus torquatus
f: Ramphastidae CHESTNUT-EARED ARACARI
With the most northerly Pteroglossus castanotis
14–16 in range of the aracaris, f: Ramphastidae
35–40 cm this species occurs in humid Aracaris are long-tailed, gregarious toucans. Most
forests from S. Mexico to have a red rump and prominent bands on the belly.
northern S. America. This species is from northwestern S. America.
 WOODPEC K ERS AN D TOUCANS BLACK-BILLED BARBET
Lybius guifsobalito
484 f: Ramphastidae
Many African barbets occur in more
open country than the forest-dwelling
species of the Americas and Asia. This
BIRDS • WOODPECKERS AND TOUCANS

species occurs in E. Africa.

71/2 in 61/2 in 9 in
19 cm 17 cm 23 cm

BLACK-SPOTTED RED-HEADED BARBET

BARBET Eubucco bourcierii
Capito niger f: Ramphastidae
f: Ramphastidae This species is found from
This barbet from northern Costa Rica to Peru. It is unusual
S. America is rarely seen, but among barbets in being
its froglike call is often heard. largely silent. 10 in
red forehead 26 cm BEARDED BARBET
Lybius dubius
BLUE-THROATED BARBET f: Ramphastidae
Psilopogon asiaticus Barbets have sensory bristles at the base
f: Ramphastidae green of their bill, which are much enlarged in
Asian barbets often forage upperparts this species from W. and C. Africa.
with other fruit-eaters in the forest
canopy. This species occurs from
the Himalayas to Thailand.
61/2 in 8 in
9 in 17 cm 20 cm
23 cm

large, yellowish 11 in
bill 28 cm

COPPERSMITH CRIMSON-FRONTED
BARBET BARBET
Psilopogon haemacephalus Psilopogon rubricapillus
f: Ramphastidae f: Ramphastidae
121/2–13 in This is a widespread barbet A small Asian barbet, this
32–33 cm found in forest edges and scrub. species is restricted to Sri
BROWN-HEADED Its incessant “tonk-tonk” Lanka and southwestern
BARBET
Psilopogon zeylanicus hammering call is a common India. It is a common species
f: Ramphastidae sound in S. Asia. that is seen in towns.
A typical Asian barbet,
this species from the
Himalayas, India, and Sri
Lanka is a fruit-eater and RED-FRONTED
especially favors figs. TINKERBIRD
Pogoniulus pusillus
f: Ramphastidae
GREAT BARBET Restricted to riverside forests of
Psilopogon virens the E. African coast, this species
f: Ramphastidae feeds on insects and fruits, tail narrows
The largest Asian barbet, including mistletoe berries. to point
this species is a noisy
bird of highland forests 4–41/4 in
10–11 cm
from the E. Himalayas
to Thailand.
red vent 4–41/4 in 41/4 in
10–11 cm 11 cm

YELLOW-RUMPED
TINKERBIRD YELLOW-FRONTED
Pogoniulus bilineatus TINKERBIRD
f: Ramphastidae Pogoniulus chrysoconus
Tinkerbirds are small, f: Ramphastidae
black and white African barbets A more widespread relative of the
with a repetitive call that continues red-fronted tinkerbird, this species
through the day. This species is occurs in dry, open woodland and
widespread south of the Sahara. savanna in sub-Saharan Africa.
 SPOT-FLANKED white-spotted
BARBET upperparts
9 in
22 cm Tricholaema lacrymosa 485
f: Ramphastidae
A barbet of damp 6–61/2 in
woodlands from 15–16 cm
C. to E. Africa, the
RED-FRONTED BARBET spot-flanked barbet mainly
Tricholaema diademata feeds on figs and berries.
beardlike bristles f: Ramphastidae
An E. African barbet related to
the spot-flanked barbet, this species D’ARNAUD’S
occurs in drier habitats. Like other BARBET
barbets, it nests in tree-holes. Trachyphonus darnaudii
9 in
f: Ramphastidae
TOUCAN-BARBET 22 cm Trachyphonus barbets
Semnornis ramphastinus from Africa live in
f: Ramphastidae open country and spend
much time on the ground.
A bird of the rainforests of D’Arnaud’s barbet occurs
Colombia and Ecuador, this species throughout E. Africa.
is intermediate between barbets and striking head
toucans. It is exclusively a fruit-eater. pattern

8 in 9 in
20 cm 23 cm

RED-AND-YELLOW
BARBET
Trachyphonus erythrocephalus
f: Ramphastidae
A typical African ground
barbet, this bird eats insects,
fruits, seeds, and even small
lizards. It often burrows into
termite mounds to nest.
FIRE-TUFTED BARBET
Psilopogon pyrolophus 61/2 in
f: Ramphastidae 17 cm
The only Asian barbet with a
graduated tail, this S.E. Asian
species also has tufts of 43/4–5 in
12–13 cm
facial bristles and a
cicadalike call.
11 in
28 cm

GREEN-BACKED
HONEYBIRD NORTHERN WRYNECK
Prodotiscus zambesiae Jynx torquilla
f: Indicatoridae f: Picidae
Honeybirds eat insects, fruits, This Eurasian woodland
and even beeswax. This African ant-eater has a weaker bill
species lays eggs in the nests of the than true woodpeckers and
white-eye, a small woodland bird, is named for its ability to
and its young kill host nestlings. twist its neck.

BAR-BREASTED 4 in
PICULET 10 cm GOLDEN-SPANGLED
Picumnus aurifrons PICULET
f: Picidae Picumnus exilis
Piculets are tiny nuthatchlike f: Picidae
members of the woodpecker This S. American piculet,
family. They use their short like other piculets, lacks stiff,
bill for taking insects from supporting tail feathers of bigger
decayed wood. This species is woodpeckers, so it spends less
from central S. America. time on vertical trunks.
4 in
10 cm
4 in 4 in 4 in
10 cm 10 cm 10 cm

OCHER-COLLARED
OCHRACEOUS PICULET PICULET SPOTTED PICULET
Picumnus limae Picumnus temminckii Picumnus pygmaeus
f: Picidae f: Picidae f: Picidae
Confined to E. Brazil, the Confined to the forests of The spotted piculet
ochraceous piculet—like other E. Paraguay, S.E. Brazil, and occurs only in the
piculets—reuses woodpecker N.E. Argentina, this piculet tropical forests of
nesting holes, as its bill is too is well camouflaged by its N.E. Brazil, where it
small to excavate its own nest. pattern and coloration. is relatively common.
 WOODPEC K ERS 7 in 7–9 in 11 in
486 AND TOUCANS 18 cm 18–22 cm 28 cm

COMMON
FLAME-BACKED
WOODPECKER
Dinopium javanense
BIRDS • WOODPECKERS AND TOUCANS

12 in
30 cm
f: Picidae
A tropical woodpecker,
GROUND this species occurs in
WOODPECKER YELLOW-BELLIED a variety of woodland
Geocolaptes olivaceus SAPSUCKER types—including
f: Picidae NUBIAN WOODPECKER Sphyrapicus varius mangrove—and ranges
Unusual in living on the Campethera nubica f: Picidae from India eastward
ground, this South African f: Picidae Sapsuckers make holes in to Borneo and Java.
ant-eating woodpecker This relative of the European trees to drink sap. This
inhabits barren, rocky areas green woodpecker occurs in dry fork-tailed species breeds
and tunnels into earth banks parts of northeastern Africa, in N. America and migrates 16–191/2 in
to make a nesting chamber. where it is often seen in pairs. to the Caribbean. 40–49 cm

PILEATED WOODPECKER
red cap Dryocopus pileatus
f: Picidae
HEART-SPOTTED 6–61/2 in This is the largest N. American
WOODPECKER 15–17 cm woodpecker. Unlike the Eurasian black
Hemicircus canente woodpecker, American members of
f: Picidae the Dryocopus genus are crested.
One of two related small, 18–221/2 in
45–57 cm
crested woodpeckers
from S.E. Asia, this
species is named for the
black, heart-shaped male has
marks on its back. red face streak

9 in
23 cm GOLDEN-GREEN BLACK WOODPECKER
WOODPECKER Dryocopus martius
Piculus chrysochloros f: Picidae
f: Picidae This large woodpecker
Typical of a group of from woodlands across
green-backed woodpeckers N. Eurasia belongs
from tropical America, to a small group of
this bird often follows predominantly black species
mixed-species flocks, that have a red crown.
gleaning bark surfaces.

white patch
12–13 in on wing
31–33 cm

91/2 in
24 cm GREEN
RED-BELLIED WOODPECKER
71/2–9 in WOODPECKER Picus viridis
19–23 cm f: Picidae
Melanerpes carolinus
f: Picidae This European member
white Like other Melanerpes of a group of Old
underparts woodpeckers, this common World green-backed
N. American species stores food woodpeckers is also
in crevices. Despite its name, called the yaffle after its
the belly only has a tinge of red. distinct yelping call.

12 in
31 cm
71/2 in
19 cm

RED-HEADED YELLOW-FRONTED ROBUST

WOODPECKER WOODPECKER WOODPECKER
Melanerpes erythrocephalus Melanerpes flavifrons Campephilus robustus
f: Picidae f: Picidae f: Picidae
A distinctive woodpecker Most Melanerpes woodpeckers The Campephilus woodpeckers
from N. America, this is have partly barred plumage. are black and white birds
an aggressive species that They may also have striking with a red head. The robust
destroys nests and eggs of colors, as does this woodpecker is restricted
other birds in its territory. S. American species. to eastern S. America.
 red patch (in male) HAIRY WOODPECKER red nape 8–9 in
Picoides villosus (in male) 20–22 cm
f: Picidae 487
A N. American relative of the 9 in
three-toed woodpecker, this 22–23 cm
11–12 in species becomes more abundant

BIRDS • SERIEMAS
28–31 cm according to the availability of
its prey—larvae of bark beetle.

GREAT SPOTTED
NORTHERN FLICKER WOODPECKER
Colaptes auratus Dendrocopos major
f: Picidae f: Picidae
7–10 in
American flickers are so 18–26 cm A widespread member
called because of the way of a Eurasian group of
their colored underwings pied woodpeckers, this
flash in flight. This species bird is common in
has red- and yellow-shafted forests and gardens from MIDDLE-SPOTTED
dark bars forms that hybridize. Europe to S.E. Asia. WOODPECKER
on wings Dendrocoptes medius
red patch f: Picidae
beneath tail A pied woodpecker
11 in 9 in
28 cm 23 cm confined to Europe and
S.W. Asia, this species
red underparts drums less than the related
iridescent green great spotted woodpecker.
upperparts
RUFOUS-TAILED
JACAMAR
Galbula ruficauda
f: Galbulidae
7 in This C. and S. American
18 cm species is typical of its 8 in
genus, characterized by 20 cm
THREE-TOED iridescent green coloration
JACAMAR above and rufous below.
CHESTNUT
Jacamaralcyon tridactyla JACAMAR
f: Galbulidae Galbalcyrhynchus
The most dull-colored jacamar, purusianus
this dry forest species nests in f: Galbulidae
earth banks and has two toes Jacamars specialize in
in front and one behind. GREAT JACAMAR 8–81/2 in catching large insects,
Jacamerops aureus 20–22 cm such as butterflies, from
f: Galbulidae the air. This is one of
This is the largest species two chestnut-colored
of jacamar. It occurs from WHITE-EARED
PUFFBIRD species from S. America.
Costa Rica to Bolivia and Nystalus chacuru
11 in feeds mainly on insects, f: Bucconidae
28 cm supplemented by small lizards. Like other puffbirds, this
species from central S.
America is a big-headed, 6 in
15 cm
“puff-bodied” bird with a
heavy bill for catching
BLACK-FRONTED small animals.
NUNBIRD RUSTY-BREASTED SWALLOW-WINGED
Monasa nigrifrons NUNLET PUFFBIRD
f: Bucconidae Nonnula rubecula Chelidoptera tenebrosa
Nunbirds are f: Bucconidae f: Bucconidae
black-bodied relatives Nunlets are small, Described as martinlike
of puffbirds. This noisy drab-colored members of when perched and batlike
S. American species often the puffbird family. This in flight, this bird of
forages beneath monkey bird is a typical northern S. America
troops, feeding on disturbed 51/2 in S. American species of sallies for flying insects
small animal prey. 14 cm vine-bordered forest. along rivers.

SERIEM AS loose plumage on

neck and breast
barred wing feathers
rarely revealed

Seriemas, order Cariamiformes, are PHYLUM CHORDATA

RED-LEGGED SERIEMA
Cariama cristata
large, long-legged, noisy ground birds CLASS AVES f: Cariamidae
Related to the extinct, predatory,
of dry savanna woodland in South ORDER CARIAMIFORMES
giant terror birds, seriemas are natives
FAMILIES 1
America, with unclear relationships SPECIES 2
of the grasslands of S. America.
This species has a sicklelike claw
to other bird orders. to dismember small prey.

Once linked with cranes, seriemas have since been placed closer to falcons. The
two species are among the largest ground-living birds of South America and rarely 30–35 in
75–90 cm
fly, except to reach the canopy of trees to nest and roost overnight. Their loud calls
make them more often heard than seen.
488 PA S S E R I N E S
This huge order makes up about Passerines combine a high metabolic rate with PHYLUM CHORDATA
60 percent of all bird species. They a comparatively large brain size, giving some
BIR DS • PASSER INES

CLASS AVES

are known collectively as perching the resilience to tolerate cold and a few the ORDER PASSERIFORMES

birds, for their specialized feet. intelligence to master the use of simple tools. FAMILIES 141

They produce helpless, naked young, which are SPECIES 6,456

Like many birds, passerines have four-toed feet reared in nests ranging from simple cups to
with three toes pointing forward and one back. elaborate mud-chambers or hanging purses of D E B AT E
As a passerine lands, a muscle automatically woven grass. A few passerines are brood parasites, OUT OF AUSTRALIA
tightens tendons running through the legs so laying their eggs in the nests of other species.
that the toes lock around a perch, securing Early DNA analysis identified two
the bird even in sleep. s ongbi r d s songbird groups: Corvida and Passerida.
The mostly Old World Corvida includes
These birds, of the order Passeriformes, are found Passerines are split into two groups, largely on the
carnivores and fruit-eaters, such as
in almost all land habitats worldwide, from dense basis of the structure of their vocal box. The first crows and birds of paradise. The global
rainforest to arid desert and even freezing Arctic group, the Suboscines, forms about a fifth of all Passerida—with more than a quarter
tundra. They range in size from the tiny American passerines. It occurs in the Old World tropics of all bird species—includes tits,
flycatchers, no bigger than hummingbirds, to the but has greatest diversity in America; it includes sunbirds, and finches. Later techniques
powerfully built Eurasian ravens. broadbills, pittas, antbirds, and tyrant flycatchers. revealed several primitive Australian
The second group, the Oscines, includes all groups, such as the lyrebirds, which do
di v ersity remaining passerine families, popularly known not belong to either of these divisions.
The order includes birds adapted to a variety of as songbirds. Birds often communicate by voice, Together with fossil evidence, this
suggests that songbirds diversified in
feeding habits. The insect-eaters have a bill like a but the structure of their vocal box enables many
Australasia before giving rise to Corvida
needle for probing foliage or with a wide gape for passerines to deliver complex songs that are and Passerida.
catching prey in flight. Other passerines have a important in courtship and defending territory.
stubby bill for cracking open seeds, while some Songs are often so distinctive that
have a long curved bill for extracting nectar. they can be used to identify species.

BROA DBI L L S ASITIES thin,

decurved bill
The families Calyptomenidae and The brush-tipped tongue
Eurylaimidae consist of forest- of these Madagascan birds, 3½ in
dwelling birds from tropical the Philepittidae, suggests 9 cm
Africa and Asia. They mostly that they may have evolved
use their wide bill to catch from nectar-feeding
insects in trees, but members of 6½–7 in ancestors: one genus is COMMON
17–18 cm SUNBIRD-ASITY
one group of Asian green broadbills now fruit-eating; the Neodrepanis coruscans
are fruit-eaters. GREEN BROADBILL other resembles the One of two long-billed birds
blue bill with Calyptomena viridis unrelated sunbirds from eastern Madagascar, this
yellow at base One of the three S.E. Asian species evolved the nectar-
green broadbills, this bird is a that feed on nectar. feeding habit of true sunbirds,
specialist fruit-eater and builds but is unrelated to them.
globular hanging nests.
10 in
25 cm
PI T TA S
Birds of this Old World family forage for insects
on the forest floor in tropical regions. The
Pittidae have a rounded body and a short bill,
and many species are brilliantly plumaged.
6 in Both adults incubate the eggs.
15 cm

8 in 7½ in
20 cm 19 cm

BLACK-AND-
BLACK-AND-RED YELLOW BROADBILL BLUE-WINGED PITTA INDIAN PITTA
BROADBILL Eurylaimus ochromalus Pitta moluccensis Pitta brachyura
Cymbirhynchus macrorhynchos An Asian insect-eating Found from S. China to Borneo Like its relatives, this
This distinctively colored S.E. Asian broadbill, this species forages and Sumatra, the blue-winged pitta from the S.
broadbill frequents forests near water. in the middle and upper levels pitta inhabits dense forest when Himalayas, India, and Sri
It builds a pouchlike nest suspended of rainforests from Myanmar breeding, but overwinters Lanka builds domed nests
from the tips of branches. to Borneo and Sumatra. in coastal scrub. on or near the ground.
MANAKINS COTINGAS 5 in
13 cm
These birds, the Pipridae, are related to red cap A N D R E L AT I V E S 489
cotingas. The brightly colored males gather A diverse group of tropical American
in groups called leks to display to females. passerines, the Cotingidae consists of

BIR DS • PASSER INES

For many species, courtship involves fruit-eating or insect-eating forest birds.
elaborate dancing movements. Females Males are brightly colored and are highly
build the nest and rear offspring alone. vocal when courting females; some species
display in trees and others on the ground.

6 in bright
15 cm yellow throat

10½–11 in
27–28 cm
BARE-THROATED
BELLBIRD FIERY-THROATED
Procnias nudicollis FRUIT-EATER
Bellbirds are named Pipreola chlorolepidota
after their metallic calls. Fruit-eaters are stocky green
5½–6 in Like other bellbirds, birds, males of which are
14–15 cm males of this eastern usually black headed and
S. American species red or yellow throated. This
have white plumage. is a small Andean species.

bare red skin

around eyes
ARARIPE MANAKIN
Antilophia bokermanni
Described as recently BLUE MANAKIN
as 1998, this critically Chiroxiphia caudata
endangered bird is One of the most colorful
found only in the Araripe birds of the rainforest of
highlands of N.E. Brazil. S. Brazil, the blue manakin
has a whining, catlike call. 9 in
22 cm

glossy
black body

4¼–5 in 3½–4 in purple-red

11–13 cm 9–10 cm patch

11–12 in
28–30 cm

PURPLE-THROATED BLACK-TAILED
FRUITCROW TITYRA
Querula purpurata Tityra cayana
Males of this distinctive This canopy bird belongs
Amazonian cotinga have to a group of big-headed
PIN-TAILED MANAKIN STRIPED MANAKIN a prominent, iridescent fruit-eaters. It nests in
Ilicura militaris Machaeropterus regulus purplish red throat patch. tree-holes, often
Males of this manakin This is an inconspicuous species It can pluck fruits while those abandoned by
from S.E. Brazil have of northern S. America. Males hovering in midair. woodpeckers.
elongated central tail feathers of the species have been
and display to females with observed making insectlike
raised rump feathers. buzzing sounds while displaying. large crest
hides the bill

brilliant
red plumage
3½ in
9 cm

GOLDEN-HEADED ANDEAN
MANAKIN COCK-OF-THE-ROCK
Pipra erythrocephala Rupicola peruvianus
The male of this The colorful crested males
S. American species displays of this large Andean cotinga
by jumping and whirring its display in groups to females.
wings—and sliding to one 11–12½ in Females nest among rocks
side or backward. 28–32 cm and rear young alone.
 T Y R A N T F LYC AT C H E R S gray head
490 A N D R E L AT I V E S
6½–8½ in 6 in 9 in
Widespread throughout the Americas, these birds 17–21 cm 15 cm 22 cm
of the family Tyrannidae often account for a third of
BIR DS • PASSER INES

all passerines in South American bird communities. brown tail

These birds are insectivorous and typically perch with pale
and wait for prey or glean it from foliage. cinnamon edge

GREAT CRESTED EASTERN WOOD PEWEE

FLYCATCHER Contopus virens
Myiarchus crinitus This bird, with its TROPICAL
A widespread tyrant flycatcher, characteristic “pewee” call, KINGBIRD
6 in this large, migratory species, hunts by sallying—taking Tyrannus melancholicus
15 cm like its relatives, catches insects off from perches to catch A large, sallying tyrant
on the wing. It often hovers insects in midair. It breeds flycatcher, this aggressively
dark brown when foraging. in eastern N. America. territorial bird breeds in open
upperparts habitats from southern
N. America to S. America.
red
underparts
4 in 7½ in 6½ in
10 cm 19 cm 17 cm

VERMILION
FLYCATCHER
Pyrocephalus rubinus COMMON TODY-FLYCATCHER CLIFF FLYCATCHER BLACK PHOEBE
A species of open country, Todirostrum cinereum Hirundinea ferruginea Sayornis nigricans
this bird forages near the This tiny C. and S. American tyrant This flycatcher from northern and This tail-wagging, tyrant flycatcher
ground. Males are vivid flycatcher is typical of a group that central S. America is swallowlike forages close to the ground in tropical
red, whereas females are forages by upward striking, but favors in its aerial foraging behavior, regions, often near water. It dives
mainly gray and white. more open habitat than its relatives. and perches on rocky outcrops. into ponds to capture minnows.

TYPICAL ANTBIRDS A N T PI T TA S A N D AU ST R A L A SI A N
From tropical American forests, these heavy-billed A N T-T H RU S H E S WRENS
birds of the family Thamnophilidae hunt insects Short-tailed antpittas spend more time on the The Maluridae are small, cock-tailed insect-eaters
near the ground, with some following army ants ground than the tree-dwelling ant-thrushes. that resemble wrens of the northern hemisphere,
to feed on insects fleeing the Both are insectivores of but are more closely related to nectar-feeding
ants. Some are long-clawed South American forests honeyeaters. Male fairy-wrens are patterned in
7 in for gripping vertical stems. and belong to the blue and black. Emu-wrens and grasswrens
18 cm families Grallariidae and are browner birds of grassy habitats.
Formicariidae, respectively.
WHITE-BEARDED
ANTSHRIKE
Biatus nigropectus MOUSTACHED ANTPITTA STRIATED GRASSWREN
This little-known, uncommon Grallaria alleni Amytornis striatus
species is restricted to 7 in A rare species found in isolated Like most other grasswrens,
S.E. Brazil, where it feeds 18 cm localities in Colombia and this C. Australian species favors
on insects in bamboo Ecuador, the moustached spinifex grass, where family 6 –7 in
forests. It is threatened antpitta inhabits undergrowth groups dart beneath bushes. 15–18 cm
by deforestation. in humid montane forest.
VARIEGATED
FAIRY WREN
TA PAC U L O S A N D GNAT E AT E R S Malurus lamberti
C R E S C E N T- C H E S T S Dumpy, short-tailed, long-legged insect-eaters 6 in
This most widespread
Australian fairy wren, like
15 cm others, builds a domed
Strong-legged and weak-flying, the of forest undergrowth, gnateaters of the family
nest. Young may stay
Rhinocryptidae and Melanopareiidae Conophagidae are secretive birds that forage close to help raise
are among the most ground adapted of South near the ground using tyrantlike the next brood.
American passerines. Some have long hind claws sallying or gleaning techniques.
to scratch for food in soil and leaf litter. They are related to antbirds.

5½–6 in COLLARED RUFOUS GNATEATER SPLENDID

14–15 cm CRESCENT-CHEST Conopophaga lineata FAIRY WREN
Melanopareia torquata More abundant than other Malurus splendens
Crescent-chests have a longer gnateater species, this bird Fairy wrens form strong
tail than tapaculos and have been from eastern S. America pair-bonds, but partners may
separated into a new family, the often moves in mixed- mate with other individuals.
Melanopareiidae. This Brazilian species flocks and uses 5 in This species is found mostly 5½ in
species occurs in arid habitats. degraded habitats. 13 cm in southern Australia. 14 cm
 BOW E R BI R DS LY R E B I R D S
A N D CAT BI R DS The Menuridae are large Australian birds that
491
white The Australasian family Ptilonorhynchidae eat ground insects. They have complex
throat consists predominantly of fruit-eaters. Male vocal organs and excel at

BIR DS • PASSER INES

bowerbirds—often brightly colored—build mimicking forest sounds.
structures called bowers to attract females. Males court females on
They mate with many females and play no display mounds by
part in raising the young. fanning their long
9–10 in tails and plumes.
9 in olive 23–25 cm 32–38 in
22 cm upperparts 80–96 cm
9 in
23 cm

GREEN CATBIRD
Ailuroedus crassirostris GOLDEN BOWERBIRD SUPERB LYREBIRD
Male catbirds—named for Prionodura newtoniana Menura novaehollandiae
their catlike call—attract The male of this small The most common lyrebird,
mates by laying leaves on species from N. Australia this species is found in forests
the ground. This species is attracts a mate by building of S.E. Australia and Tasmania.
found in New Guinea a tower of sticks up to Its lyre-shaped outer tail feathers
bright yellow and E. Australia. 10 ft (3 m) high. are adorned with notches.
underparts

AU ST R A L A SI A N 6½–7 in
16–18 cm
OV E N BI R DS
TREECREEPERS A N D R E L AT I V E S
The birds of the family Climacteridae BROWN These American birds, the Furnariidae, are
GREAT KISKADEE have evolved to resemble the TREECREEPER adept at hunting hidden invertebrates and are
Pitangus sulphuratus northern treecreepers, but are Climacteris picumnus known for the immense diversity of their nest
Widespread through This common eastern
tropical America and unrelated and, unlike them, do Australian species has a architecture. This includes stick nests, tunnel
named after its call, this not use their tail for support distinct dark-backed nests, and those that resemble clay ovens.
species is a typical sallying when climbing up trees. form in the north of the
tyrant flycatcher, but also range and a brown-
forages near the ground. backed form in the south. 7½–8 in 7–8 in
19–20 cm 18–20 cm

HON E Y E AT E R S
The birds of the family Meliphagidae are from Australia and
the southwest Pacific islands. They feed on nectar with a
10–12 in
long, brush-tipped tongue and are important pollinators 25–30 cm
of plants in that region. Other nectar-feeders, such as
sunbirds, have evolved similar features.
olive-green WHITE-EYED
wings FOLIAGE-GLEANER
Automolus RUFOUS HORNERO
7½–8½ in leucophthalmus Furnarius rufus
19–21 cm A S. American Widespread in
bird with distinctive central and southern
LEWIN’S HONEYEATER white irises, like many S. America, this species,
Meliphaga lewinii insect-eaters, this species typical of the hornero
A member of a short-billed forages in mixed flocks group, builds an
4–41/4 in group of honeyeaters, this to flush out prey. ovenlike mud nest.
10–11 cm species from eastern Australia
eats insects, fruits, and berries.
SCARLET HONEYEATER
Myzomela sanguinolenta
BLUE-FACED HONEYEATER
Entomyzon cyanotis WOODC R E EPE RS
This bird is an E. Australian A large, noisy honeyeater from The tropical American birds of
member of a group of long- Australia and New Guinea, this species
billed, flower-visiting honeyeaters is more insectivorous than many the family Dendrocolaptidae are
that often carry daubs of pollen others, but it also feeds on fruits. specialists at climbing up tree
on their forehead. trunks. They have stiffened
tails for support and strongly
clawed front toes for
5–6½ in 11½–12½ in 6½–7½ in gripping the bark.
13–16 cm 29–32 cm 16–19 cm
7½ in
TUI NEW HOLLAND 19 cm
Prosthemadera HONEYEATER SCALLOPED
novaeseelandiae Phylidonyris novaehollandiae WOODCREEPER
EASTERN SPINEBILL Although confined to Like other honeyeaters, this Lepidocolaptes falcinellus
Acanthorhynchus tenuirostris New Zealand, this species white-whiskered species from This typical buff and
Spinebills are an ancient group of is related to short-billed southern Australia and Tasmania brown woodcreeper is
heathland honeyeaters and the most honeyeaters of Australia. also takes honeydew—the found only in the forests
specialized nectar-feeders of the family. It has an extraordinary waste sugar solution produced of southeastern
This species is from E. Australia. vocal range. by certain sap-sucking insects. S. America.
 THORNBILLS PA R DA L O T E S
492 A N D R E L AT I V E S Dumpy birds from Australia,
dark facial mask
The Acanthizidae is a small family of warbler- and wrenlike, between white stripes the Pardalotidae have a
insect-eating birds from Australia and adjacent islands. stubby bill for gleaning
BIR DS • PASSER INES

It also includes Australia’s smallest bird—the sap-sucking scale insects

weebill. These birds have short wings, from trees. Brightly
a short tail, and drab-colored, colored, they nest
longish legs. 41/4–51/2 in
11–14 cm
in deep tunnels in
earth banks.

WHITE-BROWED
BUFF-RUMPED SCRUBWREN 31/4–4 in
41/4 in THORNBILL Sericornis frontalis 8–10 cm
11 cm Acanthiza reguloides Scrubwrens are birds of
Thornbills are mostly Australasian thickets. Mostly SPOTTED PARDALOTE
gray, brown, or yellow. brown, some have white head Pardalotus punctatus
This eastern species has a markings, such as this species, Three of the four species of pardalotes have
freckled forehead, like which is widespread across white spots. This highly active bird is from
many of this group. Australia and Tasmania. dry forests in S. and E. Australia.

WO O D S WA L L OW S , C ROWS A N D J AYS
BU TC H E R- BI R DS , A N D The family Corvidae, occurring worldwide,
R E L AT I V E S includes some of the biggest passerines.
From Southeast Asia, New Guinea, and Australasia, the birds They are intelligent, opportunistic birds 13–151/2 in
in the family Artamidae include wood swallows that catch that have complex social organization 33–39 cm
insects in flight and are among the few small passerines that and strong pair-bonding. Crows
can soar. The Australasian butcher-birds, currawongs, and have demonstrated tool use,
ground-dwelling Australian magpies are intelligent, highly play behavior, and EURASIAN JACKDAW
perhaps even Corvus monedula
vocal, omnivorous birds. A small crow of W. Eurasia
self-awareness. and N. Africa, this bird nests
22–27 in in cavities on crags, and is
56–69 cm found on coastal cliffs and
MASKED in urban areas.
WOOD SWALLOW COMMON RAVEN
Artamus personatus Corvus corax
71/2 in This dark-faced, thick-billed Found in a wide
19 cm wood swallow occurs in drier range of open-country
parts of inland Australia and habitats across the northern
is highly nomadic. Like other hemisphere, this is the most
wood swallows, it often widespread crow and the
gathers in large flocks. largest passerine.

10–12 in
PIED CROW 25–30 cm
Corvus albus
A relative of the raven, very long BLUE JAY
this heavy-billed bird of tail Cyanocitta cristata
AUSTRALIAN open country is perhaps This colorful jay of N. America lives
MAGPIE the most common member in tightly bonded family groups. It is fond
Gymnorhina tibicen 131/2–171/2 in
34–44 cm of the crow family in of acorns and disperses them, helping
A widespread Australian Africa and Madagascar. to distribute oak trees.
species with a highly variable
black and white plumage, this 18–20 in
bird has a varied and melodious 46–50 cm
song and is a capable mimic.
COMMON MAGPIE
Pica pica
This Eurasian bird is common
IORAS 18–19 in in habitats ranging from open
45–48 cm woodland to semidesert. It is
These rainforest birds, the Aegithinidae, are related more closely to Corvus 18 in
usually active in the high canopy. Their green crows than Asian magpies. 46 cm
or yellow coloration camouflages them in
foliage, where they glean for insects. Males
181/2–201/2 in
can engage in elaborate courtship displays. 47–52 cm

6 in ROOK CARRION CROW

COMMON IORA 15 cm Corvus frugilegus Corvus corone
Aegithina tiphia Routinely flocking This common Eurasian,
The smallest and most widespread iora, it throughout the year in largely solitary species has a
is found across tropical Asia, from India to Eurasia, this bare-faced wide-ranging diet. It feeds on
Borneo, sometimes in disturbed habitats. crow nests in colonies in small animals and vegetable
This bird builds a cup-shaped nest. trees in open countryside. matter, as well as carrion.
ORIOLES SHRIKES BUSH SHRIKES
Old World passerines related to shrikes and crows, the The family Laniidae contains predators A N D R E L AT I V E S 493
birds of the family Oriolidae live in forest canopies of open country. Many species store The family Malaconotidae is entirely
and eat insects and fruits. Many species have their prey (insects and small vertebrates) confined to Africa. They are found mostly
striking yellow and black plumage. The by impaling them on thorns. Most occur in scrubby, open woodland, and have a
females are usually greener than males. in Africa and Eurasia; two species live in hooked bill for catching large insects.
North America.
101/2–111/2 in CRIMSON-BREASTED
61/2–7 in GONOLEK
27–29 cm 17–18 cm Laniarius atrococcineus
91/2 in AUSTRALASIAN Members of the African bush shrike
24 cm EURASIAN FIG-BIRD RED-BACKED family, gonoleks have red and black
GOLDEN ORIOLE Sphecotheres vieilloti SHRIKE plumage. This species occurs
Oriolus oriolus Stout-billed relatives of Lanius collurio in southern Africa.
Breeding in woodlands orioles from Australasia, This bird breeds from
in W. and C. Eurasia, fig-birds are gregarious Europe to Siberia and
this oriole migrates fruit-eaters. This species overwinters in Africa.
southward to occurs in N. and Like other Lanius shrikes, 9 in
overwinter in Africa. E. Australia. it has a musical call. 23 cm

DRONG OS WAT T L E - E Y E S
Black, long-tailed birds of Old World tropics, A N D R E L AT I V E S
the Dicruridae sally for insects, bolting out The Platysteiridae, wattle-eyes and their
suddenly to catch their prey. relatives, form a family of insect-eating birds
They are aggressive birds, from Africa. They have a flat, hooked bill
and sometimes attack with bristles at the base. Like flycatchers,
orange- larger species to defend they snatch their prey suddenly.
red bill their nests. 10 in
26 in 26 cm
67 cm
CRESTED DRONGO
RED-BILLED Dicrurus forficatus
BLUE MAGPIE BROWN-THROATED
Like other drongos, this WATTLE-EYE
Urocissa erythrorhyncha Madagascan species has a long, Platysteira cyanea
A forest bird from the forked tail and red eyes. It has a Wattle-eyes get their
Himalayas to E. Asia, this distinctive tuft of feathers name from the red skin
species robs chicks from nests at the base of its bill. around their eyes. This
and plucks at carcasses. common species occurs in
woodland throughout 5 in
sub-Saharan Africa. 13 cm
VA N G A S
GREEN JAY A N D R E L AT I V E S
Cyanocorax yncas
This bird feeds on fruits The predatory passerines of the family Vangidae VIREOS
and seeds. S. American include the helmet-shrikes of Africa and vangas Superficially resembling American warblers,
populations of the green from Madagascar. They feed on invertebrates,
jay differ sufficiently from but somewhat thicker-billed, the members
111/2 in those of C. America to be reptiles, and frogs, and have a range of bill of the family Vireonidae are more closely related
29 cm considered as separate species. shapes—chisel-, sickle-, and dagger-shaped— to crows, and Old World orioles and shrikes.
for different prey and feeding techniques. They take insects by gleaning or fly catching,
and also eat some fruits.
WHITE
8 in HELMET-SHRIKE
20 cm Prionops plumatus
Widespread across
AZURE-WINGED sub-Saharan Africa, the
MAGPIE white-crested helmet- BLACK-CAPPED VIREO
Cyanopica cyanus shrike often gathers in small Vireo atricapilla
A sociable species, this bird groups. This bird has a wide This species breeds in N. America
12–14 in range of different calls.
31–35 cm breeds in colonies. Two separate and migrates to Mexico. Unlike
populations (in Portugal and other vireos, the sexes differ. The
in E. Asia) of this gregarious male is black-capped, while the 41/4 in
woodland bird may constitute female is gray-capped. 11 cm
separate species.
8 in
20 cm
131/2 in 43/4–5 in
34 cm 12–13 cm
RUFOUS VANGA
EURASIAN JAY Schetba rufa RED-EYED VIREO
Garrulus glandarius Common in the Vireo olivaceus
More closely related to Old forests of Madagascar, N. American populations
World crows than American this bird resembles of this very vocal vireo
jays, this colorful woodland a shrike, but is not migrate to S. America, where
bird habitually hoards closely related to the they join resident races
acorns in the fall. shrike family. of the same species.
 TYPICAL TITS 51/2–61/2 in
14–16 cm
PENDULINE TITS
494 These small, acrobatic, usually hole-nesting birds of the family These small, needle-billed birds from the family
Paridae occur in wooded habitats across North America, Eurasia, Remizidae occur in Africa and Eurasia, with
and Africa. They frequently hang upside down to glean insects one species in America. Most use cobwebs and
from foliage, and manipulate seeds and nuts to crack them open. other soft material to build flask-shaped nests
that hang from branches, often over water.
43/4–51/2 in VARIED TIT 43/4–6 in TUFTED TITMOUSE
12–14 cm Sittaparus varius 12–15 cm Baeolophus bicolor 41/4 in 31/2–41/4 in
This species is found in Like other tits, this eastern 11 cm 9–11 cm
forest habitats, including N. American species
conifers and bamboo supplements its insect diet with
forests, in N.E. Asia, BLACK-CAPPED seeds, holding them firmly to
Japan, and Taiwan. CHICKADEE smash them with its bill.
Poecile atricapillus
A typically inquisitive
and acrobatic tit, this is
a common N. American 41/2–43/4 in
bird. Like other tits, it 11–12 cm
hoards seeds for later use.

GREAT TIT BLUE TIT PENDULINE TIT VERDIN

Parus major Cyanistes caeruleus Remiz pendulinus Auriparus flaviceps
Widespread across This common bird of The only member of the Unlike most other
51/2 in Eurasia, this tit occurs broadleaf woodlands family to have a wide penduline tits, the verdin
14 cm in habitats from forest in Europe, Turkey, range across Eurasia, this makes a spherical nest.
to heathland, and has and N. Africa is a species occurs on marshes It is found in desert
a diverse range of frequent visitor to bird with trees, where it builds scrub of the S. USA
vocalization. feeders in gardens. its pendulous nest. and Mexico.

BIRDS-OF- AU ST R A L I A N ROBI NS WA X W I N G S
PA R A D I S E The Petroicidae are chunky, round-headed insect-eaters. From the family Bombycillidae, these
Mainly found in the rainforests of New Unrelated to the robins of Europe or America, they are berry-eating birds are named for the
Guinea, members of the family Paradisaeidae found from Australasia to the islands of the southwest waxy, red-tipped shafts on their
are mostly fruit-eating birds. The males Pacific. Some exhibit cooperative breeding, in which wings. Three species occur in the
display their bright, gaudy plumes in young birds help their parents to raise a new brood. cool northern forests of North
elaborate courtship America and Eurasia.
displays, and spend waxy red
most of their energy shaft on wing
on this mating ritual,
leaving the females to
rear the young alone.
5 in 6 in
13 cm 15 cm
yellow tip 7 in
121/2 in to tail 18 cm
32 cm JACKY WINTER EASTERN YELLOW ROBIN
Microeca fascinans Eopsaltria australis
LESSER BIRD- This common robin uses Common in the woodlands BOHEMIAN WAXWING
OF-PARADISE its broad bill for fly-catching. and gardens of E. Australia, this Bombycilla garrulus
Paradisaea minor It is widespread in woodland bird sallies from low perches for This sleek bird is pinkish brown, with chestnut
This species occurs across throughout Australia and invertebrates, which it snatches undertail feathers. It breeds in northern taiga forest and
N. and W. New Guinea. New Guinea. from the ground. is attracted to berried shrubs during its migration south.
Males use their long,
yellow flank plumes
and distinctive cape for
courtship displays. L ON G -TA I L E D T I T S SILKY
Birds of the family Aegithalidae are F LYC AT C H E R S
small, restless insect-eaters that The family Ptilogonatidae from Central
build dome-shaped nests that are America has just four species. These
woven with cobwebs and lined birds are named for their soft plumage
with feathers. Most species (similar to that of related waxwings)
are found in Eurasia, and their feeding habits.
yellow flank with one occurring in
plumes North America.

51/2 in
14 cm
LONG-TAILED TIT PHAINOPEPLA
Aegithalos caudatus Phainopepla nitens
The most widespread long-tailed tit, This bird of the S. US and
this woodland bird inhabits a range from Mexico nests colonially in
N. to C. Eurasia, gathering in restless woodland, but is territorial 7–81/2 in
flocks when not breeding. when breeding in deserts. 18–21 cm
MONA RC H S AU ST R A L I A N
A N D R E L AT I V E S MUDNESTERS 495
Generally long-tailed birds, members of the Monarchidae The family Corcoracidae contains
have a broad bill for fly-catching. Most species occur in two species of social birds that feed APOSTLEBIRD

BIR DS • PASSER INES

Old World tropical forests. With the exception of the on the ground and build large, Struthidea cinerea
magpie-lark, they are tree-dwellers that build cup-shaped cup-shaped nests of grass. These This ground-living bird associates
nests decorated with lichen. are held together with mud and are in flocks of 6 to 20 individuals. It
occurs in woodlands of northern 111/2–121/2 in
made in trees on horizontal branches. and E. Australia. 29–32 cm
black head

LARKS HORNED LARK

Eremophila alpestris
10–12 in This lark breeds on
26–30 cm Brown in color, with a melodious call, these
Arctic N. American and
birds from the family Alaudidae inhabit arid, Eurasian tundra. It
open habitats. Most occur in Africa, with one overwinters on coastlines
species in North America. They usually have farther south.
long hind claws that provide the stability they
need to spend so much time on the ground. 51/2–61/2 in
14–17 cm

7–8 in
18–20 cm 7–71/2 in
GREATER 18–19 cm EURASIAN
HOOPOE-LARK SKYLARK
reddish brown Alaemon alaudipes Alauda arvensis
upperparts This long-legged lark Common across
has a curved bill and Eurasia, from the
occurs in arid habitats of British Isles to Japan,
N. Africa and the Middle this open-country
East, where it habitually bird is notable for its
AFRICAN PARADISE- runs on the ground. musical aerial song.
FLYCATCHER
Terpsiphone viridis
This bird has variable color
forms, but all males have BULBULS red cheek
patch
long tail streamers. This
species occurs in savanna, Found across the warmer parts of Eurasia and
south of the Sahara. Africa, most bulbuls of the family Pycnonotidae
are gregarious, noisy, fruit-eating birds. The soft
plumage of many species is drably colored,
marked by red or yellow feathers beneath the tail.

9–10 in
23–25 cm
8 in
20 cm
61/2–15 in BLACK BULBUL
17–38 cm Hypsipetes leucocephalus
Common in forests and
long gardens in India, China,
tail and Thailand, this species
feather exists in dark-headed and
MAGPIE-LARK white-headed races. RED-WHISKERED BULBUL
Grallina cyanoleuca Pycnonotus jocosus
Unlike other members of the monarch family, A common Asian bulbul found from India to
this Australian bird spends much time on the the Malay Peninsula, this is an opportunistic
ground and builds a large mud nest. woodland species that is also found near villages.

S WA L L OW S
A N D M A RT I NS
8 in 43/4–51/2 in 43/4–6 in 6–71/2 in
20 cm 12–14 cm 12–15 cm 15–19 cm

From the family Hirundinidae,

these swiftlike birds have long
wings and a forked tail. Their
short, flattened bill and wide gape help
in taking insects on the wing. They
build mud nests or use tree-holes
or tunnels in banks. BANK SWALLOW TREE SWALLOW BARN SWALLOW
Riparia riparia Tachycineta bicolor Hirundo rustica
GREATER STRIPED SWALLOW Like other swallows, this species This N. American swallow of Occurring worldwide, this
Cecropis cucullata migrates south to overwinter in wooded swamps supplements is the most widespread
This African grassland swallow breeds the tropics. It nests in colonies its insect diet with berries. species of swallow. It was
toward the south of the continent and in riverside banks across the This enables it to breed farther originally a cave-nester, but
migrates northward to overwinter. northern hemisphere. north than other swallows. now uses buildings as well.
 BA BBL E R S , L AUGH I NG
496 T H RU SH E S , A N D R E L AT I V E S GNATCATC H E R S
Generally more gregarious, noisier, and less migratory than warblers, These small American insect-eaters from the
the babblers and laughing thrushes of the families Timaliidae family Polioptilidae are related to wrens, but are
and Leiothrichidae have evolved into a variety of warblerlike more warblerlike in appearance. In common
and thrushlike forms. Some species are brightly colored. 9 in with several members of the wren family, some
23 cm gnatcatchers cock their tail when foraging.
WHITE-EARED SIBIA 43/4 in
silver-gray Heterophasia auricularis
ear patch 12 cm
Sibias are nectar-feeding
babblers. This species is
confined to Taiwan, where its BLUE-GRAY
CHESTNUT- 6½ in distinctive call is often heard GNATCATCHER
CAPPED BABBLER 16–17 cm in mountain forests. Polioptila caerulea
Timalia pileata This N. American gnatcatcher
A bird of low thickets in 5½ in may flick its white-edged
S.E. Asia, this babbler is 14 cm tail to flush insects. Unlike
often found near water, related gnatcatcher species,
along with flycatchers males do not have dark
and other babblers. head markings.

RED-TAILED MINLA
dark red Minla ignotincta TREECREEPERS
wing patch This small babbler is similar
to a tit. It is a noisy inhabitant From the family Certhiidae, these
of montane forest canopies in are small insect-eating birds of
Nepal, China, and Myanmar.
the northern hemisphere. They
forage on vertical tree trunks
13 in
using their tail as a prop,
33 cm habitually climbing one tree and
7 in
flying to the bottom of the next.
18 cm

EURASIAN TREECREEPER
SILVER-EARED MESIA GREATER NECKLACED Certhia familiaris
Leiothrix argentauris LAUGHING-THRUSH This is the most widespread
A skulking, montane forest bird, this Garrulax pectoralis Certhia treecreeper. It is found 5 in
S.E. Asian species belongs to a group Laughing thrushes are large forest babblers with across Eurasia, from Britain 13 cm
of “song babblers,” which includes laughing calls. They often move in mixed flocks. This to Japan, in broadleaf and
minlas, sibias, and laughing thrushes. species inhabits the Himalayas and S.E. Asia. coniferous forests.

O L D WO R L D WA R B L E R S 5–6 in
13–15 cm
ICTERINE
WARBLER
5 in
13 cm
SEDGE
WARBLER
A N D R E L AT I V E S Hippolais icterina Acrocephalus
schoenobaenus
This Eurasian woodland
Several families, such as the large and widespread Sylviidae and those species has a more This is one of many
with fewer species such as Madagascar’s Bernieridae, comprise a varied musical call than other species of reed
reed warblers of the warblers (inhabitants
group of thin-billed, insectivorous birds. Some are forest species, while family Acrocephalidae. of wetland areas) that
others inhabit low bushes, dense grasses, or taller reedbed habitats. It migrates to southern breeds in Eurasia and
Many are subtly patterned and difficult to tell apart. Africa in winter. winters in Africa.

lemon-yellow
underparts
7½–9 in CAPE GRASSBIRD
19–23 cm Sphenoeacus afer 7–9½ in
18–24 cm
A bird of South African
shrubland, it belongs to an BROWN SONGLARK WRENTIT
ancient African family, the Cincloramphus cruralis Chamaea fasciata
Macrosphenidae, which This “grassbird” warbler from
evolved separately from other A dull-colored bird with
Australia in the family Locustellidae a cocked tail, the wrentit
Old World warblers. is a nomad of open habitats. Like 6 in
15 cm is the only New World
larks, it sallies skyward from member of the family
exposed perches. Sylviidae. It may be
related to parrotbills.

VINOUS-THROATED
43/4 in PARROTBILL
12 cm Sinosuthura webbiana SUBALPINE
Despite its stumpy, seed-cracking BLACKCAP WARBLER
bill, this long-tailed Asian Sylvia atricapilla Sylvia cantillans
parrotbill is a member of the Male Sylvia warblers are typically Like many Sylvia warblers,
insectivorous Old World warblers patterned with patches of black or this species breeds in scrubby
family Sylviidae. This species brown. In this widespread Eurasian 5½ in 43/4–5 in Mediterranean habitats and
occurs in China and Korea. species, females have a brown cap. 14 cm 12–13 cm winters in Africa.
BEARDED TIT GOLDCRESTS
The bearded tit forms a single-species Among the smallest of passerines, these color-crested birds of
497
family, the Panuridae. A reedbed specialist, the family Regulidae are found in cool northern forests. They
it eats insects in summer and hardens have a high metabolic rate, which forces them to feed constantly

BIR DS • PASSER INES

its stomach in winter to digest when awake. They glean tiny, soft-bodied invertebrates from
reed seeds. foliage with their needlelike bill.
6½ in
16–17 cm 3½ in
9 cm
4½ in
11 cm
BEARDED TIT GOLDCREST
Panurus biarmicus Regulus regulus
All kinglets are adapted to RUBY-CROWNED KINGLET
coniferous forests. The Eurasian Regulus calendula
goldcrest has especially grooved The red head patch of this N. American
WHITE-EYES feet and enlarged toe pads for species is visible when it raises its crown
clinging to needlelike leaves. feathers—a trait shared by all kinglets.
Most of the Zosteropidae are characterized by
a ring of white feathers around the eye. The
birds of this uniform family, closely related to
babblers, have a brush-tipped tongue and are
N U T H ATC H E S WALLCREEPER
Tichodroma muraria
nectar-feeding specialists. A N D R E L AT I V E S This central
Eurasian mountain
Nuthatches, in the family Sittidae, and the bird gleans insects
5 in
13 cm wallcreeper, sole member of the family from rocks with
Tichodromidae, are more acrobatic its pointed bill.
than treecreepers, and do not use
their tail as a support. They feed on 6½ in
16–17 cm
seeds as well as insects, sometimes
WHITE-NAPED YUHINA hoarding them in crevices.
Yuhina bakeri black stripe across eye
Now part of the white-eyes family,
crested yuhinas are adapted to take
41/4 in
nectar. The white-naped yuhina is 11 cm
an eastern Himalayan species.
5½ in
14 cm

BROAD-RINGED
WHITE-EYE
Zosterops poliogastrus
This open woodland white-eye
is restricted to isolated EURASIAN RED-BREASTED orange-brown
mountain ranges in Ethiopia, NUTHATCH NUTHATCH underparts
Kenya, and Tanzania. Distinct
Sitta europaea Sitta canadensis
races are found according 41/4 in A widespread species, this This N. American
to region. 11 cm forest bird can—like other species has a similar body
nuthatches—crack open pattern to the Eurasian
nuts by wedging them species, but males are
in cracks in tree bark. more vividly colored.
FA I RY- B L U E B I R D S
Two species of fairy-bluebirds, from the family Irenidae,
are found in Southeast Asia, where they feed on fruits— WRENS
especially figs—in forest canopies. Only With the exception of the winter
males have a vivid blue color; females wren, the family Troglodytidae is
are dull green. confined to the Americas. Most WINTER WREN
Troglodytes troglodytes
species are highly vocal, but visually 4 in The only wren whose range
inconspicuous, short-winged birds 10 cm
includes Eurasia, this bird is
bright blue that forage on insects in undergrowth; found across the northern
upper parts hemisphere. It has a large
10 in some even sleep on the ground. number of local races.
25 cm
CACTUS WREN
Campylorhynchus
brunneicapillus
The largest wren, this species
occurs in deserts of California and
Mexico, where it forages in
groups on the ground.
5½ in
ASIAN 14 cm
FAIRY-BLUEBIRD
Irena puella BEWICK’S WREN
The most widespread fairy- Thryomanes bewickii
bluebird, found from India to 7–9 in This long-tailed wren inhabits
Indonesia, this species often feeds 18–23 cm dry, open, wooded habitats in
with other fruit-eaters, such as California and Mexico. It has a
hornbills and pigeons. variety of songs in its repertoire.
 MOCKINGBIRDS CURVE-BILLED
THRASHER STA R L I NGS
498 A N D R E L AT I V E S Toxostoma curvirostre AND MYNAS
A bird of arid, scrubby
Birds of the family Mimidae occur habitats in the S. US and From the family Sturnidae, these are mostly gregarious,
throughout much of the Americas, and the Mexico, this species uses its noisy birds, many with a glossy plumage that has a
long bill to probe the soil
Caribbean and Galapagos Islands. Usually for invertebrates. metallic sheen. This family is divided into mynas and
gray or brown, these strong-legged birds are relatives from southern Asia and the Pacific, and true
highly vocal; some are accomplished mimics. starlings from Africa and Eurasia.
101/2 in
27 cm 101/2–12 in
27–31 cm

pale gray upperparts

long tail
20 in
50 cm
81/2–10 in
21–26 cm 81/2–91/2 in HILL MYNA WHITE-NECKED
21–24 cm Gracula religiosa MYNA
Found in forests of Streptocitta albicollis
NORTHERN tropical Asia, this This long-tailed, magpie-like
MOCKINGBIRD GRAY CATBIRD species is a popular cage myna is restricted to
Mimus polyglottos Dumetella carolinensis bird. It is notable for its rainforests in Sulawesi,
This N. American bird is Named for its catlike mewing melodious call and its Indonesia, and adjacent
famous for its extraordinary call, this N. American bird forages mimicking ability. islands, where it usually
repertoire of varied songs, on the ground. It overwinters in associates in pairs.
which it delivers day and night. C. America and the Caribbean.

OX PEC K E R S 10 in
25 cm
In the family Buphagidae, oxpeckers YELLOW-BILLED
OXPECKER BALI MYNA
bound low over African savannas Buphagus africanus Leucopsar rothschildi
in undulating flight. Although their Common in savanna across Found only in the
strong feet provide a tight grip on sub-Saharan Africa, this rainforests of Bali,
the hide of large animals as the birds bird habitually perches on
large mammals, eating
Indonesia, this striking
bird is highly endangered
feed, their short legs are unsuitable parasites but also 71/2–9 in by habitat destruction and
for walking on the ground. pecking at wounds. 19–22 cm the bird trade.

O L D WO R L D F LYC AT C H E R S
A N D C H ATS
5 in reddish brown
Related to thrushes, the family Muscicapidae is 13 cm base of tail
divided into two groups: true flycatchers, with a
broad bill for snatching flying insects; and chats, COMMON
which include robins, nightingales, and wheatears. STONECHAT
Some species are brightly colored, but most of these Saxicola torquatus
small birds have predominantly gray or brown plumage. Typical of chats, this is a small, 51/2 in
7 in upright, perching insectivore 14 cm
18 cm with a harsh call. It is common
in grassland in Eurasia
and Africa.
BLUE-AND-WHITE
FLYCATCHER
Cyanoptila cyanomelaena
Belonging to a large group of vivid
blue flycatchers from tropical Asia,
this species from E. Asia forages
high in forests.

71/2–81/2 in
19–21 cm
51/2 in 6–61/2 in 5–51/2 in
14 cm 15–16 cm 13–14 cm 51/2 in
14 cm
WHITE-BROWED
SHORTWING COMMON REDSTART
EUROPEAN ROBIN NORTHERN WHEATEAR MOCKING CLIFFCHAT Brachypteryx montana Phoenicurus phoenicurus
Erithacus rubecula Oenanthe oenanthe Thamnolaea cinnamomeiventris Now members of the Named for the reddish
Related to chats, this bird of Wheatears are white-rumped Belonging to a group of flycatchers family, brown tail, redstarts are
hedgerows and woodland is chats of open country. This dark-colored African chats, shortwings are running chatlike birds found mainly
found in W. Eurasia and is the most widespread this bird occurs in bushy, rocky birds found in Asian forests. in Asia. This species occurs
N. Africa. In Britain, it is a species across Eurasia. It habitats and can become tame This species occurs from the from W. to C. Eurasia and
common garden visitor. winters in Africa. near villages. Himalayas to Java, Indonesia. migrates to E. Africa.
 T H RU S H E S A N D R E L AT I V E S
Belonging to the family Turdidae, most thrushes are woodland
499
glossy blue
body birds that forage on the ground for invertebrate prey, such as
earthworms, snails, and insects. They are found worldwide,

BIR DS • PASSER INES

9 in but the majority of species occur in the Old World. Many
22 cm have melodious songs.

EUROPEAN STARLING
Sturnus vulgaris
61/2–81/2 in 9 in
Native to Eurasia, but introduced to 22 cm
N. America, this common starling 16–21 cm
roosts communally, in groups that
gather after mass aerial maneuvers.
ORANGE-HEADED THRUSH
Zoothera citrina
One of many tropical Old
SPLENDID GLOSSY 7 in World Zoothera thrushes, this
STARLING 18 cm species is found in forests from
Lamprotornis splendidus the Himalayas to Bali, Indonesia.
Widespread in sub-Saharan
Africa, this species belongs to
a group of African starlings
characterized by a
metallic sheen on black band on
their plumage. orange breast
12 in
30 cm EASTERN BLUEBIRD
Sialia sialis
A bird of eastern N. America, this is a
common species of open woodland and
fields. It sometimes nests in old
woodpecker holes.
HILDEBRANDT’S
STARLING
7–71/2 in Lamprotornis
18–19 cm hildebrandti
EMERALD This E. African
STARLING glossy starling occurs
Lamprotornis iris in wooded savanna,
This glossy W. African where it preys on large
species feeds mainly on ground insects, often
fruits, especially figs, flocking with other faint spots
but also takes ants. starling species. on breast
VARIED THRUSH
Ixoreus naevius 71/2–10 in
Found in mature coniferous forests of 19–26 cm
western N. America, this bird overwinters
in parks and gardens. Like other thrushes,
it forages in ground litter.
red or white
spot on blue
throat 8–9 in
20–23 cm
SONG THRUSH
WHITE-TAILED Turdus philomelos 8–11 in
ROBIN 7 in 20–28 cm
Found from Europe to
Myiomela leucura 18 cm Siberia, this woodland
A bird of riverine forests from and garden bird habitually AMERICAN
the Himalayas to Indochina, uses a hard surface as ROBIN
it usually keeps close to the an “anvil” to smash
Turdus migratorius
ground unless disturbed. open snail shells. Unrelated to the
European robin, this
BLUETHROAT N. American thrush
Luscinia svecica sometimes gathers in
A relative of the nightingale, the winter roosts of up to a
bluethroat breeds in damp regions quarter of a million birds.
of N. Eurasia, and migrates
to Africa and S.E. Asia. 5 in
13 cm

61/2 in
17 cm
EURASIAN
COMMON BLACKBIRD
NIGHTINGALE PIED FLYCATCHER Turdus merula 9–101/2 in
22–27 cm
Luscinia megarhynchos Ficedula hypoleuca A common long-tailed
This brown bird of This large genus of mainly thrush of woodland habitats FIELDFARE
W. to C. Eurasian thickets Asian flycatchers is ranging from Europe and Turdus pilaris
is known for its loud, closely related to chats. This N. Africa to India, the This bird breeds in N. Eurasia and
rich song, which it sings woodland species ranges 91/2–111/2 in highly territorial blackbird winters farther south, where
night or day. from Europe to Siberia. 24–29 cm often visits gardens. it flocks together in fields.
 LEAFBIRDS WA X B I L L S 4 in
10 cm
500 The fruit-eating birds of the family A N D R E L AT I V E S
Chloropseidae occur in forests in southeastern Consisting of small, highly gregarious, often
Asia. They use their brush-tipped tongue to brightly colored seed-eaters, the family
BIR DS • PASSER INES

take nectar, with which they supplement their Estrildidae occurs in tropical Africa, Asia,
diet. Male leafbirds are characteristically and Australia. Many birds live in grassland
green, with a blue or black throat. or open woodland and build domelike
nests. Both parents share 6½ in
parental duties. 16 cm
8 in
20 cm
ZEBRA FINCH JAVA SPARROW
Taeniopygia guttata Lonchura oryzivora
5½ in Native to the drier parts This vulnerable species from Java and
ORANGE-BELLIED of Australia, the zebra finch Bali frequents farmland and feeds on
LEAFBIRD 14 cm
Chloropsis hardwickei is a popular cagebird cereal crops. It has been hunted as a
around the world. rice pest and for the pet trade.
This melodious bird
inhabits the forest
canopy at high altitudes,
4 in 43/4 in
in a range from the 10 cm 12 cm
Himalayas to the Malay PURPLE
Peninsula. GRENADIER
Uraeginthus ianthinogaster
An E. African bird of
dry woodland, the purple
grenadier belongs to a
genus of predominantly
blue waxbills. COMMON
WAXBILL
W H Y DA HS Estrilda astrild
A N D R E L AT I V E S Abundant throughout
Africa, this tiny waxbill, like
4 in
African whydah birds and related indigobirds 10 cm related species, is a restless,
of the family Viduidae are cuckoolike brood flocking bird of open land, GREEN-WINGED PYTILIA
feeding on grass seeds. Pytilia melba
parasites of waxbills. The mouthparts and Male Pytilia waxbills have red splashes on
begging behavior of the chicks GREEN-BACK their wings. This African species is host to
TWINSPOT
usually resemble those of the Mandingoa nitidula blackish the paradise whydah (Vidua paradisaea),
host chicks, thus fooling bill a brood parasite.
Found in western to
the host parents. southern Africa, this thicket-
dwelling twinspot has
white-spotted underparts
and is more secretive than
other waxbills.

43/4–15 in 43/4 in 43/4 in

12–38 cm 12 cm 12 cm

black and white

“scaly” belly

EASTERN PARADISE WHYDAH SCALY-BREASTED RED-THROATED PARROTFINCH

Vidua paradisaea MUNIA Erythrura psittacea
Typical of whydahs, breeding Lonchura punctulata Parrotfinches are mostly green-bodied
males of this E. African species have This bird is common in S. Asian waxbills from S.E. Asia to the Pacific Ocean.
extraordinarily long tail feathers, scrub. Male and female are This species occurs in grasslands of the
which they use in display flights. alike in appearance. island of New Caledonia.

F LOW E R PEC K E R S OL D WOR L D DIPPERS

These dumpy birds of the family Dicaeidae S PA R ROWS Belonging to the family Cinclidae, dippers are
are related to the sunbirds from tropical Asia These stubby-billed, seed-eating birds, the the only passerines that can dive and swim under
and Australasia. Largely fruit-eating, like Passeridae, are found across Africa and Eurasia. water. They have adaptations for their aquatic
sunbirds, they take nectar from flowers. The As well as the familiar true sparrows, the group lifestyle, such as well-oiled, waterproof
flowerpeckers differ in having a shorter bill. includes snowfinches, mountain-dwelling birds feathers and oxygen-storing blood.
that occur from the Pyrenees to Tibet.
7 in
18 cm
HOUSE SPARROW 6 in
MISTLETOEBIRD Passer domesticus 15 cm WHITE-THROATED
Dicaeum hirundinaceum Originally native to DIPPER
This Australian flowerpecker Eurasia and N. Africa, the Cinclus cinclus
has a small gut to process 4–41/4 in house sparrow has adapted Widespread through temperate
mistletoe berries rapidly, 10–11 cm to human settlements Eurasia, the white-throated
and plays an important role around the world. dipper breeds near fast-flowing
in dispersing seeds of streams but may move to
this parasitic plant. slower-flowing rivers in winter.
 PIPITS AND 6 in
15 cm
5½–6½ in
14–17 cm
43/4 in WAG TA I L S 501
12 cm
BUFF-BELLIED
Present on every continent, PIPIT
Anthus rubescens
the birds of the family A typical ground-running

BIR DS • PASSER INES

Motacillidae inhabit open pipit, this species breeds in
country, where they feed on insects. RED-THROATED PIPIT Arctic tundra and winters
Most wagtails have longer tails and Anthus cervinus in fields and on coasts
olive further south.
are more brightly colored than more This pipit breeds in Arctic back
sombre pipits, and some tundra, during which it
CUT-THROAT are associated develops a colored throat:
Amadina fasciata reddish brown in males
Named for the male’s red neck with water. and pink in females.
patch, this is a common bird of dry
African woodlands, and is often 6 in 6½ in
found near human habitations. 15 cm 16–17 cm

multicolored body
6½–8 in
17–20 cm

YELLOW WAGTAIL WHITE WAGTAIL

GOLDEN PIPIT Motacilla flava yellow Motacilla alba
Tmetothylacus tenellus underparts
This widespread Eurasian A typical wagtail, this
A bird of open bush and species overwinters in Africa, species is widespread
grassland, the golden pipit India, and Australia. There are throughout Eurasia—
is confined to E. Africa, several races, including many often on farmland
from Sudan to Tanzania. with gray or black head patterns. or in towns.

SUNBIRDS
The small, fast-moving, nectar-feeding Old World
tropical birds of the family Nectarinidae are similar 4 in
to American hummingbirds, with their long, long, 10 cm
purple chest decurved bill
decurved bill and long tongue. Males are usually
brightly colored with a metallic sheen. These
birds are fiercely territorial. 6 in
15 cm

5½ in
14 cm 7 in
18 cm
brilliant
scarlet breast
STREAKY-BREASTED
SPIDERHUNTER
Arachnothera affinis
Spiderhunters are drab,
long-billed members of the
sunbird family. Like other SCARLET-CHESTED PURPLE SUNBIRD
GOULDIAN FINCH sunbirds, this S.E. Asian SUNBIRD Cinnyris asiaticus
Erythrura gouldiae species eats invertebrates Chalcomitra senegalensis Like other sunbirds, this
A brilliantly colored relative of as well as nectar. Common throughout much of species from S. Asia feeds
parrotfinches, this endangered bird sub-Saharan Africa, this large its young mostly on insects.
of northern Australia is nomadic. sunbird occupies a variety of The male loses its brilliant
Males are red- or black-faced. wooded habitats. plumage after breeding.

W E AV E R S RED-COLLARED
WIDOWBIRD ACC E N TORS
Euplectes ardens
Gregarious seed-eaters, the Ploceidae Breeding males of widowbirds Mostly ground-dwelling passerines,
build elaborate nests. Males usually take are black and some fan enlarged the narrow-billed Prunellidae are
sole responsibility for this duty; females tails during display flights. This found in Eurasia. Most species
use the nest as the basis for choosing species is widespread in are adapted to high altitudes, but
sub-Saharan Africa.
mates. Most species are African, with they move to lower altitudes
a few found in southern Asia. in winter to supplement their
4–41/4 in insectivorous diet with seeds.
10–11 cm
6 in 41/4–5 in 6–16 in
15 cm 11–13 cm 15–40 cm YELLOW-CROWNED
BISHOP DUNNOCK
RED-BILLED QUELEA Euplectes afer Prunella modularis
CHESTNUT WEAVER Quelea quelea This African bird is Unlike other accentors,
Ploceus rubiginosus Widely considered the related to widowbirds. this is a lowland bird,
The genus Ploceus includes world’s most abundant bird, Breeding males are and it does not usually
the largest number of this African species gathers vibrantly patterned; gather in flocks. It 6 in
species of weavers. This in giant flocks that can cause females and nonbreeding is widespread in 15 cm
species occurs in E. Africa. serious damage to crops. males are red or black. temperate Eurasia.
 FINCHES AND 43/4 in
502 R E L AT I V E S 12 cm

Finches of the family Fringillidae have diversified 43/4–5 in 6 in

in Eurasia, Africa, and the tropical regions of the 12–13 cm 15 cm
BIR DS • PASSER INES

Americas. From thin-billed nectar-feeders to heavy-

billed, seed-cracking grosbeaks and hawfinches,
these birds have evolved to cope with a wide
range of foods.
EURASIAN GOLDFINCH AMERICAN GOLDFINCH CHAFFINCH
Carduelis carduelis Carduelis tristis Fringilla coelebs
6½ in
17 cm Goldfinches have a thinly pointed Bright yellow goldfinches and The most common finch in
bill for taking seeds from the siskins have diversified in the Europe, the chaffinch is also
seed heads of tall plants, such Americas, especially in found throughout N. Asia.
as thistles. This species is S. America. This migratory It often forages with other
widespread in Eurasia. species is from N. America. finch species in winter.
large, white
wing patch
6–6½ in
15–17 cm
43/4 in
12 cm

GRAY-CROWNED EVENING GROSBEAK

RED CROSSBILL YELLOW-FRONTED CANARY ROSY FINCH Hesperiphona vespertina
Loxia curvirostra Serinus mozambicus Leucosticte tephrocotis Grosbeaks of the finch family have a
Crossbills use their uniquely crossed This bird belongs to a group A member of a group of mountain heavy, seed-crushing bill, a feature they
mandibles for extracting seeds from cones. of mostly yellow-colored African finches related to bullfinches, share with the unrelated grosbeaks
This species is widespread in coniferous serins and canaries. It is common this N. American species is an of the cardinal-grosbeak family.
forests across the northern hemisphere. south of the Sahara. inhabitant of high, rocky landscapes. This is a N. American species.

A M E R ICA N BL AC K BI R DS
A N D R E L AT I V E S
8½–10 in 6½–9½ in 7½–10 in
These American species of the family Icteridae superficially 21–26 cm 17–24 cm 19–26 cm
resemble the common blackbird, to which they are not
related. As a group, these strong-billed birds are more
closely related to finches. The bill has a powerful gaping
action for prying apart tough food.

RED-WINGED BLACKBIRD EASTERN MEADOWLARK

Agelaius phoeniceus Sturnella magna
COMMON GRACKLE YELLOW-HEADED A N. American bird of marshy habitats, A species from eastern N. America,
Quiscalus quiscula 11–13½ in BLACKBIRD this colonial nester often breeds in the this open-country bird builds a nest
An opportunistic N. American 28–34 cm Xanthocephalus xanthocephalus company of the larger, more dominant, on the ground, covering it with
species, this bird raids garbage This marshland bird of western yellow-headed blackbird. a grass roof.
dumps and corn crops. N. America breeds in colonies and
Its internally keeled bill builds its nests over water, perhaps
can saw open acorns. to deter predators. 6–8 in 14½–18 in
15–20 cm 37–46 cm
black head

7½–9 in 7–8 in
19–22 cm 18–20 cm

orange
underparts

BROWN-HEADED COWBIRD BALTIMORE ORIOLE BOBOLINK CRESTED OROPENDOLA

Molothrus ater Icterus galbula Dolichonyx oryzivorus Psarocolius decumanus
This N. American bird produces Feeding on insects and caterpillars This ground-nesting N. American Like other tropical American
many eggs, laying them in the nests in spring and summer, this species bird gets its name from its oropendolas, this colonial species
of other passerines. Its young are changes its diet in winter to include bubbling flight song. It migrates weaves long nests that hang from
raised by many different host species. nectar and berries. to central S. America in winter. tips of branches in open forest.
 yellow 41/4 in 4 in 5½ in
forehead 11 cm 10 cm PURPLE-THROATED 14 cm HOUSE FINCH
EUPHONIA Carpodacus mexicanus 503
Euphonia chlorotica The N. American house
Euphonias are small birds that finch is a type of rosefinch.
mainly eat fruits, especially Rosefinches are especially

BIR DS • PASSER INES

mistletoe berries. This species diverse in temperate Asia. The
is from northern S. America. males are red or pale colored.

BLUE-NAPED CHLOROPHONIA
Chlorophonia cyanea
The fruit-eating chlorophonias are
mainly green birds. This species is
widespread in forests across S. America.

8 in 8 in 6–6½ in 5½ in 9 in
20 cm 20 cm 15–16 cm 14 cm 22 cm

PINE GROSBEAK EURASIAN BULLFINCH IIWI JAPANESE GROSBEAK

Pinicola enucleator Pyrrhula pyrrhula Drepanis coccinea Eophona personata
The pine grosbeak occurs in Bullfinches have a short, thick bill This long-billed nectar-eater is A strikingly patterned
coniferous forests in the northern and a thickset head. This is the most found only in Hawaii. It is a bird bird, the Japanese grosbeak
hemisphere. The bill of this widespread of the bullfinches and of montane forests, the largest of cold northern forests
grosbeak resembles that of the is found in woodlands across populations occurring at breeds in Siberia and N. Japan
closely related bullfinches. temperate Eurasia. higher elevations. and winters in S. China.

black and white

AMERICAN streaks overall 4½–5½ in
11–14 cm
WA R B L E R S
Unrelated to warblers of the Old BLACK-AND- YELLOW-BREASTED
World, these finch-related insect- WHITE WARBLER CHAT
Mniotilta varia Icteria virens 7 in
eaters of the family Parulidae are found This N. American warbler This N. American bird is 18 cm
throughout North and South America. moves up and down tree relatively large for a warbler.
The tropical species are sedentary, but trunks like a nuthatch, It sings at night as well as
temperate ones are migratory. The males and has long hind claws during the day, and mimics
for clinging to bark. the calls of other birds.
molt their bright colors in winter.
5½ in
14 cm
5 in 41/4–5 in
yellow body 13 cm 11–13 cm

43/4–5 in
12–13 cm

YELLOW WARBLER BAY-BREASTED WARBLER HOODED WARBLER AMERICAN REDSTART

Setophaga aestiva Setophaga castanea Setophaga citrina Setophaga ruticilla
Widespread from This bird breeds in spruce Like the American redstart, An active forager, this
N. America to the forests in eastern N. America. this bird from broad-leaved N. American warbler habitually
Caribbean, there is a Its numbers fluctuate according woodland of the E. US feeds flashes its orange and black wings
large number of local to the abundance of its prey, on insects by catching flies and tail to flush out insects. It
races of yellow warblers. the spruce budworm. while in flight. also catches prey in the air.

dull brown
feet and legs 6 in 43/4 in
15 cm 12 cm

5 in
13 cm

41/4–5½ in
11–14 cm
NORTHERN PROTHONOTARY COMMON
WATERTHRUSH WARBLER YELLOWTHROAT
Parkesia noveboracensis Protonotaria citrea Geothlypis trichas GOLDEN-WINGED WARBLER
A large, tail-bobbing ground Unusually for American A bird of wet habitats, Vermivora chrysoptera
warbler of N. America, this warblers, this species of this is a migratory species A warbler of eastern N. America,
bird forages in leaf litter of densely wooded swamps nests like other N. American this bird breeds in scrubby, open
wet woodlands and nests in tree cavities—sometimes warblers. It winters in habitats and farmland, and benefits
in thickets near water. using old woodpecker holes. California and Mexico. from deforestation.
 B U N T I N G S A N D R E L AT I V E S A M E R I C A N S PA R R O W S
504 True buntings of the Old World in the family Emberizidae Birds in the family Passerellidae resemble Old World buntings rather than
are mostly ground-feeding seed-eaters. Longspurs, in both true sparrows, especially in the bill structure, with a small, sharp upper
Old and New Worlds, are similar short-legged ground mandible and often slightly kinked cutting edge. Many also have striped 7½ in
BIR DS • PASSER INES

birds of open spaces, from plains and tundra head patterns, with or without streaked flanks—others are more uniform 19 cm
to mountain tops in the family Calcaridae. in color. They search for seeds while hopping on the ground.

BLACK-HEADED
BUNTING
Emberiza melanocephala
An inhabitant of scrub and dark gray
olive groves, the black-headed body
bunting breeds in the
Middle East and
winters in India.

YELLOW-THIGHED FINCH
6½ in Pselliophorus tibialis
17 cm A noisy, tropical member of the
male yellow beneath American sparrow family, this
and has a black cap species is found only in the montane
6–6½ in rainforests of Costa Rica and Panama.
15–16 cm

9 in
22 cm

DARK-EYED JUNCO
LAPLAND LONGSPUR Junco hyemalis
Calcarius lapponicus Juncos are gray-brown, SPOTTED TOWHEE
Relatives of buntings, longspurs ground-flocking Pipilo maculatus
6 in have been named for their long N. American sparrows. This Towhees are long-tailed American
15 cm hind claw. This species has a species, locally called the sparrows. The spotted towhee is a
circumpolar distribution snowbird, is a common winter reddish-brown-sided bird found in
during the breeding season. visitor of garden bird feeders. thicket habitats across N. America.

TA N AG E R S A N D R E L AT I V E S 7 in
18 cm
An American family, the Thraupidae consist of gaudily colored
tanagers, found in the forests of tropical South America. These 5½ in 7–7½ in
14 cm 18–19 cm
finch-related birds have evolved to exploit a range of food
sources, including fruits, insects, seeds, and nectar.

VARIABLE SEED-EATER
Sporophila corvina
Belonging to a group of tropical
American, stubby-billed
seed-eaters, this bird has a
variable plumage pattern.
4¼ in
11 cm
GREEN HONEYCREEPER SCARLET TANAGER WESTERN TANAGER
Chlorophanes spiza Piranga olivacea Piranga ludoviciana
This stocky species lives in In most species of Piranga This bird breeds in western
forest canopy and uses its tanager, the males turn N. America and overwinters in
5½ in stout, decurved bill to eat red during breeding. C. America. It is the only Piranga
14 cm fruits. It is often seen in This migratory species tanager where the breeding male
BLACK-MASKED
FINCH mixed flocks of tanagers. is from N. America. is predominantly yellow.
Coryphaspiza melanotis
A ground-feeding finch of
grassland habitats in central 6 in 7 in
S. America, this species 15 cm 18 cm
has been assigned to
the tanager family.

LARGE GOLDEN-COLLARED BLUE-WINGED

GROUND FINCH TANAGER MOUNTAIN TANAGER
Geospiza magnirostris Iridosornis jelskii Anisognathus somptuosus
This seed-eater is a native A member of a group of Most mountain tanagers
of the Galapagos Islands. Andean forest tanagers of northern S. America are
The species feeds less with a yellow head streak, colored blue and yellow,
6 in from the ground than this species occurs in including this inhabitant
15 cm other ground finches. Peru and Bolivia. of montane rainforests.
 6½–7½ in pale gray band
17–19 cm above eye
505
FOX SPARROW
Passerella iliaca
Widespread throughout
N. America, this large sparrow
has a red-streaked back and
chest. It usually forages
in low vegetation.
LARK-BUNTING 7 in
Calamospiza melanocorys 18 cm
A bird of the N. American
prairies, the lark-bunting is a
ground-nesting member of
the American sparrow group.

long, rounded,
plain tail variably colored 5–5½ in
belly, streaked 13–14 cm
blackish, brown,
or red

6½–7½ in 5½–6½ in
17–19 cm 14–16 cm

WHITE-CROWNED
SPARROW SONG SPARROW CHIPPING SPARROW
Zonotrichia leucophrys Melospiza melodia Spizella passerina
This N. American bird is usually A common N. American bird from A rufous (reddish brown)-capped bird
found in low vegetation or on Alaska to Mexico, this species has a common in open woodland across
the ground. It has distinctive large number of races. It is named N. America, the chipping sparrow has
black and white head markings. for its melodious song. a distinctive “chipping” trill.

RED-CRESTED CARDINAL
Paroaria coronata
C A R D I N A L S A N D R E L AT I V E S
Birds of the Paroaria genus are found in These mostly chunky-billed birds are seed-eaters like buntings,
S. America. This is a common species and many have bright colors like tanagers. Related to both, the
in open woodland.
Cardinalidae belong to a large group of American passerines
7½ in that are derived from finches.
19 cm
8½–9 in
21–23 cm
5 in 7–8½ in
13 cm 18–21 cm

ROSE-BREASTED NORTHERN
GROSBEAK CARDINAL
Pheucticus ludovicianus Cardinalis cardinalis
breeding male has This migratory American cardinal The males of this resident of the
blue underparts grosbeak has a typically heavy bill E. US and Mexico are red, due to
and head for feeding on large seeds and on chemicals called carotenoids that are
insects, such as beetles. acquired from their diet.

RED-LEGGED green back 5½ in

HONEYCREEPER 14 cm
Cyanerpes cyaneus
Tropical American honeycreepers
feed on nectar. The most 5 in
widespread species, this bird has 13 cm
a decurved bill. Like females,
males develop a dull green
“eclipse” plumage after breeding.

5 in
BLUE-NECKED 13 cm
TANAGER
Tangara cyanicollis PAINTED BUNTING INDIGO BUNTING
This open forest bird from Passerina ciris Passerina cyanea
northern S. America belongs This species breeds in the S. US The most wide ranging of the blue
to a group of tanager species and overwinters in C. America and buntings, this species migrates between
with particularly colorful, the Caribbean. Only males have the Canada and S. America. The males lose
iridescent plumage. striking tricolored pattern. their blue plumage in winter.
506 MAMMALS
The mammals are very successful animals. They are able to occupy
ANIMALS • MAMMALS

almost every terrestrial habitat, and some visit the deep oceans
between breaths of air. However, this has not always been the case;
at 210 million years old, mammals are a relatively recent group.

PHYLUM CHORDATA Mammals owe their success to a unique combination

CLASS MAMMALIA of adaptations that allowed them to replace reptiles
ORDERS 28 as the dominant form of animal life on Earth. Like
FAMILIES 160
SPECIES About 6,300
reptiles, mammals breathe air, but unlike their scaly
ancestors, they are warm-blooded, so they are able
to burn fuel (food) in order to maintain a constant,
warm body temperature. This creates conditions in
which the internal chemical processes that sustain life
can happen most efficiently, without relying directly
on heat from the sun. Uniquely, mammals have hair,
which reduces heat loss and allows an animal to be
active in cold climates and at night. Because fur can be
molted and regrown, it is seasonally adjustable, too.
A single jawbone articulated directly
to the skull gives mammals, such as this va r i at i o n s o n a t h e m e
Tasmanian devil, a powerful bite.
The basic mammal skeleton is strong, with upright
limbs supporting the body from below. This
arrangement allows land-dwelling mammals to
walk, run, and leap, but the basic structure is highly
adaptable, and has been modified for swimming, as
in seals and cetaceans; for flying, as in bats; and for
climbing or swinging through the trees, as in some
primates. The mammalian skull has powerful jaws,
Baleen is made of a protein called with a single lower jawbone articulating directly with
keratin, and baleen whales use it to strain
sea water, trapping food in their mouth. the skull, and a diverse range of teeth adapted to
a huge variety of food types. Certain other bones,
which in reptiles contribute to the lower jaw, are put
to a new use by mammals—they have become the
three tiny bones of the inner ear, allowing for greatly
enhanced hearing. The mammalian skull also serves
to protect the brain, which is larger in mammals
than in other groups, hinting at increased processing
power. With a big brain comes intelligence, and an
By sucking milk from their mother,
young warthogs obtain all the nourishment unrivaled ability to learn, to remember, and to engage
they need during the first weeks of life. in complex behavior.
These abilities take time to fine-tune, and young
D E B AT E mammals do so during the extended period of parental
E Q U A L LY W E L L A D A P T E D care that begins while they are being fed on milk
produced by their mother’s mammary glands. These
Marsupial features are sometimes
described as primitive. This implies
unique organs evolved from sebaceous glands, which
that marsupials, whose young are originally provided secretions to condition the skin and
born in an embryonic state and develop perhaps to prevent eggs from drying out, and it is for
in a pouch, are less advanced than them that the entire group is named.
placental mammals, which bear
conventional young. In fact, the two
groups emerged at the same time FUR AS INSULATION
about 175 million years ago. The air trapped in a sea otter’s dense fur
insulates it from the cold water in which it lives.
m a m m a l g ro u p s
There are three groups of mammals—the egg-laying monotremes, the
pouched mammals (or marsupials), and the placental mammals. The
latter group is the most diverse, and is here divided into further groups.
e g g - l ay i n g ro d e n t s
mammals

508 533
pouch ed t r e e s h r e ws
mammals

509 543
sengis c o lu g o s

518 543
tenrecs and pr i m at e s
golden mole s

519 544
a a r dva r k bat s

520 562
du g on g a n d h edgehogs , moles ,
m a nat e e s a n d r e l at i v e s

521 570
hyraxes pa n g o l i n s

521 573
e l e ph a n t s ca r n i vo r e s

524 574
a r m a di llos odd-toed
u n g u l at e s

525 602
sloths an d ev en-toed
a n t e at e r s u n g u l at e s

528 608
r a bbits , whales,
hares, and por poises ,
pi k a s a n d d o l ph i n s

529 628
508 E G G - L AY I N G M A M M A L S
There are just five species in this Monotremes have a specialized snout for finding PHYLUM CHORDATA
group of mammals, which are known and eating prey. The platypus, which is partly
MAMMALS • EGG-LAYING MAMMALS

CLASS MAMMALIA

as monotremes. All of them have aquatic, has a flattened, ducklike bill covered ORDER MONOTREMATA

a specialized snout and lay eggs. with sensory receptors that allow the animal to FAMILIES 2

locate invertebrates under water, even in murky SPECIES 5

The monotremes, which are the duck-billed conditions. The long, cylindrical snout and long
platypus and the long-nosed and short-nosed tongue of the land-dwelling echidnas are ideal for
echidnas, form the order Monotremata. They probing the nests of ants and termites, and for
are found in various habitats in New Guinea, catching worms. Neither the platypus nor the
Australia, and Tasmania. echidnas have teeth; instead they have grinding
Platypuses and echidnas lay soft-shelled eggs surfaces or horny spines on their tongue.
that hatch after around ten days’ incubation. The
young feed on milk secreted from the female’s w h at ’ s i n a n a m e
mammary glands; monotremes have no teats. The word “monotreme” means “single hole” and
Infant echidnas live in their mother’s pouch makes reference to the cloaca, a single posterior
The large, webbed forefeet of the
until their spines appear and then, like young body opening into which the digestive, urinary, duck-billed platypus provide propulsion.
platypuses, stay in a burrow for several months. and reproductive tracts empty. The hind feet and tail are used to steer.

D U C K- B I L L E D P L AT Y P U S
short, dense sensitive,
The sole member of the family Ornithorhynchidae, the body fur small eyes ducklike bill
duck-billed platypus is well adapted to a semiaquatic
lifestyle. It has a streamlined body,
waterproof fur, webbed feet, and
a flattened tail. Males have a
venomous horny spur on
each hind foot.
16–25 in
40–63 cm

DUCK-BILLED PLATYPUS
Ornithorhynchus anatinus
Found in rivers and
streams in E. Australia and
Tasmania, this rare species uses
its soft bill, which is covered in
electroreceptors, to hunt
venomous spur for invertebrates.
of male

sharp spines
ECHIDNAS for defense

The family Tachyglossidae is made

up of the long- and short-beaked
echidnas. Their rounded body is 19–25 in
covered with fur and spines, and the 48–63 cm
long snout is ideal for seeking out
insects, ants, and worms.

12–18 in
30–45 cm

SHORT-BEAKED ECHIDNA EASTERN LONG-BEAKED

Tachyglossus aculeatus ECHIDNA
Widespread in Australia, Tasmania, Zaglossus bartoni
and New Guinea, the short-beaked The largest of the monotremes,
echidna lays a single egg into a this species lives in the forested
pouch on its abdomen. highlands of E. New Guinea.
POUCHED M A M M ALS 509
These mammals bear live young that in a protective pouch. Some marsupials have only PHYLUM CHORDATA
are born in a very immature state, one offspring at a time, but others may have up to

MAMMALS • POUCHED MAMMALS

CLASS MAMMALIA

and typically complete their growth a dozen or more. The phase during which young ORDERS 7

in the mother’s abdominal pouch. are in the pouch is equivalent to the gestation FAMILIES 18

period of placental mammals. SPECIES OVER 350

Marsupials inhabit a vast array of habitats from Kangaroos and some other marsupials are able
desert and dry scrub to tropical rainforest. Most to stop the development of an embryo before it D E B AT E
of them are either terrestrial or arboreal, but implants in the uterus if they already have an NOT SO PRIMITIVE?
several glide, one is aquatic, and there are two infant occupying the pouch. The pregnancy
marsupial moles that live underground. The diets continues when the pouch is vacated. Pouched mammals were once
of marsupials are equally wide-ranging: among considered primitive because their
young are born at an early stage
the different species are carnivores, insectivores, th e sev en or ders
without prolonged nourishment by a
herbivores, and omnivores. There are even Pouched mammals, known as marsupials, are now placenta inside the mother. Scientists
marsupials that feed on nectar and pollen. divided into seven major groups. These are: the now know that this reproductive
Marsupials also differ markedly in size, from American opossums, order Didelphimorphia; strategy is an adaptation to their
the tiny planigales, which are among the the shrew opossums, order Paucituberculata; environment and that pouched
smallest mammals in the world, weighing less monito del monte, the sole member of the order mammals are just as advanced as
than about ⅓ oz (4.5 g), to the red kangaroo, Microbiotheria; the Australasian carnivorous placental mammals. Kangaroos can
males of which can weigh over 200 lb (90 kg). marsupials, order Dasyuromorphia; bandicoots, support two dependent juveniles of
order Peramelemorphia; the marsupial moles, different ages, and a fertile egg that
can develop almost immediately
e a r ly d e v e l o p m e n t order Notoryctemorphia; and the Australasian
should anything happen to the other
Young marsupials are born blind and furless. marsupials of Diprotodontia, the largest order two. This strategy means pouched
They make their way through the mother’s fur of pouched mammals, which includes mammals recover quickly when harsh
and latch on to a nipple and suckle. In around the koala, wombats, possums, environmental conditions abate.
half of marsupial species, the nipples are located wallabies, and kangaroos.

SHREW OPOSSUMS MARSUPIAL SOUTHERN

MARSUPIAL MOLE
Fewer incisor teeth distinguish the members of MOLES Notoryctes typhlops
This marsupial mole is
the family Caenolestidae from other American Two species of Australian marsupial specialized for burrowing
marsupials. All eight species occur in the Andes moles form the family Notoryctidae. 41/2–51/2 in in the sandy desert and
in western S. America. 11–14 cm spinifex grassland of
Their short limbs, large claws, and horned C. Australia.
nose-shield help them to burrow. They lack
external ears and have nonfunctional eyes.

31/2–51/2 in
9–14 cm
OPOSSUMS
The New World opossums of the
family Didelphidae have a pointed
DUSKY SHREW OPOSSUM muzzle with sensitive whiskers,
Caenolestes fulginosus
Living at high altitudes in Colombia, Ecuador, and naked ears. Many species have
and Venezuela, this shrew opossum uses its a grasping, prehensile tail that
large lower incisors to kill prey. helps when climbing. Some
opossums lack pouches.

MONITO DEL MONTE

VIRGINIA OPOSSUM
The sole member of the family Microbiotheriidae, Didelphis virginiana
the monito del monte is well adapted to the cold. It 141/2–20 in The largest American marsupial, 61/2–11 in
uses both daily torpor and seasonal hibernation to 37–50 cm this species lives in grasslands 16–28 cm
conserve energy when temperatures are low or food and temperate and tropical
forests in the US, Mexico,
is in short supply. and C. America.
8–121/2 in
31/4–5 in 20–32 cm
8–13 cm BROWN-EARED
MONITO DEL MONTE WOOLLY OPOSSUM
Dromiciops gliroides Caluromys lanatus
Inhabiting cool bamboo Also known as the western BARE-TAILED WOOLLY OPOSSUM
forest and temperate woolly opossum, this arboreal, Caluromys philander
rainforest in Chile and solitary opossum inhabits This woolly opossum’s long, prehensile tail helps
Argentina, this species has moist forests in western it move through the canopy of moist rainforests
dense fur to limit heat loss. and central S. America. across eastern and central S. America.
 OPOSSUMS
510 41/4–5 3/4 in 43/4–9 in
11–14.5 cm 12–22 cm

10–16 in
MAMMALS • POUCHED MAMMALS

26–40 cm

dark fur
around WOOLLY MOUSE OPOSSUM WATER OPOSSUM
large eyes Marmosa sp. Chironectes minimus
Living in C. and S. America, this pouchless The only aquatic marsupial, this opossum
marsupial with thick, woolly fur is arboreal, of C. and S. America is also unique in that both sexes
nocturnal, and omnivorous. have a pouch, but it is only watertight in females.

31/2–5 1/2 in 4 3/4–5 3/4 in

9–14 cm 12–14.5 cm
LINNAEUS’S MOUSE OPOSSUM
Murinus murina
Widespread in forests, pampas, and
plantations in S. America, this agile,
nocturnal climber has a long, prehensile tail.

ELEGANT FAT-
prehensile TAILED MOUSE
tail OPOSSUM
Thylamys elegans
Like several other
opossums, this marsupial
from Chile stores fat
dependent in its tail as winter
young approaches.

PATAGONIAN OPOSSUM
white spot Lestodelphys halli
above eye Living in shrub, savanna, and
grassland in Argentina, the
Patagonian opossum has the
most southerly distribution
of any opossum species.

4 3/4–7 in
12–18 cm

8–13 in
20–33 cm

GRAY FOUR- GRAY SHORT-TAILED

EYED OPOSSUM OPPOSUM
Philander opossum Monodelphis domestica
This opossum has white Found in Argentina, Brazil,
spots on its forehead, Bolivia, and Paraguay, this
making it look as if it short-tailed opossum
has four eyes. It lives sometimes inhabits human
in Mexico, and dwellings, as well as forest,
C. and S. America. scrub, and grassland.

N U M BAT BILBIES 12–22 in

30–55 cm
The numbat is the only member of the family Myrmecobiidae. The family Thylacomyidae has only one species,
It has a distinctive striped coat, strong claws for digging, since the lesser bilby has been declared extinct.
and a very long tongue for extracting termites Living in arid habitats, this nocturnal marsupial
from their nests. does not need to drink water, as it absorbs
moisture from its food.
NUMBAT 9–111/2 in
Myrmecobius fasciatus 22–29 cm
Found only in eucalyptus GREATER BILBY
forest and woodland in Macrotis lagotis
Western Australia, this A burrowing species of the desert of
diurnal marsupial is a C. Australia, this marsupial has silky fur,
specialized termite-eater. a tricolored tail, and long, rabbitlike ears.
QUOL L S , DU N NA RT S , white band on
A N D R E L AT I V E S rump and chest 511
With strong jaws and sharp canine teeth, the family Dasyuridae is
made up of more than 70 large and small carnivorous marsupials.
These animals also have sharp claws, except on their big toe. fat store in
base of tail

33/4–41/4 in
9.5–10.5 cm TASMANIAN DEVIL
Sarcophilus harrisii
The largest carnivorous marsupial in
FAT-TAILED FALSE ANTECHINUS the world, this nocturnal hunter lives in
Pseudantechinus macdonnellensis a variety of habitats across Tasmania.
This nocturnal insect-eater stores fat in
the base of its tapered tail. It inhabits arid 221/2–26 in
rocky habitats of C. and W. Australia. THREE-STRIPED 57–65 cm
DASYURE
Myoictis sp.
This marsupial’s coloration
23/4–51/2 in helps it to remain
7–14 cm BROWN camouflaged on the
ANTECHINUS rainforest floor in the
Antechinus stuartii islands of Indonesia
Antechinus males die of and New Guinea.
stress and exhaustion after
their first breeding season.
This species is endemic to 71/2–91/2 in 43/4–9 in
forests in E. Australia. 19–24 cm 12–23 cm

10–16 in
26–40 cm
RED-TAILED PHASCOGALE CREST-TAILED MULGARA
WESTERN QUOLL Phascogale calura Dasycercus cristicauda
Dasyurus geoffroii Possessing a brush of black hair on its 41/4–5 in This carnivore from W. and C. Australia
Also called the chuditch, this nocturnal red-based tail, this carnivorous marsupial 10.5–12.5 cm inhabits arid and semiarid habitats,
hunter from S.W. Australia is mainly ground- inhabits woodland in S.W. Australia. such as desert, heath, and grassland.
dwelling, although it can climb trees. It stores fat in its tail.
2–3 in
5–7.5 cm
23/4–4 in 2–3 in 21/4–31/2 in
7–10 cm 5–7.5 cm 6–9 cm

KULTARR
Antechinomys laniger
This fast, agile marsupial
uses its large hind feet to NARROW-NOSED PLANIGALE INLAND NINGAUI FAT-TAILED DUNNART
bound across woodland, Planigale tenuirostris Ningaui ridei Sminthopsis crassicaudata
grassland, and semidesert This flat-headed, nocturnal, rodentlike A shrewlike marsupial with a pointed snout, Living in open grassland in southern
habitats in southern marsupial lives in low shrub and dry this nocturnal predator hunts insects in arid Australia, this small, nocturnal
and C. Australia. grassland in S.E. Australia. spinifex grassland in C. Australia. marsupial stores fat reserves in its tail.

BANDICOOTS 9–15 in
22.5–38 cm
11–14 in
28–36 cm
SPINY BANDICOOT
These omnivorous marsupials are Echymipera kalubu
found across Australia and New Guinea. This forest-dwelling,
Characterized by fused second and third nocturnal insect-eater of
New Guinea has a conical
toes on their hind feet and three pairs of snout, spiny coat, and
lower incisor teeth, the Peramelidae also a hairless tail.
have short, coarse, or spiny hair.
coarse, yellowish
brown fur
melanistic (black)
12–171/2 in form of species 101/2–14 in
31–44.5 cm 27–35 cm

LONG-NOSED EASTERN BARRED

BANDICOOT BANDICOOT
Perameles nasuta Perameles gunnii
Found in rainforest and Named for the creamy SOUTHERN BROWN BANDICOOT
woodland in eastern bands of fur on its Isoodon obesulus
coastal Australia, this flanks, this bandicoot This short-nosed bandicoot lives
nocturnal bandicoot lives in grassland and in the shrubby heathland in southern
forages for food by grassy woodland in Australia and several islands, including
digging for insects. Australia and Tasmania. Kangaroo Island and Tasmania.
 KOA L A PYG M Y P O SSU M S
512 The sole member of the family large, white, These small, nocturnal marsupials with
Phascolarctidae, the koala is rounded ears a prehensile tail are omnivorous, feeding
adept at climbing because on insects, fruits, nectar, and pollen.
MAMMALS • POUCHED MAMMALS

of its powerful forearms, Four species of the family Burramyidae

opposable fingers and toes, and are endemic to Australia; the fifth lives
sharp, curved claws. It sleeps in Australia and New Guinea.
for up to 20 hours each day
because its diet of eucalyptus 4¼ in
leaves is low in nutrients. 10.5 cm

LONG-TAILED
PYGMY POSSUM
Cercartetus caudatus
This arboreal
marsupial is found
in temperate rainforest
dense in New Guinea and
fur N.E. Queensland.

26–32 in
65–82 cm

4–5 in
10–13 cm
KOALA
Phascolarctos cinereus dull,
Feeding almost exclusively grayish brown
on eucalyptus leaves, upperparts
the koala lives in the forests
and woodlands of MOUNTAIN
E. Australia. It is solitary PYGMY POSSUM
and nocturnal. Burramys parvus
A ground-dweller of
rocky, upland habitats in
Australia, the mountain
pygmy possum hibernates
under snow for several
months during winter.

R I NGTA I L S
A N D R E L AT I V E S
This family consists of ringtail possums and the
lemurlike greater glider. The Pseudocheiridae
are all arboreal and are specialized
leaf-eaters. They have an enlarged
pouch at the start of the large
intestine for fermentation of
long, curved cellulose from their diet.
claws

WOM BATS COMMON WOMBAT

Vombatus ursinus COMMON RINGTAIL
Stocky marsupials with a short Found in forest, Pseudocheirus peregrinus
heathland, and coastal This agile marsupial
tail and short limbs, wombats have scrub in S.E. Australia, lives in a wide range of
large forepaws and long claws for this species is capable of habitats in E. Australia and 111/2–14 in
burrowing. The family Vombatidae digging tunnels up to Tasmania. It has become 29–35 cm
feed on coarse grass, which they 655 ft (200 m) long. a pest in New Zealand.
35–45 in
grind using strong jaws, and digest 90–115 cm
with an elongated gut. 33–43 in 14–22 in
84–111 cm 35–55 cm

silky fur mottled

brown and gray
LEMUROID RING-
SOUTHERN TAILED POSSUM
HAIRY-NOSED WOMBAT Hemibelideus lemuroides
Lasiorhinus latifrons Found only in a
An inhabitant of central small area of rainforest
southern Australia, this wombat in N.E. Queensland,
lives colonially in warrens Australia, this species
but feeds alone. of ringtail is nocturnal.
C U SC U S A N D R E L AT I V E S SULAWESI BEAR
CUSCUS large eyes assist
The family Phalangeridae includes cuscuses, brushtail possums, and their Ailurops ursinus sight at night 513
The largest of the cuscuses,
relatives. Most members are arboreal, with an opposable thumb on the this species inhabits the
hind limbs and a prehensile tail. In cuscuses, part or all of the tail is naked, canopy of temperate rainforest
whereas in brushtail possums the tail has fur. in Sulawesi and several other
Indonesian islands.

16½–29 in
42–74 cm
18½–22½ in
COMMON SPOTTED 47–57 cm
CUSCUS
Spilocuscus maculatus
This cuscus is sexually
dimorphic (males and
females look different)—
only the male has spots. It
lives in the rainforests
of New Guinea and
N.E. Australia.
19½–21½ in
49–54 cm

MOUNTAIN
strong, BRUSHTAIL POSSUM
curved claws Trichosurus cunninghami
The mountain brushtail
furry tail possum inhabits dense,
wet forests in S.E. Australia,
13–23½ in typically living at altitudes
33–60 cm above 985 ft (300 m).

CUSCUS
Phalanger sp.
12–18½ in Cuscuses of this genus
SCALY-TAILED POSSUM 30–47 cm live on New Guinea and
Wyulda squamicaudata nearby islands. They live
Found only in the Kimberley in N.W. Australia, at different altitudes
this nocturnal possum is solitary and bears to avoid competition
just one young at a time. with one another.

grizzled, GLIDING AND

gray fur
STRIPED POSSUMS
The Petauridae include striped possums and gliders,
13½–18 in
except the greater glider (left). Gliders have a thin,
34–45 cm furred membrane between their fore and hind
limbs. Striped possums emit strong smells and
GREATER have an elongated fourth finger to probe for
GLIDER wood-boring beetles.
Petauroides volans gray body with
The largest gliding dark stripe
marsupial, this species along its back
can travel distances 9½–11 in
of more than 330 ft 24–28 cm
6–6½ in
(100 m) between trees. 12½–16 in 15–17 cm
It lives in E. Australia. 32–40 cm

LEADBEATER’S
POSSUM
GREEN RINGTAIL Gymnobelideus leadbeateri
Pseudochirops archeri Found in moist,
Named for its thick, STRIPED POSSUM high-altitude forests
greenish fur, this solitary Dactylopsila trivirgata in Victoria, Australia,
possum is found only in Skunklike in appearance and odor, this possum feeds on
rainforests in the far north this possum is a nocturnal tree-dweller from insects, and sap and
of Queensland, Australia. N.E. Queensland, Australia, and New Guinea. gum from trees.
DAINTREE RIVER
RINGTAIL POSSUM 6–8½ in club-shaped tail
Pseudochirulus cinereus 15–21 cm
Resembling a lemur, prehensile tail thick tail SUGAR GLIDER
this ringtail possum lives in Petaurus breviceps
the montane tropical rainforest
in the Daintree River area in Partial to the sweet sap
N.E. Queensland, Australia. of eucalyptus trees, the
sugar glider is native
to N.E. Australia,
13½–14½ in New Guinea, and
34–37 cm neighboring islands.
 HONEY POSSUM F E AT H E R-TA I L E D
514 This diminutive species is the only GLIDERS
member of the family Tarsipedidae. The family Acrobatidae contains three
It has fewer teeth than other possums species—the feather-tailed or pygmy
MAMMALS • POUCHED MAMMALS

and a long, brush-covered tongue glider, the broad-toed feathertail glider,

for probing flowers. and the feather-tailed possum. All have
tails fringed with rows of stiff hairs.

HONEY POSSUM FEATHER-TAILED GLIDER

Tarsipes rostratus Acrobates pygmaeus
2–23/4 in
Inhabiting heathland and This species is the smallest gliding 5–7 cm
woodland in S.W. Australia, this marsupial. It lives in E. Australian
possum is a specialized feeder 21/2–31/2 in forests, where it feeds on nectar.
on nectar and pollen. 6.5–9 cm long, pointed snout

K A NGA ROOS
26–36 in
66–92 cm A N D R E L AT I V E S
The kangaroos and wallabies of the
family Macropodidae are medium to large
animals, with long hind legs for jumping. 21/4–41/2 ft
The hind feet lack a first toe, a fleshy 0.7–1.4 m
sheath encompasses the second and third
toes, forming a short grooming claw, 221/2–421/2 in
57–108 cm
and the strong, elongated fourth
and fifth toes are weight-bearing.

COMMON WALLAROO
red-brown
Osphranter robustus
RED KANGAROO Widespread throughout most of mainland
upperparts Osphranter rufus Australia, the common or hill wallaroo
The largest extant marsupial, often seeks shade in rocky outcrops.
this kangaroo is widespread in
Australia, where it lives in
savanna grassland and desert.

231/2–34 in
60–85 cm
18–21 in
45–53 cm

PARMA WALLABY AGILE WALLABY

Notamacropus parma Notamacropus agilis
Native to the mountains of the Great Unusually for wallabies, this species occurs
Dividing Range in Australia, the parma in both Australia and New Guinea, where it
wallaby inhabits a range of forest habitats. inhabits grassland and open woodland.

joey in pouch 21/4–71/4 ft

0.7–2.2 m

3–71/2 ft
0.9–2.3 m

EASTERN GRAY
RED-NECKED KANGAROO WESTERN GRAY KANGAROO
WALLABY Macropus giganteus Macropus fuliginosus
Notamacropus rufogriseus
Widespread in E. Australia, The only kangaroo that does not reproduce
This species lives in this species lives in dry woodland, using the method of delayed implantation of the
coastal forest and scrub scrub, and shrub. A subspecies embryo in the uterus, this species occurs in
in southeastern Australia, is found in Tasmania. southern Australia, including Kangaroo Island.
including the islands of
Tasmania and the Bass Strait.
long tail used
for support at
rest, for balance
when moving
POTOROOS R AT- K A N G A RO O
This family contains potoroos, bettongs, and rat kangaroos, small The musky rat-kangaroo is the sole member of the
515
marsupials with many similarities to the larger Macropodidae. family Hypsiprymnodontidae. It is relatively primitive;
However, unlike them, the Potoroidae as adults have a single unlike other kangaroo species, it has a first toe, which is

MAMMALS • POUCHED MAMMALS

serrated upper and lower premolar in each side of the jaws. 12–14 in opposable and helps it grip.
30–36 cm 6–11 in
10–16 in 15–28 cm
26–41 cm
MUSKY
RAT-KANGAROO
LONG-NOSED POTOROO BRUSH-TAILED BETTONG Hypsiprymnodon moschatus
Potorous tridactylus Bettongia penicillata This diurnal species
Inhabiting heath and forest This bettong lives in forest and inhabits tropical rainforest in
in S.E. Australia, this potoroo grassland in S.W. Australia. It has N. Queensland, Australia,
has strong, curved claws that help a prehensile tail that it uses for where it forages for fallen
it to dig for underground fungi. moving nesting material. fruits, seeds, and fungi.

191/2–27 in 131/2–43 in RED-NECKED PADEMELON 111/2–25 in

49–69 cm 34–109 cm Thylogale thetis 29–63 cm
A forest wallaby from eastern
Australia, this species ventures
to the forest edge at night to feed
on grass, leaves, and shoots.

muscular
thighs
NORTHERN NAIL-TAILED BRIDLED NAIL-TAIL WALLABY
WALLABY Onychogalea fraenata
Onychogalea unguifera This nocturnal wallaby was once 211/2–30 in 16–21 1/2 in
Also called the sandy nail-tailed wallaby, thought to be extinct; the only 54–77 cm 40–54 cm
this medium-sized kangaroo occurs remaining wild population is in a
across N. Australia. small area of Queensland, Australia.

12–151/2 in long, narrow

31–39 cm sole of foot

22–303/4 in
55–78 cm

coarse, RUFOUS HARE-WALLABY BROWN DORCOPSIS QUOKKA

dark fur Lagorchestes hirsutus Dorcopsis muelleri Setonix brachyurus
Previously inhabiting the This forest-dwelling Rare in mainland Australia,
Australian mainland, the mala or kangaroo is endemic to the this small marsupial is
rufous hare-wallaby now exists in low-altitude rainforests of found on Rottnest and
26–34 in the wild on only two islands in western New Guinea, and Bald Islands, off the
66–85 cm Western Australia. three offshore islands. southwestern coast. DORIA’S TREE
KANGAROO
reddish-brown Dendrolagus dorianus
upperparts The heaviest of the
SWAMP WALLABY 20–34 in GOODFELLOW’S tree-dwelling marsupials,
Wallabia bicolor 50–85 cm TREE KANGAROO Doria’s tree kangaroo
Darker in color than other wallabies, the Dendrolagus goodfellowi frequently descends to the
swamp wallaby lives in tropical and temperate Also called the ornate ground. It lives in montane
forests, and swampland in E. Australia. tree-kangaroo, this forest in New Guinea.
species is native to the
mountainous rainforests
of New Guinea, where it
feeds on leaves and fruit.
161/2–28 in
42–71 cm

20–24 in nonprehensile LUMHOLTZ’S

51–62 cm tail TREE KANGAROO
Dendrolagus lumholtzi
BRUSH-TAILED ROCK WALLABY The smallest of
Petrogale penicillata the tree-kangaroos,
This resident of S.E. Australia has this marsupial is found
roughened, padded hind feet that help it in the rainforests of
to grip when jumping in rocky habitats. N. Queensland, Australia.
 BIG BOOMER
Male red kangaroos, known as
boomers, are much larger than
females, sometimes weighing twice
as much. Only male red kangaroos
are red; the females, known as blue
fliers, have bluish-gray fur all over.
The males fight for access to mates,
their battles taking the form of
boxing matches.

velvety nose

glands in the neck

and chest produce a
dark, scented secretion,
which male kangaroos
rub on bushes to assert
their dominance

kangaroos lick
their forearms
when hot—this
cools the blood
close to the skin
elastic tendon
stores energy
when the leg
is flexed, and
releases it to
power the
next hop
R E D K A N G A RO O 517
Os ph rant er r uf u s
Kangaroos evolved in Australia, filling the niche occupied elsewhere by grazing animals such

MAMMALS • POUCHED MAMMALS

as antelope. In common with those species, kangaroos have a big stomach to take on large
quantities of grass. Living out in the open is risky for all herbivores, and kangaroos share with
antelope a number of adaptations for avoiding predators. They live in herds (called mobs), have
sharp senses, are tall enough to scan their surroundings for danger, and possess a wonderful turn
of speed. The red is the largest and swiftest kangaroo, able to hop at more
than 30 miles (50 km) an hour. Only the female has a pouch, in which she size 2¼–4½ft (0.7–1.4 m)
carries her young—known as a joey—for the first seven months of its life. habitat Scrubland, desert
Red kangaroos are well adapted to drought—they are able to eat scrubby distribution Australia
diet Herbivorous
saltbush, which is toxic to other animals.

EYE
The eyes are widely spaced toward
the sides of the head, giving a very
wide field of vision. This aids the
kangaroo’s awareness of predators.
STRONG FIGHTER
The male kangaroo has
a broad, muscular chest.
The forelimbs are used
to punch rivals in fights,
but the most forceful
blows come from
double kicks of
EARS the hind feet.
The ears are large and
sensitive, and can be
swiveled in different
directions to focus on
sounds that might mean
danger is approaching.

HIND FOOT
tail acts as a The scientific name
counterbalance Macropus literally
when hopping, and means “big foot.”
as a supportive Each hind foot has
fifth limb when CLAWS four toes: an outer,
standing still The feet have surprisingly weight-bearing pair,
sharp claws. These provide and an inner pair used
grip when hopping, but for grooming.
also serve as weapons, and
as combs for grooming.

HOPPING
Red kangaroos hop
on their enormous
hind legs with
apparently effortless
grace. They are able to
cover up to 30 ft (9 m)
in a single bound.
518 SENGIS
Formerly known as elephant shrews, Found only in Africa, sengis occupy a range of PHYLUM CHORDATA
sengis form a single family. They are habitats, from forests and savanna to extremely
MAMMALS • SENGIS

CLASS MAMMALIA

superficially similar to the true shrews, arid deserts. Some take shelter in rock crevices ORDER MACROSCELIDIDEA

although they are not related. or nests of dried leaves, while the larger species FAMILIES 1

dig out shallow burrows. SPECIES 20

Distinguished by an elongated, flexible snout, Sengis are primarily insectivorous, although

sengis are small, with long legs and tail. They some species take spiders and worms. They use D E B AT E
move either on all fours or, especially if they their sensitive nose to root on the ground and AN ORDER OF THEIR OWN
need to move fast, by hopping. Sengis live in under leaf litter for food, extending their long
pairs within a defined area, although there is tongue to scoop up their prey. In the past, sengis have been linked
little interaction between the partners. Males with shrews and hedgehogs, rabbits
and pikas, and even ungulates. Now
and females even sometimes make separate nests, ac t i v e yo u n g
in their own order Macroscelididea,
each defending the jointly held territory against Depending on species, a pair of sengis may breed genetic evidence suggests their
intruders of its own sex. Active mostly by day, several times during the year. Litters are small, closest relatives are tenrecs and
many sengis maintain a network of cleared paths numbering between one and three young, golden moles in the order Afrosoricida.
around their territory, which they patrol for prey which are well-developed at birth
and use as quick escape routes if threatened. and rapidly become active.

SENGIS hind legs

The Macroscelididae occur in a wide range of African habitats, longer than
forelegs
from deserts to mountains to forests. Largely insectivorous,
sengis (or elephant shrews) use their elongated, movable
snout to search for prey, which they flick into the mouth
with the help of their tongue.

41/2–43/4 in
11–12 cm BUSHVELD SENGI
Elephantulus intufi
Widespread in arid shrubland in southern
Africa, this sengi occupies large territories, which
long, it defends vigorously against rivals of the same sex.
furless tail
41/2–5 in 4–8 in 5–6 in
11–13 cm 10–20 cm 12.5–15 cm
4–43/4 in
10–12 cm

WESTERN ROCK SENGI RUFOUS SENGI DUSKY-FOOTED SENGI NORTH AFRICAN SENGI
Elephantulus rupestris Elephantulus rufescens Elephantulus fuscipes Petrosaltator rozeti
Inhabiting rocky shrubland Found in southern and eastern Africa, Frequenting hot, dry grassland in C. Africa, The only N. African elephant shrew,
in S.W. Africa, this species this sengi has long, sandy-reddish fur this poorly known species may actually this desert species slaps its very long
is active by day, leaving the and a tail as long as its head and body. belong to a different genus of sengis. tail and drums its feet when alarmed.
shade of a rock crevice
to foray for food. rounded ears
61/2–81/2 in 10–111/2 in 4–43/4 in
16–21 cm 25–29 cm 10–12 cm

ROUND-EARED SENGI
Macroscelides proboscideus
A medium-sized sengi, this
southern African species
occupies some of the world’s
driest habitats. Strictly
monogamous, males
FOUR-TOED SENGI BLACK AND RUFOUS SENGI strongly guard their mates.
Petrodromus tetradactylus Rhynchocyon petersi
Found in a range of moist habitats, This giant elephant shrew inhabits the
this species is one of the most coastal forests of E. Africa. Its orange
widespread elephant shrews. It foreparts graduate through deep red
occurs from C. to South Africa. to black on the rump.
TENRECS AND GOLDEN MOLES 519
Three families make up the order are also burrowing tenrecs. Until recently, tenrecs PHYLUM CHORDATA
Afrosoricidae—the golden moles that are and golden moles were classified with true moles,

MAMMALS • TENRECS AND GOLDEN MOLES

CLASS MAMMALIA
exclusively burrowers, the tenrecs and shrews, and hedgehogs because of their largely ORDER AFROSORICIDA

shrew tenrecs that have evolved into a insectivorous diet and often similar appearance. It FAMILIES 3

variety of habitats, and the otter shrews is now known that the order Afrosoricida evolved SPECIES 55

independently and is most closely related to the

that live in riverine and wetland areas. elephants, hyraxes, and sirenians.
Most species in this order are native to mainland
Africa, with tenrecs also found in Madagascar. u n u s u a l f e at u r e s
Golden moles are all very similar in appearance, with Tenrecs and golden moles have features once
a cylindrical body and other anatomical adaptations considered primitive but now recognized as
to suit their burrowing lifestyle. In contrast, tenrecs adaptations to harsh environments. Such features
display diverse characteristics that reflect the wide include a low metabolic rate and body temperature.
range of environments they occupy. Various species These mammals also have the ability to enter a state
of tenrecs are found in tropical forests, where they of torpor—for up to three days—to save energy
A Grant’s golden mole feasts on a
may be terrestrial or semiarboreal. The otter shrews in cold conditions, and possess highly efficient locust. At night it forages for food on
are aquatic, living in streams and rivers; and there kidneys, which reduce the need to drink water. the surface, feeding mainly on termites.

GOLDEN MOLES
These moles resemble both true moles (Talpidae) from Europe, Asia, and North 31/2–41/2 in
9–11 cm
America, and marsupial moles (Notoryctemorphia) from Australia, as all have similar
burrowing habits. The Chrysochloridae from southern Africa have short legs with
powerful digging claws, dense fur that repels moisture, and toughened skin, especially
on the head. They have nonfunctional eyes covered with skin, and lack external ears.

soft, dense,
glossy fur

JULIANA’S GOLDEN MOLE

Neamblysomus julianae
Unique to the dry highlands of South Africa,
typically in sandy soils, this species frequents
well-irrigated gardens within its range.

31/2–43/4 in
9–12 cm

CAPE GOLDEN MOLE

Chrysochloris asiatica
Although secretive, this is a common species
in parts of South Africa. It uses the enlarged
second toe on its front feet for digging.

leathery pad
protects nostrils
and aids burrowing

3–31/4 in
7.5–8.5 cm
webbed toes to
kick soil backward eyes covered with
thick layer of skin
4–51/2 in
10–14 cm
HOTTENTOT GRANT’S DESERT
GOLDEN MOLE GOLDEN MOLE
Amblysomus hottentotus Eremitalpa granti
This golden mole inhabits Inhabiting the southwest
tunnel systems up to 655 ft African coastal dunes, one of
(200 m) long, using the the world’s driest habitats, this
large second and third toes species “swims” through sand
on its front feet for digging. rather than building tunnels.
 TENRECS
520 From Africa and Madagascar, tenrecs have diverse body forms,
resembling the unrelated shrews, mice, hedgehogs, and otters. 4–7 in 7–14 in
They range in weight from 3/16 oz (5g) to more than 2¼ lb 10–18 cm 18–35 cm
MAMMALS • TENRECS AND GOLDEN MOLES

(1 kg). Largely nocturnal and with poor eyesight, these

insectivores use their sensitive whiskers to locate food.
LOWLAND STREAKED TENREC COMMON TENREC
LESSER white-tipped Hemicentetes semispinosus Tenrec ecaudatus
HEDGEHOG TENREC spines on back A tailless tenrec, the coarse, spiny A large terrestrial tenrec, with
Echinops telfairi coat of this species is distinctively red-brown fur and spines along the
Its hairs modified into two-tone—black, with yellow body, the females have up to 29 teats,
spines, this species bears stripes and crown bristles. more than any other mammal.
a remarkable resemblance
to a true hedgehog, sharing
its defense of rolling
into a ball.
moderately
large ears
4–61/2 in
10–17 cm

4–4 3/4 in
10–12 cm
RICE TENREC
Oryzorictes sp.
A burrower with well-developed
forelimbs, long claws, and small eyes
and ears, this tenrec can become
numerous in marshes and rice paddies.

51/2–9 in
14–23 cm
GREATER
HEDGEHOG TENREC
Setifer setosus
sharp claws Widespread throughout Madagascar, even in
urban habitats, this species has a diverse diet,
from insects, earthworms, and carrion to fruit.

A A R DVA R K
A A R D VA R K Inhabiting savanna and bushland in sub-Saharan
Africa, aardvarks are solitary and nocturnal.
AARDVARK
Orycteropus afer
The yellowish skin of the
Powerful front legs with flattened claws are used aardvark is often stained
The only species in the Orycteropodidae, the red by soil, in which it
to dig into the nests of ants and termites for food. burrows and forages. It
aardvark is adapted for digging. It has an arched also has a sparse covering
back, thick skin, long ears, and a tubular snout. of bristly hairs.
3 –4 1/2 ft
0.9–1.4 m
The aardvark’s toes PHYLUM CHORDATA hair lighter
(four on the front foot CLASS MAMMALIA colored
on body
and five on the hind) ORDER TUBULIDENTATA

each have a flattened nail FAMILIES 1

SPECIES 1
used to dig burrows and
excavate insect nests,
which it detects through its keen sense of smell.
It uses its long, thin tongue to catch huge numbers
of insects. Its teeth are very unusual, and are one of
the main reasons for it being classified in its own
order. The aardvark is born with incisors and canines at
the front of the jaw, but when these fall out they are not
long, blunt,
replaced. The rear teeth lack enamel, are open-rooted, shovel-shaped
and grow throughout life. claws
DUGONG AND HYRAXES 521
M A N AT E E S Furry and rotund, modern hyraxes belong to the
only extant family in the order Hyracoidea. Hyraxes

MAMMALS • HYRAXES
A small order of fully aquatic herbivores, sirenians were once the primary terrestrial herbivores in the
inhabit a range of tropical habitats, from swamps Old World. Now just five species remain.
and rivers to marine wetlands and coastal waters.
Widely represented in the PHYLUM CHORDATA
Also known as sea cows, PHYLUM CHORDATA
fossil record of Asia, Africa, CLASS MAMMALIA
sirenians are supremely CLASS MAMMALIA and Europe, some extinct ORDER HYRACOIDEA
adapted to an aquatic way ORDER SIRENIA species of hyrax were more FAMILIES 1

of life. Their front legs are FAMILIES 2 than three feet in height. SPECIES 5

paddlelike and adapted for SPECIES 4 Unlike most browsing and

steering; the hind legs are not grazing animals, hyraxes use their cheek teeth (rather than
visible, being restricted to two small remnant bones floating in the incisors) to slice vegetation, before chewing it. With a relatively
muscle. The body is therefore streamlined, and the flattened tail indigestible diet, they have a complex but efficient digestive
provides propulsion. A layer of blubber below their skin provides system in which bacteria break down tough plant matter.
insulation, but to help counteract its natural buoyancy, the bones With a number of primitive mammalian features, such as
are dense and the lungs and diaphragm extend for the full length poor control of their internal body temperature, hyraxes show
of the spine. The result is a hydrodynamic body, albeit rather several affinities with elephants: their upper incisor teeth are
slow-moving, capable of fine adjustments to shift its position in the often extended into short tusks, their soles have sensitive pads,
water. All four species are considered to be at risk of extinction. and they have high brain function.

DUGONG HYRAXES
There is only one species in the The Procaviidae are rotund, short-tailed 17 ¼–221/2 in long,
44–57 cm silky fur
Dugongidae. Slow-moving herbivores from Africa and the Middle
marine herbivores, dugongs East. Hyraxes are often seen basking in
graze on submerged the sun, and huddling together in
seagrass beds, using the groups for warmth. They
downturned muscular will also seek shade if
snout to uproot food. too hot. Behaving in
this way provides
8¼–10 ft hyraxes with an
2.5–3 m DUGONG
Dugong dugon effective means of
From the Indo-Pacific region, especially around thermoregulation.
Australia, the dugong’s cylindrical body has no
dorsal fin or hind limbs. The tail has flukes.
WESTERN TREE
HYRAX
Dendrohyrax dorsalis short
M A NAT E E S A usually dark, semi-
snout
arboreal hyrax from W. and
With a shorter snout than dugongs, the C. Africa, this species has
Trichechidae also differ in the shape of a distinctive white patch on
its lower back and rump.
their tail, which is paddle-shaped rather
than fluked. Rather sluggish, they spend 121/2–22 in
much of the day sleeping under water, 32–56 cm
surfacing every 20 minutes to breathe.
121/2–231/2 in
32–60 cm
AMAZONIAN
MANATEE 8–12 3/4 ft FLORIDA MANATEE BUSH HYRAX
Trichechus inunguis 2.5–3.9 m Trichechus manatus latirostris Heterohyrax brucei
A freshwater manatee The largest living sirenian, the There are 24 subspecies of
from the Amazon Basin, Florida manatee inhabits both bush hyrax in rocky habitats
this species usually has fresh water and coastal waters across Africa. It feeds on grass
a characteristic white of the S.E. US. It feeds on and fruit, and also catches and
chest patch. aquatic plants. 151/2–23 in eats small vertebrates.
39–58 cm

6 1/2–10 ft 8–12 3/4 ft SOUTHERN TREE ROCK HYRAX

2–3 m 2.5–3.9 m HYRAX Procavia capensis
ANTILLEAN MANATEE Dendrohyrax arboreus From Africa and the
Trichechus manatus manatus An able climber from Middle East, this
Ranging from Mexico to southern Africa, usually colonial hyrax basks for
Brazil, this manatee is smaller inhabiting hollow trees, long periods. Moist,
and more likely to be found this hyrax is known for rubbery foot pads
farther from the shore than the its shrieking territorial provide grip on smooth
Florida subspecies. calls at night. rocky surfaces.
 BRAIN POWER
Long acknowledged for having a good
memory, the elephant is now known
to solve problems, use tools, show
emotions, and work with others,
making it one of the most intelligent
of all terrestrial animals.

elongated nose and

upper lip form the
mobile trunk
 A F R I C A N S AV A N N A 523
ELEPHANT
L oxodont a af r icana
A mature male African savanna elephant is larger than any other living land mammal and
in exceptional cases can weigh as much as eleven tons. The head alone can exceed 880 lb
(400 kg), including the brain, eyes, trunk, ears, tusks, and teeth. Females are smaller and
form strongly bonded herds, comprising a dominant female, close relatives, and their
offspring. Bull elephants form less cohesive groups led by older males, but in the breeding
season, when they come into musth, they are more solitary and will fight for access to
receptive females. Elephants are long-lived and may survive well into their 60s.

EYE TRUNK SKIN

Long eyelashes protect the small, The trunk has short sensory
laterally-positioned eyes, which can see hairs along its length that
clearly for only about 33 ft (10 m). Other enhance its responsiveness
senses compensate for this limitation. to touch.

FINGERLIKE TIP
The opposing fingerlike
projections at the tip of the
trunk are extremely sensitive
EAR and dextrous. This allows
The large, thin- the African savanna elephant
skinned ears help to pick up objects as small
to regulate body as a peanut.
temperature. They
are also used to
communicate moods,
such as aggression.
TAIL
Elephants swing their tail
to deter insects. The air
FOOT turbulence created prevents
Elephant feet have sensory them from landing, and any
receptors along their margins. that do are swatted.
They pick up low-frequency
underground sounds that can musth glands are only
travel over long distances. found in elephants and
are present in both sexes

the tusks are

enlarged second
incisor teeth

size 81/2–13 ft (2.6–4 m) LARGE AND HEAVY

habitat Dry woodland, A bulky body supported by
scrub, and savanna grassland straight, columnar legs might
distribution Mainly E. and suggest the elephant is a large abdominal
S. Africa, south of the Sahara slow-moving animal. It can, cavity contains
diet Leaves, roots, bark,
however, reach speeds of about 62 ft (19 m)
fruits, and seeds
15mph (24kph) when charging. of intestines
524 ELEPHANTS
Largest of all land mammals, elephants draw in water and serve as a snorkel when PHYLUM CHORDATA
are distinguished by their colossal swimming. Sensory organs in the trunk pick up
MAMMALS • ELEPHANTS

CLASS MAMMALIA

bulk, long flexible trunk, large ears, scents and vibrations, so aiding communication. ORDER PROBOSCIDEA

and curving ivory tusks. An elephant’s large ears are also used for FAMILIES 1

communication and when spread out are a sign of SPECIES 3

The only members of the Proboscidea, elephants aggression. However, constant flapping helps an
inhabit grassland and forest in tropical Africa and elephant to lose heat. Tusks are incisor teeth that D E B AT E
Asia. Sometimes weighing more than five tons, grow continually. They are used to dig for roots TWO SPECIES OR ONE?
elephants have a modified skeleton to bear their and salt, clear pathways, and mark territories.
weight. The limb bones are extremely stout, and The splitting of the African elephant
the toes are splayed around a pad of connective s o c i a l s t ru c t u r e s into two species, the savanna elephant
tissue. This large frame needs vast amounts of Male and female elephants have different social and the smaller forest elephant,
was originally based on physical
food and elephants eat for up to 16 hours a day, behavior. Females form herds of related cows
differences, but now there is sufficient
consuming up to 550 lb (250 kg) of vegetation. and their calves, led by an elderly matriarch. genetic evidence to substantiate this
Bull elephants form short-term associations, division. However, some scientists are
ea rs , nose , an d mou t h but will fight with reluctant to accept this idea because
The elephant’s muscular trunk, a fusion of the any intruding the two species sometimes interbreed
nose and upper lip, is remarkably versatile and rival during the where their ranges overlap.
is able to grasp morsels of food. It can also breeding season.

ELEPHANTS 73/4–11 ft
2.4–3.4 m
Including elephants and the extinct mammoths, the
Elephantidae form a family of large herbivores, with
a long trunk, large ears, thick skin, and tusks. The 81/2–13 ft
large ears 2.6–4 m
trunk is used in feeding, drinking, bathing, and
in social interactions. Elephants make a range
of sounds, from trumpeting to subsonic
rumblings, enabling them
to communicate over
long distances.

long, curved
tusks

ASIATIC ELEPHANT
Elephas maximus AFRICAN SAVANNA ELEPHANT
Domesticated for forest operations and Loxodonta africana
ceremonial use, this species has smaller ears The largest living land animal, this
and a more arched back than African species. elephant has a large head and ears. Both
Females usually lack tusks as do some males. sexes have well-developed, curved tusks.

51/4–91/2 ft
1.6–2.9 m

muscular trunk
projections

AFRICAN FOREST
ELEPHANT
Loxodonta cyclotis
With five toenails
on the forelimbs
and four on the hind
limbs, this small elephant
semicircular with straight tusks is
toenails from the tropical forests
of Central Africa.
ARM ADILLOS 525
Recognizable by their armored shell— Armadillos have few natural predators. Despite PHYLUM CHORDATA
unique among mammals—armadillos the threats from human hunters and habitat loss,

MAMMALS • ARMADILLOS
CLASS MAMMALIA

occur in various shapes, sizes, and the range of some armadillo species is expanding. ORDER CINGULATA

colors. All are native to the Americas. Although their armor is heavy, armadillos FAMILIES 2

are effective swimmers. By inflating the stomach SPECIES 20

Armadillos, sole members of the order Cingulata, and intestines with air, they can increase their
range across a variety of habitats, feeding mostly buoyancy and cross small bodies of water.
on insects and other invertebrates. Despite having Armadillos can hold their breath under water
short legs, armadillos can run quickly and burrow for several minutes, allowing them to walk
with their strong claws to avoid predators. Their over a stream bed rather than swim across it.
main defense consists of a bony carapace, covered
with horny plates, across their upperparts. In most habits
species, there are rigid shields over the shoulders Most species of armadillo are nocturnal, though
and hips, with a varying number of bands they occasionally emerge during the day as well.
separated by flexible skin covering the back and These animals are largely solitary, associating with
Contrary to popular belief, not all
flanks. This allows some species to roll into a ball one another only during the mating season. Males armadillos can roll into a ball in defense—
to protect their vulnerable, furry underparts. sometimes display aggression toward rivals. only species of the genus Tolypeutes can.

ARMADILLOS 91/2–221/2 in
24–57 cm
LONG-NOSED
ARMADILLO
Dasypus sp.
The only surviving family in their order, the The seven species of this
Dasypodidae are found in the Americas. Covered genus inhabit shaded,
with bony plates across their upperparts, they use rocky areas. Unlike other
their sharp claws to dig for invertebrates and to armadillos, they have
sparse, yellowish fur,
excavate burrows. Of the 20 or so species, mainly on their bellies.
some are able to roll into a ball when
threatened, in order to protect
their vulnerable, soft, long, pointed
furry underparts. snout used
long tail in for foraging
proportion to body

8–12 in 10–16 in
20–30 cm 26–40 cm
well-developed
9–12 in claws
22–31 cm

PICHI
Zaedyus pichiy
A small, dark armadillo
ANDEAN HAIRY ARMADILLO LARGE HAIRY ARMADILLO with thick dorsal plates,
Chaetophractus vellerosus Chaetophractus villosus this species wedges itself
Unusual in having copious hair between its scales, From arid habitats in southern S. America, this into its burrow when
this mammal inhabits high-altitude S. American species has long, coarse hairs between the 18 or threatened, presenting its
grasslands. It is hunted for its meat and shell. so plates of protective armor across its back. jagged scales for defense.

PINK FAIRY ARMADILLO

16–20 in 41/2–6 in Chlamyphorus truncatus 21/4–31/4 ft
40–50 cm 11–15 cm This tiny armadillo 0.75–1 m
from C. Argentina
is mainly subterranean,
“swimming” through loose
sand; its head is shielded
SIX-BANDED ARMADILLO to reduce abrasion.
Euphractus sexcinctus 12–15 in
More active by day than most armadillos, this 30–38 cm
brownish-yellow species forages in grassland and rounded,
forest, eating plant and animal matter. pink ears

NORTHERN NAKED-TAILED GIANT ARMADILLO

ARMADILLO Priodontes maximus
jointed bands on Cabassous centralis The largest armadillo, this
blackish upperparts The protective plates of this species has tough armor
C. and S. American armadillo do not and uses its long, curved
extend to its tail. It relies mainly on third front claw to dig
dense cover to avoid predators. for food and in defense.
526 S I X- B A N D E D
ARMADILLO
Euph rac tu s sexcinc tu s
MAMMALS • ARMADILLOS

Despite its name, this species of armadillo may have anything from six to
eight armored bands around its middle. The upper part of the animal’s body
is covered by a sturdy carapace. The carapace is made up of plates of bone
topped with a thin covering of horny material, and it is embedded
in the armadillo’s skin. The bands act as joints in the carapace, giving the
animal flexibility, although it is unable to curl completely into a ball like size 16–20 in (40–50 cm)
some other species. The six-banded armadillo is diurnal, spending the habitat Forest and savanna
daylight hours trundling about its home range in search of food. It has distribution S. America,
a varied diet, ranging from roots and shoots to invertebrates and carrion. mainly south of Amazon Basin
diet Omnivorous

SHIELDED EYE
The armadillo’s eyes are protected by
the headshield at the expense of a slightly
restricted field of vision. As it has poor
eyesight anyway, this makes little difference.

NOSE
Armadillos have an acute sense of
smell. They sniff and snuffle their
way to most meals, and can easily
detect food buried in the soil.

head is narrow and

pointed and protected by
a shield made of fused
bony plates

STRONG LEGS
The armadillo has
short but powerful
legs. When digging,
soil loosened by the
front feet is kicked
out of the hole by
the hind feet.

HAIRY SKIN
The bony carapace is
interrupted by six to
eight bands, which are
separated by flexible
skin. Long, bristly hairs
sprout from between
the bands – this is one
of the hairy armadillos.

TAIL CLAWS
Glands at the base of the Long, strong claws allow the
armadillo’s tail release armadillo to dig rapidly into
scent through small hard ground. In a matter of
holes in the armor minutes it is able to excavate
plates. The scent marks a trench deep enough to
the armadillo’s territory. bury itself in.
 ARMORED DIGGER parts of the body not
The distinctive carapace offers some defense against covered in bony plates
predators, although armadillos prefer to run away if sprout a sparse coat
possible. When digging, the carapace protects the of long hairs
skin from abrasion. Armadillos dig to find food
(both plant and animal matter), and to make
burrows in which they live or
briefly hide themselves.
528 S L O T H S A N D A N T E AT E R S
Although very different in form and grind up their food. A sloth may take about a PHYLUM CHORDATA
habits, sloths and anteaters have one month to completely digest a meal, as the fibrous
M A MM ALS • SLOTHS AND ANTEATERS

CLASS MAMMALIA

characteristic in common: they share leaf matter passes slowly through several stomach ORDER PILOSA

a lack of normal mammalian dentition. compartments and is broken down by bacteria. FAMILIES 4
SPECIES 16

With just one exception, members of this order, arboreal life

the Pilosa, are largely arboreal creatures: only the The long, prehensile tail of the arboreal anteaters
giant anteater is terrestrial. They are all native to allows them to lead a more active lifestyle than
Central and South America. the sloths. On the ground, all the arboreal species
The two anteater families feed on ants, move only with difficulty, although some are
termites, and other insects. Anteaters have no excellent swimmers. When walking, their
teeth at all, relying on their long, sticky tongue mobility is restricted by their enlarged, curved
to capture insects, which are then crushed in the claws, which the sloths use like hooks to hang
mouth before being swallowed. from branches. Anteaters have enlarged claws
The slow-moving sloths are herbivorous. They only on their front feet. These claws are used
To defend themselves members of the
have no incisors or canines, but instead have a to tear open insect nests in search of food, and genus Tamandua stand up on their hind legs
number of cylindrical, rootless teeth, used to are formidable defense weapons. and lash out with their powerful front limbs.

T H R E E -TO E D S L O T H S T WO -TO E D S L O T H S 211/2–35 in

54–88 cm
Generally smaller and slower moving than two-toed Unlike three-toed sloths, the Megalonychidae
sloths, the arboreal and mostly nocturnal Bradypodidae 23–28 in have only two toes on their forefeet, a more
have three toes on each foot, bearing 59–72 cm prominent snout, and no tail. Similarly arboreal
long, curved claws that help them to and mainly nocturnal, they usually descend
hang from branches. Their long, from trees head first.
shaggy fur has a green tinge, two toes on
front feet
due to algae growing on it.

MANED SLOTH
Bradypus torquatus
This small Brazilian sloth has long,
dark fur—especially around its head
and neck—which often harbors
algae, ticks, and moths.

BROWN-THROATED SLOTH
Bradypus variegatus
An inhabitant of the forests of
C. and S. America, this sloth is the most LINNAEUS’S
TWO-TOED SLOTH
widespread in its family. The female Choloepus hoffmanni
attracts her mate with a shrill scream. This solitary, S. American
211/2–35 in herbivore is found in the tropical
54–88 cm forests of the Orinoco and
201/2–211/2 in
52–54 cm Amazon river basins, where it
SOUTHERN spends most of its time sleeping
TWO-TOED SLOTH or resting in the forest canopy.
Choloepus didactylus
18–30 in
45–76 cm This large herbivore from S. America swims
well, even crossing rivers. Its main predators
are large raptors, such as harpy eagles.

SI L KY A N T E AT E R S
The Cyclopedidae, though well-represented in the fossil
record, has seven extant species. Large, curved front claws
shaggy, and a prehensile tail help silky anteaters live on trees,
coarse coat
where they nest in holes. They feed on ants and termites.
PALE-THROATED SLOTH
Bradypus tridactylus SILKY ANTEATER
Found in the rainforests of Cyclopes sp.
S. America, this species usually Arboreal, nocturnal, and slow moving, silky anteaters live in
lives alone and may rest for more forested areas of Central and South America. Their claws 71/2–9 in
than 18 hours a day. provide an effective defense when threatened by predators. 19–22 cm
GI A N T A N T E AT E R A N D R E L AT I V E S 181/2–30 in
47–77 cm
Native to Central and South America, the Myrmecophagidae
529
have an elongated snout and a long tongue. They use their
powerful claws to rip open termite mounds

MAMMALS • RABBITS, HARES, AND PIKAS

and ant hills. Lacking teeth, they catch
insects with their sticky saliva and SOUTHERN TAMANDUA
spine-covered tongue. Tamandua tetradactyla
This solitary species has a
prehensile tail. It forages for food at
any time of day, and is active for
huge, about 7 hours in every 24.
bushy tail

stiff, strawlike long, tubular

hair snout

31/4–41/2 ft
1–1.4 m

GIANT ANTEATER
Myrmecophaga tridactyla
The largest anteater, with a tail almost as long
as its body, this species uses its long, sticky
tongue to catch up to 30,000 insects a day.

R A BBITS, H A RES, AND PIK AS

Two herbivorous families make up the system. They produce two types of feces: moist PHYLUM CHORDATA
order known as Lagomorpha. Their pellets, which are eaten to gain further nutrients CLASS MAMMALIA

apparent similarities to rodents result from the food, and dry pellets passed as waste. ORDER LAGOMORPHA
FAMILIES 2
from adaptation to a common lifestyle. SPECIES OVER 90
e s c a p i n g p r e dat o r s
These species occupy a variety of habitats, from Pikas, the most rodentlike species in this order,
tropical forest to Arctic tundra. All are terrestrial take refuge from predators in burrows and crevices
herbivores or browsers. They are gnawing animals after raising the alarm with a whistle. Rabbits and
and share the diet of many rodents. Like those hares, in contrast, have long ears to detect danger
of rodents, the teeth of rabbits, hares, and pikas and powerful limbs to flee from predators. These
continue to grow throughout life—they are animals have large eyes positioned high on each
worn down by chewing. However, there are side of the head to provide almost 360-degree
fundamental differences between the groups: vision. When a predator is observed,
rabbits and their relatives have four incisors in hares drum their hind legs on the
the upper jaw, while rodents have only two. ground in warning.
To survive the winter, the pika collects
Relying on food that is relatively slow to break various plants, creates a hay pile of dried
down, these animals have a modified digestive food, and stores it in its den.

RABBITS AND HARES EUROPEAN RABBIT

Oryctolagus cuniculus
Native across much of the world, the Leporidae Native to the Iberian penninsula, this
species has been introduced for meat
have long, movable ears and enlarged hind legs and fur worldwide, with devastating
to detect and escape predators. Large eyes reflect effects on local habitats and wildlife.
their mainly nocturnal habits. Rabbits often inhabit
permanent burrow systems, whereas the more
14–15 in 6–12 in
solitary hares make only transitory shelters. 36–38 cm 15–30 cm

furry feet
5–7 in
13–18 cm LOP-EARED
10–15 in RABBIT
DWARF RABBIT 25–38 cm ANGORA RABBIT Oryctolagus cuniculus
Oryctolagus cuniculus Oryctolagus cuniculus First bred in England in
One of the smallest breeds, Valued for its long, the 19th century,
the dwarf rabbit occurs in soft hair, which is spun into this type has long,
many colors and patterns. Its yarn, this breed originates lop-eared (floppy) ears.
rounded face and small ears from Anatolia There is variation
make it a popular pet. (in modern-day Turkey). in both color and size.
530
M A MM ALS • SLOTHS AND ANTEATERS

hairs of the coat are

long, coarse, and
grooved and grow
away from the belly

strong, elongated
fore- and hind limbs are
about the same length

TOPSY-TURVY WORLD
Adapted to living upside down in the
forest canopy, Linnaeus’s two-toed
sloth rarely ventures to the forest
floor other than to relieve itself every
3–5 days. Strong fingers with
clawlike hooks for grasping branches
allow it to eat, mate, give birth, and
even sleep in this position.
 LINNAEUS’S 531
T WO -T O E D S L O T H
Choloepu s hof fman n i

M A MM ALS • SLOTHS AND ANTEATERS

Small and arboreal, Linnaeus’s two-toed sloth is far removed from its giant ground-
dwelling ancestors, which only became extinct about 10,000 years ago. It is one of
the slowest-moving mammals and sleeps for about 13 hours a day, often curled into a
ball in the fork of a tree, waking at night to forage for leaves. The sloth’s fur is unusual
in supporting an ecosystem that includes unique microorganisms, green algae, and
various insects. In damp conditions, the coat may acquire a green hue due to the algae
growing in the grooves of the hairs.
size 211/2–35 in (54–88 cm)
habitat Tropical lowland forest
distribution Orinoco and
EYE TEETH
Amazon basins, S. America The eyes are The five upper and four
diet Mainly leaves large and forward lower teeth are peglike,
facing, but the lack enamel, and grow
eyesight is poor. throughout life. There
Sloths are are no incisors.
nearsighted,
color blind,
and have poor
visual acuity.

NOSE EAR
A short snout with a large, The sloth’s hearing is
snub nose at its tip provides poor and limited to lower
the sloth with an excellent frequencies. The external
sense of smell. part of the ear is tiny and
buried in the fur.

CLAWS
The forelimb terminates in two
digits, each with a 31⁄4–4 in
(8–10 cm), hooklike claw. The
hind limb has three clawed digits.

MIDDLE PARTING
Due to their upside-down lifestyle, the
fur parts in the midline of the chest and
abdomen and falls down toward the
back, so rain will run off.
 RABBITS brown fur, sometimes
532 AND HARES turns reddish long, black-tipped ears

20–28 in
MAMMALS • RABBITS, HARES, AND PIKAS

141/2–16 in 50–70 cm
37–40 cm

DESERT COTTONTAIL
Sylvilagus audubonii
Found in arid regions of the S.W.
US and N. and C. Mexico, the
desert cottontail nests above
ground rather than in a burrow.

EUROPEAN HARE
Lepus europaeus
A grazer in summer and a browser of bark and
buds in winter, this hare is shy and solitary, except
during its springtime “boxing” courtship. Females
fight off males until ready to mate.

MARSH RABBIT 161/2–17 1/2 in

Sylvilagus palustris 42–44 cm
Inhabiting wetlands of
N. America, this species is a strong
swimmer. It walks instead of hopping, SNOWSHOE HARE
unlike most of its relatives. powerful hind legs 14–201/2 in Lepus americanus 22–28 in
36–52 cm Adapted to harsh N. American 55–70 cm
winters, this species has a white
22–26 in winter coat for camouflage,
56–66 cm
and large hind feet to allow
movement on soft snow.
WHITE-TAILED ARCTIC HARE
JACKRABBIT Lepus arcticus
Lepus townsendii Adapted to polar and
This species ranges extensively mountainous areas,
across western N. America. The this hare survives with
northern population turns white thick fur that turns
in winter, while the southern white in winter, and by
animals develop whitish patches digging snow holes
only on their flanks. for shelter.

201/2–24 in 22–26 in
52–61 cm 55–67 cm

BLACK-TAILED 20–22 in 18–22 in

JACKRABBIT 51–55 cm 45–55 cm
Lepus californicus
Widespread on western
N. American prairies and
farmland, this species is prone CAPE HARE
to huge local fluctuations. It has ANTELOPE JACKRABBIT MOUNTAIN HARE Lepus capensis
a black tail and black ear-tips. Lepus alleni Lepus timidus Common in open habitats
Very long ears and insulated, From polar regions and mountains of Africa and the Middle
reflective fur help this large in Europe and Asia, this hare has East, this species is very
hare to keep cool in its desert a white winter coat and a tail similar to the closely
grassland habitats in Mexico. that stays white all year round. related European hare.

PIKAS
61/2–81/2 in
The Ochotonidae are small 16–21 cm
herbivores that live in rocky
long mountainsides and open
legs steppes in North America and AMERICAN PIKA
Asia. Alert to predators, they Ochotona princeps
produce high-pitched alarm This N. American
montane scree dweller
calls as they run for cover dries food piles in
in crevices and burrows. the sun, and stores
them in its den for
the winter.
RO D E N T S 533
Ranging from tiny mice to animals t h e i m pac t o f ro d e n t s PHYLUM CHORDATA
the size of a pig, rodents live in almost Several rodent species carry fatal diseases that

MAMMALS • RODENTS
CLASS MAMMALIA

every habitat type. They make up have killed millions of people. Others consume ORDER RODENTIA

nearly half of all mammal species. or contaminate huge quantities of stored food FAMILIES 34
SPECIES About 2,500
meant for humans. The house mouse has become
Members of the order Rodentia are distinguished the world’s most widespread wild mammal
by a pair of prominent upper and lower incisor through its close association with humans. D E B AT E
teeth (often orange or yellow in color), which This species inhabits every continent, except R O D E N T O R G A N I Z AT I O N
continue to grow throughout life. Gnawing— a Antarctica, even surviving in mines and cold
characteristic behavior of all rodents—wears the stores. Some rodents cause damage to crops The huge diversity of species in the
teeth away at the same rate as they grow. Rodents order Rodentia makes the task of
or trees, or burrow in inconvenient places.
classifying them rather complicated.
have no canine teeth, just a long space between Beavers can transform entire habitats, affecting With 34 families of modern rodent
the incisors and the three or four cheek teeth in hundreds of other animal and plant species. to organize, experts in zoological
each jaw. The scientific classification of rodents is On the positive side, rodents are a vital food classification have divided them into
also based on other features of their teeth and source for predatory animals and, in some two subgroups, according to differences
jaws that are not visible externally. countries, for humans as well. Many small rodents, in their skulls, teeth, and jaws. The
such as hamsters, are specially bred as pets. Sciurognathi (squirrel-jaws) are globally
speci es di v ersi t y widespread, and include all squirrels,
To suit their various lifestyles, rodents often have beavers, and mouselike rodents. The
special adaptations such as webbed toes, big ears, Hystricognathi (porcupine-jaws) includes
the cavies, porcupines, chinchillas,
or long hind feet for a bounding gait. Some
and the capybara, and these are
rodents burrow, others live in trees or in the mostly restricted to the southern
water, but none live in the sea. Many species hemisphere and the tropics.
inhabit desert regions, where they may never
drink, deriving all the water they need from food.

MOU N TA I N SQUIRRELS AND

B E AV E R CHIPMU NKS
9–12 in
Only one species of the once Occurring almost everywhere, 23–30 cm
widespread family Aplodontiidae except in the polar regions, 8–10 in
20–26 cm
has survived into modern times. Australia, and the Sahara Desert,
The mountain beaver, or members of the family Sciuridae
sewellel, lives in burrows in range from tropical rainforests to GRAY SQUIRREL EURASIAN RED
the damp forests and plantations Arctic tundra, and from the tops of Sciurus carolinensis SQUIRREL
of western North America. trees to underground tunnels. This Common in the E. US, Sciurus vulgaris
family includes typical bushy-tailed this species was introduced to When this squirrel molts into
parts of Europe, where it is slowly its summer coat, the long
flattened head
tree squirrels, many species of displacing the native red squirrel. hairs on its ears are lost.
12–16 in
30–40 cm burrow-digging ground squirrels,
chipmunks, and marmots. They gliding
SOUTHERN FLYING membrane
feed mainly on nuts and seeds. SQUIRREL
Glaucomys volans
AMERICAN RED A strictly nocturnal
SQUIRREL native of the eastern
Tamiasciurus hudsonicus US, this squirrel lives
The chattering calls in tree holes and attics,
and sharp barks of this 61/2–8 in often in groups
red squirrel are common 17–20 cm during winter.
sounds of coniferous
woodlands in Canada
and the N. US. 41/2–51/2 in
11–14 cm

121/2–151/2 in 8–10 in
32–39 cm 20–26 cm

PREVOST’S
SQUIRREL
MOUNTAIN BEAVER Callosciurus prevostii
Aplodontia rufa PALE GIANT SQUIRREL GRIZZLED GIANT SQUIRREL There are 17 subspecies
12–15 in Ratufa affinis Ratufa macroura
This most primitive 30–38 cm of these tricolored
type of living rodent Of the four species of giant One of the world’s largest tree squirrels. Prevost’s is
inhabits forested coastal squirrel, this species inhabits the squirrels, this species occurs in S. India found in Malaysia, Borneo,
mountains of W. Canada forests of the Malay Peninsula, and Sri Lanka. It eats fruits, Sumatra, and nearby
and the US. Borneo, and Sumatra. flowers, and insects. islands, such as Sulawesi.
 SQUIRRELS CAPE GROUND
SQUIRREL
COLUMBIAN
GROUND SQUIRREL
534 AND CHIPMUNKS Geosciurus inauris Urocitellus columbianus
This coarsely haired squirrel A rather large, bushy tailed ground
escapes the extreme temperatures squirrel, this species forms colonies in
of the semideserts of South Africa meadows and forest edges from Idaho
MAMMALS • RODENTS

by sheltering in burrows. north into W. Canada.

long,
bushy tail
6–8 in
15–20 cm

GOLDEN-MANTLED
GROUND SQUIRREL
Callospermophilus lateralis
Appearing like a larger version of the
chipmunk, this species is a common 10–12 in
sight in forests and mountains 25–30 cm
of the W. US.
short,
9–11 in 43/4–6 in 43/4–6 in hairy tail
22–28 cm 12–15 cm 12–15 cm

61/2–101/2 in
17–27 cm
GAMBIAN SUN
SQUIRREL
Heliosciurus gambianus HOPI CHIPMUNK EASTERN CHIPMUNK
A common species in savanna Tamias rufus Tamias striatus
woodlands across Africa, from Chipmunks from different parts of This striped, ground-living
Senegal to Zimbabwe, this N. America form separate species. species is a common sight in forest
squirrel feeds mainly on the This species occurs in Utah, Colorado, campgrounds of the E. US, where it
seeds of Acacia trees. and adjacent parts of Arizona. has become quite tame.

B E AV E R S DORMICE 23/4–6 in
There are only two species of beavers in the family Castoridae. The Gliridae occur in wooded habitats across 7–15 cm
Both are exploited for their fur. One is widespread across Europe, sub-Saharan Africa, and patchily
North America, and the other in patches throughout most in Central Asia, with a single species in
of Europe. Both build dams out of stones, mud, and trees, Japan. Resembling small nocturnal
creating habitats for other species. squirrels, all but one are soft-
glossy, furred, arboreal species. Many
brown fur
BEAVER are threatened or are declining
Castor sp. in numbers.
European and
N. American beavers
are different species, AFRICAN
but both live a similar, DORMOUSE
semiaquatic life in Graphiurus sp.
rivers and lakes. There are 15 species
of African dormice.
They are all similar in
21/2–4 ft appearance and inhabit
0.8–1.2 m forested areas of
sub-Saharan Africa.
flat, scaly tail
bushy tail

GOPHERS
Members of the family Geomyidae, found in North
America, live alone in shallow burrows that allow them
to access roots and leaves. They carry their food in
cheek pouches and store it in underground chambers.
31/4–8 in BOTTA’S POCKET GOPHER
8–20 cm Thomomys bottae
A small burrowing creature of soft
soil and grassy areas, this species digs
up mounds of earth, which can
damage farm machinery.
 POCKET MICE AND
14–20 in K A NGA ROO R AT S 535
35–50 cm
Mostly common, with a few rare species, the Heteromyidae are found in
varied habitats from Canada to Central America, with kangaroo rats living 4 in

MAMMALS • RODENTS
gingery brown fur mainly in deserts. Pocket mice typically run on four feet, but kangaroo 10 cm
rats hop on two large hind feet.
MARMOT longer hairs
Marmota sp. of tail tuft
Several species of marmots
inhabit mountain grasslands and
rocky areas of N. America, while
some occur in Eurasia. Marmots
hibernate for up to eight months, DESERT
POCKET MOUSE MERRIAM’S KANGAROO RAT
depending on the species. 23/4–31/2 in Chaetodipus penicillatus Dipodomys merriami
7–9 cm This species is one of many small nocturnal Bounding away with its tail outstretched,
rodents that inhabit the open, sandy deserts this nocturnal inhabitant of the N. American
121/2–16 in of the S.W. US and N. Mexico. deserts looks like a miniature kangaroo.
32–40 cm
BLACK-TAILED
PRAIRIE DOG
Cynomys ludovicianus JUMPING MICE MEADOW JUMPING MOUSE
Zapus hudsonius
Active during daytime,
this ground squirrel lives The families Dipodidae (jerboas), Zapodidae Found in the cool grasslands of
in large groups in “towns” (jumping mice), and Sminthidae (birch mice) northern N. America, this species
consisting of extensive 23/4–41/4 in also has isolated populations in the
communal burrows.
are made up of rodents that have a long tail and 7–11 cm mountains of Arizona and New Mexico.
powerful hind feet that enable them to jump
81/2–101/2 in around like miniature kangaroos.
21–27 cm
HARRIS’S ANTELOPE
SQUIRREL
Ammospermophilus harrisii
An agile inhabitant of the LESSER EGYPTIAN JERBOA
Sonora Desert and northern Jaculus jaculus
Mexico, this rodent is active This desert species is found across
in the heat of the day, but N. Africa, from Senegal to Egypt, 41/2–5 in
hibernates during winter. south to Somalia, and east to Iran. 11–13 cm

VOL E S , L E M M I NGS , russet fur

A N D M U SK R ATS on back

About 765 chubby, short-tailed rodents, including

hamsters, voles, lemmings, and muskrats, make up
the family Cricetidae. They are found worldwide,
EDIBLE DORMOUSE 5–71/2 in
Glis glis 13–19 cm from western Europe to Siberia and the Pacific coast. 31/4–51/2 in
Known in Germany as “the seven-sleeper,” 8–14 cm
this dormouse hibernates for more than
six months. It is eaten in Mediterranean
countries, such as Slovenia.
31/2–43/4 in
9–12 cm

short BANK VOLE

ears Myodes glareolus
43/4–9 in Active mostly at dawn and
12–23 cm night, this typical vole inhabits
scrub, woodlands, and gardens
COMMON VOLE across most of W. Europe
Microtus arvalis EURASIAN WATER VOLE and east into Russia.
A common burrowing inhabitant Arvicola amphibius
of grasslands across most of In Britain and parts of Europe, this vole occurs near
N. Europe, this vole also occurs waterside habitats, but in Russia and Iran, it lives away
farther east and in Russia. from water, digging extensive burrow systems.
10–12 in
25–30 cm
COMMON DORMOUSE 41/2–6 in 31/4–43/4 in
Muscardinus avellanarius 11–15 cm 8–12 cm
An excellent climber and
jumper, this dormouse feeds on
flowers, fruits, and insects
at night, in shrubby woodland
across Europe.

21/4–31/2 in NORWAY LEMMING STEPPE VOLE MUSKRAT

6–9 cm Lemmus lemmus Lagurus lagurus Ondatra zibethicus
The main small mammal of the European The only species in its Originally an inhabitant of rivers,
densely tundra, the lemming’s fluctuating genus, this vole lives in the dry, ponds, and streams in N. America,
furred population affects the breeding success grassy plains that stretch from the muskrat has been introduced to
tail of many Arctic predators. Ukraine to W. Mongolia. Europe and is now widespread.
 VOL E S , L E M M I NGS , M IC E , R ATS ,
536 A N D M U SK R ATS A N D R E L AT I V E S
One fifth of all mammal species belong
2¾–4¾ in to this huge family. The Muridae are
MAMMALS • RODENTS

7–12 cm distributed almost worldwide, including

the polar regions. Some species carry serious
diseases, and others are significant agricultural
pests. However, several are used in medical
research and are commonly kept as pets.

4–4¾ in
relatively STRIPED DWARF HAMSTER 10–12 cm
long tail Cricetulus barabensis
By raiding crops to gather seeds and grain, this
rodent can become a serious pest in farming areas. NORTHEAST AFRICAN
Farmers destroy its burrows while plowing. SPINY MOUSE
Acomys cahirinus
Like other spiny mice, this rodent has stiffened
2¼–3¼ in
6–8 cm hairs on its body for protection and very thin skin,
which permits easy cooling in its hot, dry habitat.

61/2–121/2 in 3¼–5 in
17–32 cm 8–13 cm
ARABIAN
SPINY MOUSE
ROBOROVSKI’S DESERT COMMON HAMSTER Acomys dimidiatus
HAMSTER Cricetus cricetus This mouse was previously considered
Phodopus roborovskii This burrowing species lives alone the same species as the N.E. African one,
From the dry grasslands of C. Asia, this tiny and hibernates underground through 31/2–51/2 in
except that it occurs east of the Red Sea. 9–14 cm
animal is popular as a pet. During the breeding the winter. Its burrows may contain 4–7 in
season, it can produce three to four litters. up to 145 lb (65 kg) of stored food. 10–18 cm

golden orange fur

4¾–61/2 in
12–17 cm

SHAW’S JIRD
Meriones shawii
Common in deserts of
N. Africa and the Middle MONGOLIAN JIRD
East, this rodent does not Meriones unguiculatus
hibernate, but survives the In the wild, this gerbil lives in the
winter by storing up to 22 lb dry steppes of C. Asia, forming
(10 kg) of food in its burrows. large social groups. Many are
now kept as pets.
31/2–5 in
9–13 cm

4¾–61/2 in LONG-HAIRED GOLDEN HAMSTER

12–17 cm GOLDEN HAMSTER Mesocricetus auratus
Mesocricetus auratus Originally from Syria, PALLID GERBIL
The golden hamster has been where it is now an Gerbillus floweri
selectively bred to create endangered species, this Like most small desert rodents, this
exotic forms, such as this hamster has become a gerbil has a pale body. It is widespread in
albino variety, that would favorite household pet in N. Africa and the Middle East, where its
not survive in the wild. Europe and N. America. coloration provides good camouflage.

4–5 in
31/2–4¼ in 4¾–8 in 10–13 cm
9–11 cm 12–20 cm

FAT-TAILED JIRD
WHITE-FOOTED MOUSE HISPID COTTON RAT Pachyuromys duprasi
Peromyscus leucopus Sigmodon hispidus Like many small, desert-living
A very common and adaptable creature, This short-lived, surface-dwelling mammals, this species stores fat in
this deer mouse inhabits almost every type herbivore occurs in grassland habitats its tail. The tail has no fur, allowing
of land habitat in central and the E. US. in the S. US and Mexico. it to radiate excess body heat.
 pale fur, with grayish- 31/2–5 in
brown markings 9–13 cm
537
NORTHERN LUZON
GIANT CLOUD RAT
Phloeomys pallidus

MAMMALS • RODENTS
The two species of
cloud rat inhabit high
forests of the Philippines,
but are rarely seen. They
are the largest of the YELLOW-NECKED
mouselike rodents. FIELD MOUSE
Apodemus flavicollis
15–17 in This nocturnal woodland species
38–43 cm is difficult to distinguish from the
wood mouse over much of its
European range.
151/2–171/2 in
39–44 cm SOUTHERN LUZON
GIANT CLOUD RAT
Phloeomys cumingi
Like the smaller northern
species, this rat spends the day
in hollow trees or burrows,
and has one young at a time.
31/2–4¼ in
9–11 cm

WOOD MOUSE
Apodemus sylvaticus
This is the most abundant wild mouse in
Europe, living in every terrestrial habitat,
2¾–4 in white even in the mountains.
HOUSE MOUSE 7–10 cm
Mus musculus underparts
A highly adaptable species, this small, small ears
slim rodent has followed humans
all around the world, even
living in the sub-Antarctic. ALBINO DOMESTIC
MOUSE
Mus musculus
The captive-bred albino house
2¾–4 in
mouse has become common as
7–10 cm a pet, and is also widely used in
medical and scientific research.

long whiskers on nose,

robust body eyebrows, and cheeks
HARVEST MOUSE 2–3¼ in
Micromys minutus 5–8 cm
Europe’s smallest mouse, this
species lives in a variety of
grassy habitats, including
reed beds and cornfields.

41/2–10 in
11–26 cm

BROWN RAT BLACK RAT

Rattus norvegicus Rattus rattus
Also called the common rat, this is now Often called the “ship rat” because of its ability
a global pest, having traveled on ships to live aboard ships, this is also the species
and colonized even remote islands. whose fleas transmit bubonic plague.

grayish- 31/2–51/2 in 12–14 in

brown fur 9–14 cm 30–35 cm

51/2–111/2 in
14–29 cm

TYPICAL STRIPED GRASS MOUSE MALAGASY GIANT JUMPING RAT

Lemniscomys striatus Hypogeomys antimena
This distinctive, boldly marked species is The sole species of giant hopping rat, this is
a common inhabitant of grassy habitats the largest rodent in Madagascar, found only
across most of sub-Saharan Africa. in the sandy forests of the west coast.
 BA M BOO R ATS SPRINGHARES SOUTH AFRICAN
SPRINGHARE
538 A N D R E L AT I V E S Resembling rabbits in size and behavior, Pedetes capensis
This species leaves its
The family Spalacidae includes blind mole rats, bamboo rats, root the Pedetidae differ in producing only one burrow at night to nibble
rats, and zokors. Big, protruding incisors characterize the blind young at a time. They breed all year round, on the grass and herbs
MAMMALS • RODENTS

mole rats. Adapted to an underground life, they have no external but living in dry, open habitats makes them found in the dry regions
vulnerable to predators. of southern Africa.
eyes and ears. The bamboo rats of eastern Asia have visible eyes.
61/2–14 in 6–10 in 13–18 in
17–35 cm 15–26 cm 33–46 cm

13–18 in
33–46 cm
long,
tufted tail

GREATER MOLE RAT

Spalax microphthalmus LESSER BAMBOO RAT EAST AFRICAN SPRINGHARE
This blind species has sensory bristles Cannomys badius Pedetes surdaster
running from its snout to its eye Found from Nepal to Vietnam, this single Springhares escape their noctural predators with
sockets. It is native to the steppes species of its genus digs deep burrows in kangaroolike leaps. Less common than Pedestes capensis,
of Ukraine and S.E. Russia. forests, grassy areas, and sometimes gardens. this species lives on the Serengeti plain in Africa.

A F R ICA N MOL E - R AT S N E W WOR L D

Living underground, the Heterocephalidae (naked mole-rats) and the 2 3/4–41/2 in
PORC U PI N E S
Bathyergidae (mole-rats) burrow in sand and soft soils to feed on roots, 7–11 cm Tree-dwelling animals in American forests,
using their protruding front teeth as shovels. Their lips close behind the Erethizontidae have short spines, which are
the teeth to keep the mouth from filling with dirt. generally less than 4 in (10 cm) long. Most have
long, protruding a prehensile tail to grip branches.
incisor teeth

2–41/4 ft
NAKED MOLE-RAT 0.6–1.3 m
Heterocephalus glaber
This highly social animal lives in
a colony, where each individual NORTH AMERICAN
performs different, specialized jobs PORCUPINE
to help the colony as a whole. Erethizon dorsatus
This forest dweller is found
throughout N. America,
from Alaska to Mexico. Its
long, spines are hidden among
rounded its shaggy fur.
tail
NAMAQUA DUNE
MOLE-RAT
Bathyergus janetta 171/2–22 in
Native to Namibia and 44–56 cm
BRAZILIAN
southwestern S. Africa, this PORCUPINE
species uses its forefeet rather Coendou prehensilis
4–71/2 in than its teeth to dig burrows. This nocturnal species
10–19 cm COMMON MOLE-RAT 61/2–91/2 in inhabits forested areas of
Cryptomys hottentotus 17–24 cm S. America and Trinidad.
Found from Tanzania to S. Africa, this It sleeps by day and
common species lives in soft soils and forages for leaves and
farmland, where it feeds mainly on roots. shoots at dusk.

OL D WOR L D
PORC U PI N E S
spines are
Found across most of Africa and southern Asia, modified
the 11 species of the family Hystricidae family live hair 30–39 in
75–100 cm
in burrows. They are covered with long, stiff
quills that protect them against most
predators. When attacked, they often
rattle their quills as a reminder of how
sharp and impenetrable these are.

CRESTED 18–361/2 in
PORCUPINE 45–93 cm
Hystrix cristata
Widespread across the CAPE PORCUPINE
northern half of Africa, Hystrix africaeaustralis
except the Sahara, the Found in savanna across most of southern
crested porcupine is a Africa, this species forages at night, alone or
familiar nocturnal rodent. in groups, sniffing out roots and berries.
V ISCAC H A S A N D large ears help
regulate body
CHINCHILLAS temperature long whiskers
539
aid spatial
All six species of the family Chinchillidae from South awareness
America have a prominent tail and large hind feet. They
normally live in social groups, inhabiting burrows or
rocky outcrops. Most species are now quite rare because
of their exploitation for fur and their status as pests.

CHINCHILLA
Chinchilla sp.
Widely prized for their fur,
chinchillas have a fine, thick
coat that insulates them 12–18 in
against the cold in their 30–45 cm
Andean home.

9–15 in
22–38 cm

MOUNTAIN VISCACHA
bushy tail Lagidium viscacia
balances This agile rodent has dense fur to protect
the body
it from the cold at night. It lives on
steep, rocky, mountain slopes.

DA SSI E R AT CA N E R AT PAC A R A N A
This single species in the family Petromuridae The two species of the family Thryonomyidae The single species of the family
is found only in southern Africa. Its peculiar, have pale brown, coarse, flattened hair that blends Dinomyidae is a shy, bulky,
flat skull and soft ribs are adaptations to living with their surroundings in dry grass and cane slow-moving, and almost
in crevices and under stones, thickets. Cane rats produce two small defenseless animal. It lives
and distinguish it from all families of well-developed 16–30 in alone or in pairs in montane
other rodents. young per year. 41–77 cm forests, where it is preyed
upon by jaguars and humans.
CANE RAT
DASSIE RAT Thryonomys sp. PACARANA
Petromus typicus There are two species Dinomys branickii
Living on dry, rocky hillsides, of African cane rats: one Threatened by loss of its
this rodent emerges from lives in savanna grasslands S. American forest habitat,
5–10 in crevices at dawn and dusk to and the other in reedbeds and hunted for food, the pacarana 28–32 in
13–25 cm forage for seeds and shoots. and marshes. is now an endangered species. 70–80 cm

GUINEA PIGS, ROSETTE GUINEA PIG

Cavia porcellus
MARAS, AND Several coat variations occur in
CAPYBARA pet guinea pigs. In this one, the
fur forms substantial spiky
Including some of South America’s most whorls over the body.
widespread and abundant rodents, the 8–16 in long ears
Caviidae occur from mountain meadows 8–16 in 20–40 cm
to tropical flood plains, and breed all year BRAZILIAN GUINEA PIG 20–40 cm
Cavia aperea 231/2–32 in
round. All but the maras and capybara Guinea pigs mostly live in lowland 60–80 cm
are short-legged and dumpy animals. habitats, but this species is also found
8–16 in in the Andes, from Peru to Chile.
20–40 cm

PATAGONIAN
MARA
Dolichotis patagonum
Extensive shared
burrow systems and
communal breeding
LONG HAIR GUINEA PIG SHORT HAIR GUINEA PIG habits are among the
8–16 in unusual features of these
Cavia porcellus Cavia porcellus 20–40 cm long-legged rodents.
Often kept as a pet, in captivity this First domesticated more than 500 years
long-haired breed needs grooming to ago for food, guinea pigs have since
prevent its fur from becoming tangled. become popular pets worldwide.
C R E S T E D P O RC U PI N E 541
Hys t r i x cr is tata
With its long, barbed quills raised in threat, the crested porcupine

MAMMALS • RODENTS
is an intimidating prospect. A predator that has had the painful experience
of being spiked is unlikely ever to attempt another attack. Lions, hyenas,
and even people have been known to die from infected quill injuries. Despite
this impressive means of defense, the porcupine is a peaceable and rather
nervous animal, easily startled, and more likely to flee danger
than stand its ground. Porcupines live alone or in family groups,
sharing extensive burrow systems. Crested porcupines occur
throughout much of northern Africa. The species was also
once widespread in southern Europe—the population found
in Italy may be a relic of former times or the result of a more
recent introduction, perhaps by the Romans.
size 18–36½ in (45–93 cm)
habitat Savanna grassland,
EYE light woodland, rocky terrain
A porcupine’s eyesight is poor, distribution Northern
but then there is often little to Africa as far south as Tanzania,
see in the darkness of an African except Sahara Desert; also Italy
SPIKY HAIRDO EAR night. It uses its sharp senses of diet Mainly roots, fruits, and
Raising the quills creates an The small ears hearing and smell to find its way. tubers; occasionally carrion
illusion of size. A cornered are largely hidden in
porcupine will stand tall and the coarse hair. The
try to bluff its way out of porcupine has good
trouble. If this fails, it will turn hearing, which it uses MOUTH AND TEETH
tail and charge backward, to avoid danger, Porcupines have the specialized
shoving its spiky rear end shuffling off into the gnawing teeth typical of rodents,
in the face of its attacker. night if it hears enabling them to chew tough roots
another animal and tubers. The muscles that control
approaching. the jaws are immensely powerful.
QUILLS-A-QUIVER
Porcupine quills are greatly
enlarged hairs. They can
be raised by larger versions
of the tiny muscles that create
goosebumps in our own skin.

like its quills, the

porcupine’s thin coat
of coarse hair can
also be raised
in alarm

RATTLING TAIL FEET AND CLAWS

The quills of the tail are swollen Porcupines walk on the flat soles of their
and hollow. The porcupine shakes feet, with a slightly clumsy, shambling
them when alarmed and the soft gait. The soles are hairless and padded.
rattling sound warns its enemies They have short toes and strong claws,
that this is a well-armed opponent. well suited to digging.
 CAPYBARA PAC A S
542 The largest rodent, the capybara, is one of four species in the Caviidae subfamily This family includes two species of nocturnal rodent
Hydrochoerinae. Capybaras generally breed once a year; the offspring are born at the from Central and South America. The Cuniculidae
end of the wet season, when the grass is most nutritious. They may live up to six years. resemble small pigs, as they poke around on the
forest floor for fruits, seeds, and roots.
coarse fur CAPYBARA
dries quickly Hydrochoeris 20–30 in
hydrochaeris 50–75 cm
As big as a pig, this is
the world’s largest rodent.
It lives a semiaquatic LOWLAND PACA
life in the swamps Cuniculus paca
of S. America. Mainly a forest inhabitant,
this rodent lives in the
31/4–41/4 ft northern part of S. America,
1–1.3 m from Mexico to Paraguay.

DEGU S , ROC K R AT S ,
A N D R E L AT I V E S
The molar teeth of these small, silky-furred rats form
a figure-eight shape when worn, giving them their
Latin name. The Octodontidae are widespread in the
southern part of South America.

61/2–9 in
16–22 cm

small, rounded
ears
DEGU
Octodon degus
H U T I A S , SPI N Y R AT S , A N D C OY PU This species is found on the western slopes of the Andes
in Chile. Its tail breaks off easily if caught by a predator.
A Central and South American family, the Echimyidae comprises 99 species that live
in a variety of habitats, and includes tree-dwelling and burrowing spiny rats, and the
semiaquatic coypu. Although they vary in size, fur type, and dentition as well as diet, AG OU T IS
phylogenetic and molecular analyses suggest they belong in the same group.
Active during the day, these long-legged, running
rodents of the family Dasyproctidae are very shy.
61/2–12 in 12–17 in
16–30 cm 30–43 cm They breed all year round but have only two young
at a time, which can run within an
hour of birth.

RED-RUMPED AGOUTI
Dasyprocta leporina
Found in forested areas of
northeastern S. America and
the Lesser Antilles, this
SPINY RAT DESMAREST’S CUBAN HUTIA species can be identified by
Proechimys sp. Capromys pilorides its light orange rump.
These rodents have a spiny This hutia is common in Cuba. Other surviving 19–231/2 in
48–60 cm
protective coat. This feature has evolved hutia species face extinction because of a loss 19–231/2 in
in parallel to the spiny mice of Africa. of their habitat and hunting. 48–60 cm

17–23 in
181/2–23 in 43–58 cm
47–58 cm prominent
incisor teeth

COYPU CENTRAL AMERICAN AZARA’S AGOUTI

Myocastor coypus AGOUTI Dasyprocta azarae
This animal has Dasyprocta punctata An inhabitant of the forests
long, distinctively shaggy fur With a range that extends from of S. Brazil, Paraguay, and
rounded tail and huge orange front Mexico south to Argentina, this N. Argentina, this agouti
teeth. It also has webbed agouti mainly feeds on fruits, barks when alarmed.
hind feet for swimming but may also eat crabs. Pairs are It eats a variety of
and a thick, scaly tail. thought to stay together for life. seeds and fruits.
TREE SHREWS 543
These small mammals look and behave their name, most of them are only partly PHYLUM CHORDATA
like squirrels. They are active in the arboreal. Their mixed diet includes insects,

MAMMALS • COLUGOS
CLASS MAMMALIA

daytime and spend much of their time worms, fruits, and sometimes small mammals, ORDER SCANDENTIA

foraging for food on the ground. reptiles, and birds. FAMILIES 2

Some tree shrews are solitary, while others SPECIES 23

Tree shrews are native to the tropical rainforests live in pairs or groups. They are rapid breeders,
of Southeast Asia. They are unrelated to the true raising their young in nests in tree crevices
shrews and have their own order, Scandentia. or on branches. The female pays her brood
Sharp claws on all their fingers and toes enable little attention, making only the occasional
tree shrews to climb trees rapidly but, despite short visit to suckle them.

PE N -TA I L E D TREE SHREWS

TREE SHREW Members of the family Tupaiidae
The family Ptilocercidae has only one have a long snout, which is used
member, the Southeast Asian pen-tailed to locate insects and other
tree shrew. The species is named for its invertebrates as well as fruits long snout
long, feathery tail, which resembles a quill and leaves. Tree shrews also
pen. This aids balance when climbing. have sharp claws on all their
fingers and toes, helping
them to climb trees rapidly.
5–6 in 61/2–81/2 in
13–15 cm 17–21 cm
PEN-TAILED
TREE SHREW LARGE TREE SHREW elongated claws
Ptilocercus lowii Tupaia tana help hold on to
This species has a rather Tree shrews are diurnal tree branches
spindly tail with a inhabitants of S.E. Asian
brushy tip, unlike most forests. This species is found
other tree shrews, which in Borneo and Sumatra
have a thick, bushy tail. and on nearby islands.

COLUGOS
The two species that form the order to tree. The distance covered in a glide may PHYLUM CHORDATA
Dermoptera are gliding, rather than be more than 330 ft (100 m). Colugos live in CLASS MAMMALIA

flying, mammals. They are found in the canopy, hanging upside down from branches ORDER DERMOPTERA

the rainforests of Southeast Asia. or sheltering in cracks or hollow trees by day FAMILIES 1

and emerging at night to feed on fruits and leaves. SPECIES 2

The colugos are distinguished by a furry They are almost unable to move on the ground.
membrane that stretches from the neck to the The teeth of colugos are unlike those of any
tips of the fingers and tail. When the limbs are other mammal. In the lower jaw, the teeth
splayed this membrane spreads out, allowing the are arranged like combs and are believed to
lemur to maneuver in the air as it glides from tree be used for grooming as well as for feeding.

F LY I N G L E M U R S PHILIPPINE
FLYING LEMUR
Cynocephalus volans
The two members of the Cynocephalidae The only local species
have forward-facing eyes that provide depth found in the forests of the
perception This helps them judge distances S. Philippines, where it feeds
when gliding from tree to tree. They also mainly on young leaves.
have unusual comblike lower teeth, used to
strain food such as fruits and flowers. furry flight
membrane

SUNDA COLUGO
Cynocephalus variegatus large, forward-
facing eyes
This lemur lives alone or in small groups,
inhabiting tree-holes or resting high
on treetops in the tropical forests of 131/2–161/2 in 131/2–161/2 in
S.E. Asia and the Indonesian islands. 34–42 cm 34–42 cm
544 P R I M AT E S
The order to which humans belong, than other primates. The Haplorrhini includes PHYLUM CHORDATA
primates have a large brain relative the New and Old World monkeys and the apes,
M A MM ALS • PRIM ATES

CLASS MAMMALIA

to their size and forward-facing eyes many of which are diurnal, and so more reliant ORDER PRIMATES

which give them 3-D vision. on vision than the Strepsirrhini. FAMILIES 16
SPECIES 506

With the exception of a few species, including s o c i a l pat t e r n s

humans, primates are restricted to tropical and Most primates are highly social, living in small D E B AT E
subtropical regions of the Americas, Africa, and family groups, single-male harems, or large P R I M AT E H O T S P O T S
Asia. They range in size from a mouse lemur, mixed-sex troops. Many species are characterized
weighing 1 oz (30 g), to a gorilla weighing up by high levels of male competition for females. Today scientists list more species of
to 440 lb (200 kg). Sexual selection favoring the largest or most primates than a decade ago—most
the result of subspecies (geographical
Primates rely more on vision than on smell. dominant males has led to sexual dimorphism—
variants) being recognized as new
Many are arboreal and have characteristics such differences between males and females in body species. In the Amazon basin
as stereoscopic vision, allowing good judgment size or features such as canine teeth. The different populations of monkeys separated
of distance for jumping between trees; opposable sexes may also be differently colored, a form by rivers and mountain ranges can
thumbs and a prehensile tail for grasping branches; of dimorphism known as sexual dichromatism. differ in subtle ways—such as in the
long legs for leaping; and long arms for swinging. Most New World monkeys are monogamous, structure of their chromosomes. Some
Some primates have specialized diets, but many with both parents sharing responsibility for scientists believe that these monkeys
other species are omnivorous. raising their young. Old World monkeys tend to diverged when their forest habitats
live in groups dominated by related females, and became isolated due to geological
activity many thousands of years ago.
m a j o r g ro u p s the males take little or no part in parental duties.
Such forests may contain many species
Primates are divided into two suborders. Primates are generally slow to mature fully and isolated in this way and are a particular
The Strepsirrhini includes the mostly nocturnal slow to reproduce, but they are relatively long- focus of conservation groups aiming
lemurs, lorises, galagos, and their relatives. These lived. The larger apes have a potential lifespan of to protect biodiversity.
species have a better developed sense of smell up to 45 years in the wild, and longer in captivity.

GA L AG OS large,
thick fur
Native to sub-Saharan Africa, the members movable ears
of the family Galagidae inhabit a wide range
of woodland and forest habitats, including
scrub and wooded savanna. They have
longer hind legs than
forelegs, which help
them to take large leaps
between trees in forests.
These primates often
wash their hands and feet
with urine, which may help
to improve their grip and leave
scent-trails. All are nocturnal.

huge eyes

11–18½ in
28–47 cm

SILVERY GREATER GALAGO

Otolemur monteiri BROWN GREATER GALAGO
This species has the same range as the Otolemur crassicaudatus 10–16 in
brown greater galago but prefers denser One of the largest galagos, this primate lives in a variety 26–40 cm
vegetation. Despite its name, melanistic of forests in southern Africa. This omnivore uses its
(black) forms are common. comblike teeth to scrape gum from trees.
LORISES, POTTOS, 7–8½ in
18–21 cm
A N D A N G WA N T I B O S 545
These small, nocturnal omnivores have a short tail, and their
forelimbs and hind limbs are of equal length. The Lorisidae

M A MM ALS • PRIM ATES

have opposable thumbs for grasping branches and they move
much more slowly and sedately through the trees than the
galagos—climbing rather than leaping.
RED SLENDER LORIS
Loris tardigradus
dense fur Native to Sri Lanka, this
slender primate moves carefully
through the forest canopy
on its long limbs.

opposable thumb

dark rings
around
huge eyes 9–10 in
22–26 cm
vicelike grip
12–13½ in 8–9 in 12–16 in
30–34 cm 20–23 cm 30–40 cm

SUNDA SLOW GOLDEN

LORIS ANGWANTIBO
Nycticebus coucang Arctocebus aureus
As its name suggests, PYGMY SLOW LORIS WEST AFRICAN POTTO Also called the golden
this loris moves slowly and Nycticebus pygmaeus Perodicticus potto potto, this species inhabits
deliberately through the This species inhabits thick tropical A shy species, this primate lives in thick the understory of moist
trees. It inhabits tropical rainforests and bamboo forests in Laos, rainforests across equatorial Africa. A bony shield lowland forest in equatorial
forests in S.E. Asia. Cambodia, Vietnam, and S. China. at the nape of its neck protects it against predators. W. and C. Africa.

SENEGAL
BUSH BABY TA R SI E R S
Galago senegalensis
Widespread across
The small Tarsiidae are named for
C. Africa and parts of large ears their greatly elongated anklebone
E. Africa, the lesser, or or tarsus. These arboreal species
Senegal, bush baby inhabits have long limb bones, elongated
arid savanna woodland. fingers, and a long, thin tail. Their
round head has very large eyes,
4¾–8 in 4¾–6½ in which help them to see at night
12–20 cm 12–17 cm
while hunting for insects.

silky fur

PHILIPPINE
TARSIER
Tarsius syrichta
Endemic to a variety of rainforests
and scrub in the Philippines, this tarsier
possesses the largest eyes relative to
its body size among all mammals.

MOHOLI BUSH BABY

Galago moholi
This small, shy, galago lives 4½–5½ in 4½–5½ in
in small groups in southern 11–14 cm 11–14 cm
Africa. Lithe and agile, it
leaps through woodland,
where it eats insects WESTERN TARSIER
and tree gum. Tarsius bancanus
DEMIDOFF’S 4¾–6½ in Adapted for clinging and
BUSH BABY 12–17 cm
Galagoides demidovii leaping between trees,
long tail this species, also called
Also called the dwarf the Horsfield’s tarsier, lives
bush baby, this small in the tropical rainforests
primate uses its long hind of Sumatra and Borneo.
legs to jump through
rainforest canopies of long,
W. and C. Africa. slender tail
 LEMURS
546 Found in forests across Madagascar, the family Lemuridae
is made up of the most typical lemurs: mainly arboreal and
quadrupedal. Most are cathemeral, meaning they are active 16–16½ in 15½–18 in 16–20 in
40–42 cm 39–46 cm 40–50 cm
M A MM ALS • PRIM ATES

during the day and night. In several

species, males and females are
differently colored.

GREATER RING-TAILED BLACK-AND-WHITE

BAMBOO LEMUR LEMUR RUFFED LEMUR
Prolemur simus Lemur catta Varecia variegata
One of the rarest Living in groups of up to The largest lemur,
15–16 in lemurs, this species 25 individuals, this lemur this species eats a high
38–40 cm
lives in S.E. Madagascar, often spends time on the proportion of fruits.
thick, where it feeds almost ground. Its diet consists Unusually for lemurs,
woolly fur exclusively on of fruits, vegetation, it makes a leafy nest
giant bamboo. sap, and bark. for its young.

14–16½ in 15–16½ in
35–42 cm 38–42 cm

BANDRO RED COLLARED

Hapalemur alaotrensis LEMUR
Also called the Lac Alaotra RED-BELLIED LEMUR Eulemur collaris
bamboo lemur, this Eulemur rubriventer This lemur has a scent
critically endangered species A monogamous species, this gland on its wrist, with
lives only in papyrus marsh lemur lives in small groups that which it perfumes its
and reedbeds around Lake consist of a mated pair and long, furry tail for use
Alaotra, Madagascar’s their dependent young. in communication.
largest lake.

tail is as
long as body

15½–16½ in 12½–14½ in
39–42 cm 32–37 cm

WHITE-HEADED
LEMUR
Eulemur albifrons
Only the male of this
species has distinctive
white fur around its
black face; the female has red cheek
a gray face. patch in male

MONGOOSE
LEMUR
Eulemur mongoz
Primarily nocturnal
during the dry season,
the mongoose lemur
becomes more diurnal
BLACK LEMUR at the start of the
Eulemur macaco wet season.
In this sexually dichromatic
species, the colors of males and
females vary. Only the male is grasping hands
black; the female is gray-brown
with white ear tufts.

15–18 in
38–45 cm
M O U S E , DWA R F, PALE FORK-MARKED
LEMUR
10–12 in
26–30 cm
A N D F O R K- Phaner pallescens 547
MARKED LEMURS Adapted to eat tree gum,
this lemur has a long tongue
Among the smallest of all primates, the and large premolars

M A MM ALS • PRIM ATES

for chiseling bark.
members of the family Cheirogaleidae have
short limbs and large eyes. All are nocturnal, 9–12 in
22–30 cm
arboreal inhabitants of forests in Madagascar,
and become torpid to survive the dry season.

short
4¾–6 in limbs
12–15 cm

GRAY GREATER
MOUSE LEMUR EASTERN RUFOUS DWARF LEMUR
Microcebus murinus MOUSE LEMUR Cheirogaleus major
An omnivore, this lemur’s diet Microcebus rufus This solitary lemur
includes insects, flowers, and fruits. Inhabiting a range of forest feeds mainly on fruits
The young are carried in their habitats, this omnivorous and nectar. It stores fat
mother’s mouth if moved (rather species consumes a variety 4–6 in in its tail during the
than clinging to her fur). of fruits, insects, and gum. 10–15 cm wet season.

SPORT I V E L E M U R S SI FA K A S A N D 16–20 in
40–50 cm
The family Lepilemuridae consists of the sportive R E L AT I V E S
lemurs of Madagascar. These medium-sized lemurs The largest lemurs—the indri and sifakas—and the
have a prominent muzzle and large eyes, and are smaller avahis or woolly lemurs form the Indriidae
strictly arboreal and nocturnal. Their low-energy, of Madagascar. All members have long, powerful limbs
leafy diet means that they are among the least active for leaping between trees, and, with the exception of
of all primates. the indri, a long tail.

9–10 in face mostly bare, with white

23–26 cm fur along muzzle

16½–20 in
7½–10 in 42–50 cm
19–26 cm
BLACK-STRIPED
SPORTIVE LEMUR
Lepilemur dorsalis long hind VERREAUX’S SIFAKA
WHITE-FOOTED limbs Propithecus verreauxi
SPORTIVE LEMUR This species lives
Lepilemur leucopus in moist forest in Found in S.W. Madagascar, this
Grayish above and white below, this N.W. Madagascar and sifaka is capable of leaping through
16½–20½ in cactuslike vegetation on its long
lemur spends lengths of time perched nearby islands. It has 42–52 cm
vertically on tree trunks, in between a blunt muzzle and legs without being injured.
bouts of foraging for food. small ears.
25–28 in
64–72 cm
AY E -AY E
There is only one species in the Daubentoniidae—
the aye-aye of Madagascar. This nocturnal primate
has large, bare ears; a shaggy coat; very long fingers;
and continually growing incisor teeth.

12–14½ in
30–37 cm
MILNE-EDWARD’S COQUEREL’S
AYE-AYE SIFAKA SIFAKA
Daubentonia Propithecus edwardsi Propithecus coquereli INDRI
madagascariensis Occurring in small family Like other sifakas, this Indri indri
The aye-aye uses its groups in southeastern primate crosses open The largest member of the
elongated middle finger Madagascar, this sifaka has a ground by skipping on lemur group, this is
for locating and extracting large opposable thumb for its hind legs, using its the only species to have
insect larvae from dead wood. clinging to tree trunks. forearms for balance. a short, vestigial tail.
 HOW L E R , SPI DE R , A N D 19–25 in
48–63 cm
548 WO O L LY M O N K E Y S
The howler, spider, and woolly monkeys, and muriquis of
the Atelidae are the largest New World monkeys. All have
M A MM ALS • PRIM ATES

a prehensile tail that is used as a fifth limb when moving

through the trees. Spider monkeys have longer limbs
than the other members of the family.

20–28 in GUATEMALAN
50–71 cm BLACK HOWLER
Alouatta pigra
Found in the Yucatán
Peninsula of Mexico,
Belize, and Guatemala,
this species forms
groups of up to
11 individuals.

large larynx
for howling
VENEZUELAN
RED HOWLER
Alouatta seniculus
This howler monkey has an
enlarged hyoid bone in the
neck that allows it to make
calls that can be heard
several miles away.
18–25 in
46–63 cm

MANTLED HOWLER
Alouatta palliata
Named after the “mantle” of long
guard hair on its flanks, this howler
monkey lives in C. America and
12–25 in northern S. America.
30–64 cm

12–25 in 18–31 in
31–63 cm 46–78 cm
COLOMBIAN BLACK GEOFFROY'S SOUTHERN
SPIDER MONKEY SPIDER MONKEY MURIQUI
Ateles fusciceps rufiventris Ateles geoffroyi Brachyteles arachnoides
Like other spider monkeys, this Active during the day, this Also known as the woolly
subspecies does not have fruit-eating species lives in spider monkey, this species,
thumbs on its hands. It is found relatively large groups of up found in Brazilian forests, is
in Colombia and Panama. to 35 individuals in forests critically endangered due
throughout C. America. to the loss of its habitat.

GRAY HUMBOLDT’S
WOOLLY MONKEY WOOLLY MONKEY
Lagothrix cana Lagothrix lagotricha
A powerfully built species, One of the largest New World
this monkey lives in large monkeys, this species lives in
troops in the primary forests lowland primary forest in the
of Brazil, Bolivia, and Peru. upper Amazon basin.

hairless 18–26 in 18–26 in

tail tip 45–65 cm 46–65 cm
NIGHT MONKEYS
Also called owl monkeys, the members 14–16½ in 12–15 in 549
35–42 cm 30–38 cm
of the Aotidae are the only nocturnal
monkeys in the New World. They are BLACK-HEADED NORTHERN

M A MM ALS • PRIM ATES

small, with large eyes set in a flat, round NIGHT MONKEY NIGHT MONKEY
Aotus nigriceps Aotus trivirgatus
face with dense woolly fur. They have This monogamous species Also called the three-striped
a well-developed sense of smell. inhabits the central and upper monkey, this species is most
Amazon basin in primary and active on moonlit nights. It
secondary forests of Brazil, lives in forests in Venezuela
Bolivia, and Peru. and northern Brazil.

TITI, SAKI, AND black hair

tipped
12½–16½ in WHITE-FACED SAKI
Pithecia pithecia
UAKARI MONKEYS
32–42 cm
with white Male white-faced or Guianan
sakis are black with pale fur
This family contains a range of small- to medium-sized around the face, whereas
monkeys. The Pitheciidae are diurnal, arboreal, and social. females are gray-brown.
All share a common dental makeup, including large,
splayed canine teeth that help them to deal with
tough seeds and fruits.

13½–16½ in
34–42 cm

14–21 in 14½–19 in
36–53 cm 37–48 cm
BLACK-BEARDED
SAKI MONK SAKI
Chiropotes satanas Pithecia monachus
This species is known A shy primate, this species
from S. Amazonia. The males lives in the high canopy of forests
possess a beard and large in N.W. Brazil, Peru, Colombia,
swellings on their forehead. and Ecuador.
RIO TAPAJÓS SAKI hairless,
Pithecia irrorata red face
Also called the bald-faced saki,
this species lives in W. Brazil,
N. Bolivia, and E. Peru. It eats
predominantly seeds.

long,
shaggy coat

12–16½ in 11–14 in
31–42 cm 28–36 cm

BLACK-FRONTED COLLARED TITI MONKEY

TITI Callicebus torquatus
Callicebus nigrifrons Also known as the yellow-handed titi,
A fruit-eater, this titi is an this species prefers unflooded forests
inhabitant of Atlantic coastal on sandy soils in Brazil. It feeds
forest around São Paulo mostly on fruits and seeds.
in S.E. Brazil.

10½–13½ in 14–22½ in
27–34 cm 36–57 cm
14–22 in
35–56 cm
RED BALD-HEADED
UAKARI
COPPERY TITI SPIX’S BLACK- Cacajao calvus rubicundus
Plecturocebus cupreus HEADED UAKARI Several subspecies of the
Found in rainforests of the Cacajao ouakary bald-headed uakari inhabit
S.W. Amazon Basin, the A highly social species, this seasonally flooded forests of
coppery titi is monogamous uakari lives in groups of 30 the Amazon basin. The redness
and territorial. It feeds or more individuals in the of the face is thought to signal
mainly on fruits. N.W. of the Amazon basin. the health of the monkey.
 M A R MOSE T S A N D TA M A R I NS
550 The Callitrichidae consists of relatively small social monkeys, which inhabit
a variety of forests throughout tropical and subtropical Central and South
America. They are all diurnal and arboreal and have forward-facing eyes
M A MM ALS • PRIM ATES

and a short muzzle. Marmosets and tamarins have a long, nonprehensile

tail, lack the third molar, and have claws instead of nails.

7½–10 in
19–25 cm

8–9 in
20–23 cm

GOELDI’S
MONKEY SILVERY
Callimico goeldii MARMOSET
Inhabiting dense forest Mico argentatus
undergrowth, such as A specialized gum-eater
that of bamboo forest in (gumivore), this marmoset
the upper Amazon, this has huge ears, narrow jaws,
monkey makes forays into and short canine teeth for
the tree canopy for fruits. cutting into tree bark.

COMMON MARMOSET
Callithrix jacchus
The female of this species generally
mates with two males, both of
whom assist with parental care
of the usually twin offspring.
6½–8½ in
16–21 cm

long, curved claws

4¾–6 in
12–15 cm
7–9 in
18–23 cm 8–9 in
20–23 cm

GEOFFROY’S
TUFTED-EAR MARMOSET
Callithrix geoffroyi
Also called the white-headed marmoset, BLACK-TUFTED-
EAR MARMOSET PYGMY MARMOSET
this species uses scent-marks to deter Callithrix penicillata Cebuella pygmaea
others from using the holes it makes in
tree bark to extract gum. A monogamous species, The smallest monkey in the world, this
this primate is diurnal and lives species eats tree gum in seasonally flooded
high in the rainforest canopy, forests of the upper Amazon basin.
where it feeds on tree sap.

fine, yellow
hair on back

9–10 in 8½–11 in 9–13 in

23–26 cm 21–28 cm 23–33 cm
white moustache
WHITE-LIPPED BARE-FACED
TAMARIN TAMARIN
Saguinus labiatus Saguinus bicolor
EMPEROR TAMARIN The dominant female of Also called the pied
Saguinus imperator this species releases chemical tamarin, this tree-dwelling
Distinguished by its long, white signals, called pheromones, that primate lives in lowland
moustache, this tamarin lives in suppress reproduction forest in the central
the tropical forests in Peru, in other female members Amazon region near
Brazil, and Bolivia. of the group. Manaus, Brazil.
 8–10 in 8½–11 in
20–25 cm 21–28 cm
GOLDEN-HANDED TAMARIN
551
Saguinus midas
Found in northeast S. America, this
tamarin has brightly colored hands and

M A MM ALS • PRIM ATES

feet, with claws on all digits except
the big toe, which has a nail.

COTTON-TOP
TAMARIN
Saguinus oedipus
Inhabiting a very restricted
range in N.W. Colombia
and Panama, this species
feeds mainly on insects 8–10½ in
and fruits. 20–27 cm
GOLDEN LION
TAMARIN
Leontopithecus rosalia
Among the most
endangered monkeys in
banded tail the world, this species
is found only in the
Atlantic coastal forest
of S.E. Brazil. 9–10 in
22–26 cm
SADDLE-BACK
TAMARIN
Saguinus fuscicollis GOLDEN-HEADED
This tamarin lives in secondary LION TAMARIN
forest and forest edges in the 10–13 in Leontopithecus chrysomelas
26–33 cm
upper Amazon basin, foraging Restricted to the Atlantic forest of southern
for insects, fruits, nectar, and Bahia in N.E. Brazil, this primate raises its
tree sap and gum. mane to look bigger when in danger.

SQUIRREL MONKEYS
AND CAPUCHINS
13–18 in
33–45 cm
10–14½ in
25–37 cm
The family Cebidae once contained several monkey naked face
groups, but now only the squirrel monkeys and
capuchins remain. They have large ears, a long
tail, and forelimbs that are shorter
than the hind limbs. Squirrel
monkeys, of which there are
eight Central and South
American species, belong
to the genus Saimiri.
The capuchins WHITE-HEADED
comprise two CAPUCHIN
genera, the tufted Cebus capucinus
Sapajus and the The only capuchin found in
prehensile C. America, the range of
untufted Cebus. tail this primate extends from
14½–18 in Honduras to the coasts of
37–46 cm
Colombia and Ecuador.

WEEPER CAPUCHIN
Cebus olivaceus
Native to north-central S. America, this bright yellow limbs
capuchin often uses its prehensile tail to
support its body while feeding with its hands.
nails on digits

10½–12½ in
27–32 cm
long tail with GUIANAN SQUIRREL
BLACK-CAPPED a black tip MONKEY
SQUIRREL MONKEY Saimiri sciureus
Saimiri boliviensis A gregarious species, this monkey lives
Males of this species become fatter in large groups in a wide variety of
around the neck and shoulders during the forests in northern S. America to
breeding season, when they compete for mates. north-eastern S. America.
552 GUIANAN SQUIRREL
MONKEY
Saimir i sciu reu s
M A M M A L S • P R I M AT E S

Inquisitive and intelligent, the Guianan squirrel monkey has the largest brain
mass to body mass of any primate. It is also very social, living in large
mixed-sex groups of about 15–50 individuals.
Guianan squirrel monkeys are synchronous
breeders, with all young being born over a
single week during the wet season (January–
size 10–141⁄2 in (25–37 cm)
February) after a gestation period of about six months. The young initially cling to habitat Lowland rainforest, mangroves
their mother’s abdomen but then start riding on her back at about two weeks old. distribution French Guiana, Guyana, Suriname, and Brazil
Their development is rapid, and the youngsters are weaned at six months and are fully north of the Amazon
diet Mainly insects, spiders, fruits, and flowers
independent four months later. Squirrel monkeys can live for up to 20 years.

EYE
Sharp vision and good NOSTRILS
depth perception are a Broad, laterally facing nostrils
result of having large, and a short nasal region belie
close-set, forward-facing an excellent sense of smell.
eyes. These features are Scent is important for social
of vital importance to a communication between
monkey that spends most members of the group and for
of its time in the trees. finding mates and lost infants.

TEETH
The Guianan squirrel monkey has 36
small, sharp teeth. The canines are
longer and more prominent than the
other teeth, and males have larger
upper canines than females.
FOOT
HAND Unlike the hand, the foot has a large, opposable digit—the
Although the hand is used to grip branches, big toe. When used in conjunction with the other four
hold food, self-groom, and perform other toes, it allows the foot to grasp branches.
tasks, the fingers do not move independently,
so the thumb and first finger cannot be
opposed and used to pick things up.

BALANCING ACT
The tail is at least as long as the
head and body lengths combined.
It is used to help with balance on
tree branches, as squirrel monkeys
move on all fours.
body covered
by a short,
thick coat
 FACIAL FEATURES
The face of the Guianan squirrel
monkey has several distinctive 553
features: the white hair above
“Gothic” arches of each eye forms a slightly
white hair meet to pointed arch, the ears are hairy

M A M M A L S • P R I M AT E S
form deep V-shape and have ear tufts, and there
between the eyes are conspicuous dark hairs
(vibrissae) just above the eyes.
 OL D WOR L D MON K EYS 17–21 in
43–53 cm
TOQUE MACAQUE
Macaca sinica
554 Widely distributed across Africa and Asia, the Cercopithecidae have The smallest of the macaques,
this species is endemic to
closely spaced, downward-facing nostrils (catarrhine) and flattened the wet forests on the
nails. Most are diurnal and arboreal; however, baboons are mainly island of Sri Lanka.
M A MM ALS • PRIM ATES

terrestrial. Guenons, baboons, and macaques are omnivorous, with

strong jaws, cheek pouches, and a simple stomach. Colobus and leaf
monkeys are leaf-eaters (folivorous) with a complex stomach, and
lack cheek pouches.

211/2–25 in
54–64 cm

forward-facing
eyes give
3-D vision

141/2–26 in
37–66 cm
171/2–221/2 in
44–57 cm

CELEBES CRESTED MACAQUE BARBARY MACAQUE

Macaca nigra Macaca sylvanus
Endemic to Indonesia’s Sulawesi Island, The only macaque outside Asia,
this monkey has bare, pink swellings on its the barbary macaque lives in the
rump. These become particularly swollen high cedar and oak forests of
in females that are ready to mate. Algeria and Morocco.

lionlike
mane
12–25 in
31–63 cm

16–24 in
40–61 cm

CRAB-EATING
MACAQUE
Macaca fascicularis
Besides eating crabs, as
its name suggests, this
omnivorous S.E. Asian
species also forages LION-TAILED MACAQUE
for insects, frogs, Macaca silenus
fruits, and seeds. Endemic to the Western Ghats,
mountains in S.W. India, this
arboreal macaque lives mainly
in wet, monsoon forests.

RHESUS MACAQUE
Macaca mulatta
This macaque lives in dry, 19–26 in
open areas from W. 48–65 cm
Afghanistan through India to
N. Thailand and China. Adults
swim distances of up to 1/2 mile
(0.8 km) between islands.

STUMP-TAILED MACAQUE
Macaca arctoides
Both arboreal and terrestrial,
this species is found in tropical
and subtropical wet forests
sandy coloration in S.E. Asia.
 14–231/2 in
35–60 cm thick fur
555

M A MM ALS • PRIM ATES

17–29 in
43–74 cm

SOUTHERN BONNET MACAQUE 18–251/2 in JAPANESE MACAQUE

forearms of similar PIG-TAILED MACAQUE Macaca radiata 46–65 cm Macaca fuscata
length to hind legs Macaca nemestrina Often found close to Inhabiting the most northerly range
This monkey inhabits humid human habitation, this of any nonhuman primate, the
areas, including rainforest resident of S. India is Japanese macaque uses hot springs
and swamps in S.E. Asia. Its omnivorous and may rely to keep warm in winter.
diet consists mainly of fruits. on humans for food.

opposable 8–27 in 131/2–201/2 in

thumbs 20–68 cm 34–52 cm

WHITE-THROATED
GUENON
Cercopithecus albogularis
Also called the Sykes’
monkey, this tree-dwelling RED-TAILED MONKEY
omnivore is distributed Cercopithecus ascanius
in E. and S.E. Africa, An arboreal species of moist forest habitats in
including Zanzibar and C. Africa, the red-tailed monkey has large
the Mafia Islands. cheek pouches, in which it stores fruits.

18–28 in long,
45–70 cm furry tail

161/2–231/2 in
42–60 cm

L’HOEST’S MONKEY
Allochrocebus lhoesti
Also known as the mountain 16–23 in
monkey, this arboreal guenon 40–58 cm
lives in moist primary forests
in the highlands of C. Africa.
DIANA MONKEY
Cercopithecus diana
An inhabitant of the high canopy
of primary forests in W. Africa, the
Diana monkey rarely descends
to the ground. DE BRAZZA’S
151/2–28 in
39–71 cm MONKEY
Cercopithecus neglectus
This is a semiterrestrial
species of C. African swamp
forests. Males are larger
than females and have a
distinctive blue scrotum.
15–25 in
38–63 cm

18–26 in 22–34 in
45–66 cm 55–85 cm

NORTHERN BLACK-
BLUE MONKEY MONA MONKEY GOLDEN-BELLIED MANGABEY CRESTED MANGABEY
Cercopithecus mitis Cercopithecus mona Cercocebus chrysogaster Lophocebus aterrimus
Social groups of this African native A tree-dwelling species, this This mainly terrestrial mangabey lives An arboreal species that prefers
number up to 40 individuals and monkey inhabits rainforest in rainforests and swamp forests in the rainforest, this mangabey is
consist of a dominant alpha male, and mangrove forests from Congo basin. It forages by day in found in the Democratic
several females, and their offspring. Ghana to Cameroon. groups of up to at least 35 individuals. Republic of Congo.
 OL D WOR L D MON K EYS 19–35 in
556 48–88 cm

16–26 in
40–66 cm

12–28 in
30–70 cm

VERVET MONKEY
Chlorocebus pygerythrus 10–18 in
An inhabitant of savanna and open woodland, GRIVET 26–45 cm
this monkey is distributed from Ethiopia Chlorocebus aethiops
through E. Africa to South Africa. A semiterrestrial species, the grivet PATAS ANGOLAN TALAPOIN
lives in N.E. Africa. It is characterized MONKEY Miopithecus talapoin
by a green tinge on its upper face. Erythrocebus patas The smallest of the
speckled The long limbs and short Old World monkeys, the
olive-gray fur digits of this primate make it arboreal talapoin lives in
well-adapted for running. It wet and swampy forests
is found from W. to E. Africa. in W. and C. Africa.
blue flanges
on either side 18–33 in baboon tail typically
of the nose 45–83 cm appears broken

DRILL olive-gray
Mandrillus leucophaeus body
22–43 in A large terrestrial monkey
55–110 cm
of lowland mature
rainforest, the drill is
found only in Cameroon,
MANDRILL Nigeria, and the Republic
Mandrillus sphinx of Equatorial Guinea.
Found in rainforests in
western C. Africa, this
species has distinctive 20–34 in
facial markings that are 51–85 cm
duller in females and
juveniles than in males.

20–43 in
50–114 cm

CHACMA BABOON
Papio ursinus
One of the largest baboons,
this species inhabits
woodland, savanna,
steppe, semidesert,
and montane habitats YELLOW BABOON
across southern Africa. Papio cynocephalus
An opportunistic baboon, this omnivore’s
diet includes seed pods, roots, insects, and other
monkeys. It lives in S. and E. Africa.

red-brown face 20–35 in

50–90 cm

20–37 in
50–95 cm

14–34 in
35–86 cm
HAMADRYAS
BABOON
Papio hamadryas GUINEA BABOON OLIVE BABOON
This species is found Papio papio muscular limbs Papio anubis
in E. and C. Africa, One of the smallest baboons, help to run fast This species forms troops
particularly Ethiopia. this species also has one of the of up to 100 baboons in the
The male has a long smallest ranges, confined to savanna and grassland steppe
silver-gray shoulder cape. western equatorial Africa. of central sub-Saharan Africa.
20 –30 in GOLDEN SNUB-NOSED
50–75 cm MONKEY
Rhinopithecus roxellana 557
This monkey has thick fur,
which helps it to survive in 191/2–30 in
high montane forests in 49–75 cm

M A MM ALS • PRIM ATES

W. and C. China.
mantle of white “cloak”
white hair over back

18 1/2–33 in 24–30 in
47–83 cm 61–76 cm
GELADA
Theropithecus gelada
A grassland grazer from the highlands of
Ethiopia, this primate has a distinctive
bare patch of skin on its chest.

PROBOSCIS MONKEY
Nasalis larvatus
A competent swimmer, this monkey
inhabits mangrove and lowland riverine
rainforest in Borneo. It is named after
the male’s large nose.

18 1/2–27 in
47–68 cm

ANGOLA COLOBUS GUEREZA

Colobus angolensis Colobus guereza
A predominantly arboreal Also called the eastern
species, this monkey inhabits black-and-white colobus, this
a variety of forest habitats species is widespread in the
in Angola, Congo, and other moist, tropical forests of
neighboring countries. C. and E. Africa.

long tail
with a white,
bushy tip

16–31 in
41–78 cm

NORTHERN PLAINS GRAY LANGUR bright orange

Semnopithecus entellus coloration
Also called the Hanuman langur,
this gray monkey is found in S. Asia,
including India and Pakistan.

23–25 in 17–26 in
58–64 cm 43–65 cm

JAVAN LANGUR
Trachypithecus auratus
Most males and females
TUFTED GRAY LANGUR of this species are black
Semnopithecus priam in color. However, some
Found in southeast India and Sri Lanka, individuals retain their
this langur lives in many different habitats. juvenile orange coloration
Its diet consists mainly of tree foliage. into adulthood.
558 MANDRILL
Mand r illu s s phin x
The mandrill is the largest of all monkeys, and the male is particularly
M A MM ALS • PRIM ATES

spectacular. Mandrills live in troops that usually contain a dominant

male, several females, a gaggle of youngsters, and various
non-breeding, lower-ranking males. Sometimes several
groups merge to form troops of 200 or more. Mandrill
society has a strict hierarchy. Individuals advertise their
status with colorful patches of skin on the face and rump.
A dominant male is fearsome-looking, and he has a
temperament to match. The brightness of the skin pigments
is controlled by hormones and color is a good indicator of size 22–43 in (55–110 cm)
strength and ferocity. A rival must be very sure of himself habitat Dense rainforest
before challenging such a magnificent animal, and serious distribution Western Central
fights tend to occur only between well-matched individuals. Africa, from S. Cameroon to
S.W Republic of Congo
diet Mainly fruit

LONG, HARD GLARE NOSTRILS

Forward-facing eyes provide the mandrill with In a mature male, the skin
stereoscopic vision. They see in full color, which around the nostrils and
helps them to locate ripe fruit and to distinguish the center of the snout
the visual signals of other individuals. is scarlet. Females and grooves on either
young mandrills have side of the nose
a black nose.

TEETH
The long canine teeth are used
principally for fighting and in
display. The cheek teeth are
smaller, with knobbly surfaces,
which are used for grinding
up plant material.

GRASP HIND FEET

Mandrill thumbs are short, but fully The hind feet resemble the hands, in that
opposable—like those of the great apes— the toes are long and able to grasp. Mandrills
for grasping and manipulating objects. are good climbers, and often sleep among the
The fingers are long and very branches of trees. PLAIN RUMP
strong, with stout nails. All mandrills have a hairless
short, tufted tail rump and a short tail. The rump
of the subordinate male has less
color than the alpha male.

relatively short
ALL FOURS hind legs
Mandrills spend most
of their time at ground
level, where they move ALPHA MALE
around on all fours, The skin color displayed
and typically roam 3–6 long, by an individual changes
miles (5–10 km) a day. powerful depending on breeding
arms condition and mood.
The alpha male displays
the most vivid red and
blue hues on its
face and rump.
 prominent brow ridges 559
shade the eyes from bright
sunlight

M A MM ALS • PRIM ATES

coat is made up of
long, coarse hairs

POWERFUL
PROFILE
The mandrill’s head is
bulky, accommodating
powerful jaw muscles
needed to crush tough
plant foods. Its ears are
relatively small, though
it has excellent hearing.
Only high-ranking males
have an orange beard.
 GIBBONS AGILE GIBBON
Hylobates agilis
560 The gibbons, or lesser apes, of the family Although its coat
coloration varies, all
Hylobatidae are medium-sized fruit-eating agile gibbons have white
primates. They have no tail and characteristically eyebrows, and males
move by brachiation—swinging between trees also have white cheeks.
using their very long forearms. Every day gibbon This species lives in
Thailand, Indonesia,
families sing to reinforce family and pair bonds and Malaysia.
and announce territorial ownership. Some species black cap
have an enlarged throat sac to amplify the sound. 18–25 in of female
45–64 cm

SILVERY JAVAN PILEATED GIBBON

GIBBON Hylobates pileatus
Hylobates moloch Female pileated gibbons
Endemic to western Java, 17½–25 in are silver-gray with a
Indonesia, males and 44–64 cm black face, chest, and cap;
females of the silvery males are black. They are
gibbon are silver-gray found in Thailand,
with a dark cap. MÜLLER’S GIBBON Cambodia, and Laos.
Hylobates muelleri 17½–25 in
44–64 cm
18–25 in Also known as the gray
45–64 cm gibbon, this species lives
in Borneo. Monogamous
pairs spend an average
of 15 minutes per day
singing duets.

white feet
and hands

32 in
81 cm
WESTERN
HOOLOCK GIBBON
Hoolock hoolock
Male hoolock gibbons are black, 16 ½–23 in
but females are tan with dark 42–59 cm
brown cheeks. They are found
in China, N.E. India,
and N.W. Myanmar.

pronounced hair naked palms

on crown of male
18–25 in
NORTHERN WHITE- 45–64 cm
18–25 in CHEEKED GIBBON
45–64 cm Nomascus leucogenys
Northern white-cheeked
gibbons are creamy
colored when they are
born. They change color
by two years of age. LAR GIBBON
Hylobates lar
Variable in coat coloration, the
white-handed or lar gibbon lives
in forests in Thailand, Malaysia,
Sumatra, Myanmar, and Laos.

silvery white
saddle of hair
28–35in extends to rump
71–90 cm and thighs

SIAMANG
BUFF-CHEEKED Symphalangus
CRESTED GIBBON syndactylus
Nomascus gabriellae The largest of the
Male buff-cheeked gibbons are black gibbons, the siamang lives
with pale cheeks; females are buff on the island of Sumatra,
with a black cap. They are found in Indonesia, and on the
Cambodia, Laos, and Vietnam. Malay Peninsula.
HUMANS AND APES BORNEAN
ORANGUTAN
The family Hominidae contains the largest primates—the great Pongo pygmaeus very long
561
A large arboreal fruit eater, the arms
apes and humans. Orangutans are arboreal, whereas chimpanzees, Bornean orangutan lives in the
gorillas, and humans spend the majority of their time on the canopy of primary rainforest

M A MM ALS • PRIM ATES

ground. Chimpanzees and gorillas move on all fours by “knuckle- on the island of Borneo.
walking.” None of the great apes has a tail. Males are generally
larger than females and all species have a relatively large braincase. 28–38 in
72–97 cm
coarse, shaggy
red-brown coat
EASTERN GORILLA
39 –47 in Gorilla beringei
101–120 cm
The largest primate, the two
eastern gorilla subspecies inhabit
montane cloud forest and grasping
lowland forest in the eastern hands and feet
Democratic Republic of Congo,
Rwanda, and Uganda.
domed
forehead

heavily built
body

2 7–39 in
68–99 cm

SUMATRAN
ORANGUTAN
Pongo abelii
The largest arboreal
primate, the Sumatran
orangutan is restricted
to fragments of primary
tropical forest in
28–33 in N. Sumatra.
70–83 cm

BONOBO
Pan paniscus
Slighter than the
common chimpanzee, 4–7 ft
the bonobo or pygmy 1.2–2.1 m
chimpanzee lives in
humid tropical forest
in the Democratic
Republic of Congo.

WESTERN GORILLA
Gorilla gorilla
Two subspecies of western
gorilla live in lowland
tropical and swamp
forests in western
C. Africa. Adult males
are called silverbacks.

401/2–42 in
103–107 cm

28–38 in
70–96 cm

COMMON HUMAN
CHIMPANZEE Homo sapiens
Pan troglodytes Characterized by a bipedal
Four subspecies of the common posture and a lack of body hair,
chimpanzee are distributed in dry humans permanently inhabit
strong hands and moist forests and savanna every terrestrial habitat, with
and feet woodlands in equatorial Africa. the exception of Antarctica.
562 B AT S
The only mammals capable of powered image of their surroundings from returning PHYLUM CHORDATA
flight, bats are primarily nocturnal. echoes. Those species that emit echolocation calls
M A MM ALS • BATS

CLASS MAMMALIA

Many species use echolocation to through their nose usually have elaborate facial ORDER CHIROPTERA

navigate and find food. ornamentation, called noseleaves, for focusing the FAMILIES 21

sound. Bats have very sensitive hearing, often SPECIES About 1,400

Bats are found worldwide in many different habitats, highly tuned to the frequency of their returning
including tropical, subtropical, and temperate echoes. Some species listen to prey-generated D E B AT E
forests, savanna grasslands, deserts, and wetlands. sounds, such as the rustle that an insect makes E V O L U T I O N A R Y D E B AT E
Most fruit bats, as their name suggests, eat fruits. when walking on a leaf.
All other bats, once referred to as microbats, are Morphological and genetic analyses of
mainly insect eaters. However, some bats drink h a b i t s a n d a da p tat i o n s the relationships between bat families
do not always agree with one another.
nectar and eat pollen, a few suck blood, and some Bats are very social animals, living in colonies of
Genetic studies suggest that all bats
eat vertebrates, such as fishes, frogs, and bats. hundreds or thousands and, exceptionally, millions evolved from one common ancestor
Their greatly elongated arm, hand, and finger of animals. They roost in trees and caves or in and that flight evolved only once.
bones support an elastic wing membrane for flight. buildings, bridges, and mines. Temperate species However, molecular studies, although
Many bats also have a tail membrane between their either migrate to warmer climates or hibernate still proposing two bat groups, also
legs. Bats typically rest upside down, hanging from during winter. They may also become torpid if food indicate that some echolocating bats,
their strong toes and claws. becomes short in other seasons. Many interesting such as the horseshoe bats, are more
reproductive adaptations have developed, including closely related to non-echolocating fruit
t h e s e n s e o f e c h o l o c at i o n the storage of sperm, delayed fertilization, and bats (Yinpterochiroptera) than to other
echolocating microbats (Yangochiroptera).
Fruit bats rely predominantly on vision and smell, delayed implantation to ensure that the young
This is an apparent conundrum unless
whereas microbats use a specialized sense called are born at the optimum time of year. echolocation has evolved twice or has
echolocation to avoid hitting objects, and to detect later been lost in the fruit bats.
prey in the dark. They emit pulses of sound
through their mouth or nose and form a “sound”

F RU I T BAT S
The bats of the family Pteropodidae
are distributed across the tropical and
subtropical regions of the Old World.
They have a doglike face with simple
ears and large eyes. They use vision
and smell to locate their food, except
for members of the genus Rousettus,
which echolocate by producing
tongue clicks. These bats feed on
fruits, nectar, and pollen. They
have claws on both the thumb
and the second finger.

FRANQUET’S
EPAULETTED FRUIT BAT
Epomops franqueti
Also called the singing fruit bat because
2–3 in 4¼–7 in of the high-pitched call of the males, this
5–7.5 cm 11–18 cm species occurs in W. and C. Africa.

1½–3¼ in 3¼–4½ in
4–8 cm 8–11 cm

LESSER
BLOSSOM BAT LONG-TONGUED SHORT-NOSED
Syconycteris australis FRUIT BAT FRUIT BAT
This specialized nectar-feeder Macroglossus minimus Cynopterus sphinx
occurs from Papua New Occurring in S.E. Asia, This is the only fruit bat
Guinea to eastern coastal this fruit bat feeds on that makes tents from
Australia. It has a pointed nectar and pollen from palm leaves. It is found
muzzle and a brush-tipped elastic skin flowers, using its across S.E. Asia and the
tongue for probing flowers. membrane long tongue. Indian subcontinent.
 EGYPTIAN ROUSETTE hind limbs have
Rousettus aegyptiacus hooklike claws
Capable of echolocating by tongue for roosting 563
clicks, this species occurs across
Africa and the Middle East, except 7–8½ in 4–7½ in
in the Sahara Desert. 18–21 cm 10–19 cm

5–8 in
13–20 cm

4¾–6½ in
12–16 cm

HAMMER-HEADED GEOFFROY’S MOLUCCAN WAHLBERG’S

FRUIT BAT ROUSETTE NAKED-BACKED EPAULETTED
Hypsignathus monstrosus Rousettus amplexicaudatus FRUIT BAT FRUIT BAT
Males of this species are Like other Rousettus species, Dobsonia moluccensis Epomophorus wahlbergi
much larger than the females this S.E. Asian bat eats Widespread in the Found in forests and savanna
and have a greatly enlarged fruits and nectar. It roosts Moluccan Islands, this across sub-Saharan Africa, this
snout. This bat is found in together with thousands bat is rare in the far species has white patches on
W. and C. Africa. of others in caves. north of Australia. its shoulders and eyebrows.

6½–12 in 5–8 in 6–8 in 6½–9½ in 5½–9 in

16–30 cm 13–20 cm 15–20 cm 16–24 cm 14–22 cm

large eyes
AFRICAN STRAW- assist in
LITTLE RED COLORED FRUIT navigation
FLYING FOX BAT
Pteropus scapulatus Eidolon helvum
This nomadic species from Colonies of this fruit bat
Australia primarily feeds can contain 1 million
on eucalyptus blossoms. individuals. This migratory
It is occasionally found species is widespread in
in Papua New Guinea. sub-Saharan Africa.

outstretched
fingers thick leg fur
down to ankle

RODRIGUES LYLE’S FLYING FOX

FLYING FOX Pteropus lylei
Pteropus rodricensis Occurring in Cambodia,
An inhabitant of mangrove Thailand, and Vietnam,
and rainforest, this flying the Lyle’s flying fox can
fox is found only on severely damage trees
Rodrigues Island in by stripping them
the Indian Ocean. of their leaves.

9–12 in
23–30 cm
9–11½ in
22–29 cm

9–10 in 7–11 in
22–25 cm 18–28 cm

9–11½ in
23–29 cm

SPECTACLED GRAY-HEADED
FLYING FOX FLYING FOX
LARGE FLYING FOX INDIAN FLYING FOX BLACK FLYING FOX Pteropus conspicillatus Pteropus poliocephalus
Pteropus vampyrus Pteropus medius Pteropus alecto This fruit bat inhabits The largest bat in Australia,
The largest of all bats, this Found throughout India and The black flying fox has a primary and secondary tropical this flying fox lives in
flying fox is found across parts of S.E. Asia, this species wingspan of more than three rainforests in Indonesia’s Moluccan communal roosts, known
the mainland and islands roosts in large colonies in feet. It lives in Indonesia, Islands, New Guinea, and as camps, in rainforest
of S.E. Asia. forests and swamps. New Guinea, and N. Australia. N.E. Queensland, Australia. and woodland.
564 LY L E ’ S
F LY I N G F OX
P t eropu s lylei
M A MM ALS • BATS

Lyle’s flying fox is a medium-sized

representative of the Old World fruit bat
family, the Pteropodidae. Fruit bats are social
animals; many hundreds may gather in roost
trees during the day to rest and groom, dispersing
at dusk to find ripe fruits. Although they may cause
some damage to trees, many fruit bats are important
pollinators and seed dispersers of tropical plants, including
many commercial crops. Fruit bats are found in the
size 6½–9½ in (16–24 cm)
tropical regions of Africa, Asia, and Australia, although this habitat Forests
species is only found in Cambodia, Thailand, and Vietnam. They distribution S.E. and E. Asia
inhabit forested areas, including mangroves and fruit orchards. diet Fruits and leaves

all bats have a thumb claw,

CANINE FACE but only Old World fruit
Fruit bats have large eyes for navigation and a bats have a claw on the
large nose for sniffing out fruits, pollen, and second finger, too
nectar—giving them a doglike face. Species that
use echolocation have larger ears and smaller eyes.

CLINGING CLAWS
Bats’ sharp, curved claws
are perfect for clinging
to tree branches. When
roosting, tendons lock in
place to ensure the bat’s
claws stay bent without the
muscles needing to contract.

HAND WING
A bat’s wing is formed of elongated forearm
and finger bones supporting a thin elastic
membrane attached to the sides of the body.
It has a large surface area, giving lift in flight.

TAIL OR NO TAIL? WELL WRAPPED

Pteropus bats lack a tail, but some When resting, most fruit bats
have a partial membrane supported hang upside down with their
by cartilaginous spurs called calcars, leathery wings folded around
which protrude from the ankle. their bodies. Bats roosting in
the open risk overheating, so
they flap their wings and cover
WALKING UPSIDE DOWN themselves in saliva to help
Fruit bats use their large clawed keep cool in hot weather.
thumbs to help them move along
the branches of their roost tree.
The claw can also be used to
manipulate fruit when feeding.
 reddish brown,
foxy coloring

HANGING AROUND
Few bats are able to move
well on the ground, let
alone take off from a flat
surface. By hanging upside
down they are able to
take flight quickly. During
the day, they hang upside
down to sleep, nestled
together for warmth.

large eyes are adapted upright, foxlike ears detect

for excellent vision, sounds beyond the range of
especially at night human hearing
 HOR SE SHOE BAT S
566 Species of the family Rhinolophidae are
distributed throughout southern Europe,
Africa, Asia, and Australasia. Their noseleaf
M A MM ALS • BATS

is characteristically horseshoe-shaped. They nose leaf

have the most sophisticated echolocation
of all bats, involving highly specialized
sound transmission and reception.
1 1/2–13/4 in
LESSER HORSESHOE BAT 3.5–4.5 cm
Rhinolophus hipposideros
Found across Europe, N. Africa,
and W. Asia, the lesser
horseshoe bat is one of the
smallest bats in the world.

elongated
finger bone relatively short,
broad wings

21/4–2 3/4 in
5.5–7 cm

MEHELY’S GREATER
HORSESHOE BAT HORSESHOE BAT
Rhinolophus mehelyi Rhinolophus ferrumequinum
A cave-dwelling, medium-sized The largest of the European
rhinolophid, this bat is patchily horseshoe bats, this species’ range
11/2–2 1/2 in distributed across S. and E. Europe, extends from Europe eastward
4–6.5 cm and the Middle East. across Asia to Japan.

L E A F -NOSE D BATS M OU S E -TA I L E D BAT S

Found throughout much of Africa, Asia, and The tail of these bats is nearly as long as their
Australasia, the Hipposideridae contains the body and is a distinguishing feature of the family
Old World leaf-nosed bats. They have complex Rhinopomatidae. Mouse-tailed bats also have
noseleaves and, as with horseshoe bats, their a fleshy muzzle, and their large, simple
hind legs are poorly developed, making ears join at the base.
them unable to move quadrupedally.

COMMERSON’S 4 –4 1/2 in
LEAF-NOSED BAT 10–11 cm
11/2–2 1/4 in Hipposideros commersoni
4–6 cm This forest-dwelling bat roosts in
hollow trees across the island of 2–3 1/2 in
Madagascar. It is one of the largest 5–9 cm
leaf-nosed bats, weighing up to
61/4 oz (180 g).

MOUSE-TAILED BAT
Rhinopoma sp.
SUNDEVALL’S Six species of fast-flying,
LEAF-NOSED BAT insectivorous mouse-tailed
Hipposideros caffer bat live in arid and semiarid
A savanna species, this regions of N. Africa, the
leaf-nosed bat roosts in caves Middle East, and India.
and buildings throughout
Africa except in the Sahara first finger
and central forested regions. (thumb)

HOG -NOSE D BAT M EGA DE R M AT I DA E

The sole member of the family Craseonycteridae, the The family Megadermatidae comprises six fairly
tiny hog-nosed bat has long, broad wings that allow it large echolocating bat species. Carnivorous and
to hover. It lacks a tail and calcars (cartilaginous insectivorous, they have large ears and eyes
extensions of the ankle). and a broad tail membrane but little or no tail.
4–5 in
10–13 cm
1–11/4 in KITTI’S HOG-NOSED BAT
3–3.5 cm Craseonycteris thonglongyai GHOST BAT
One of the world’s smallest mammals, Macroderma gigas
and often called the bumblebee bat, Endemic to N. Australia, the ghost bat is one of
Kitti’s hog-nosed bat lives in riverside the largest microbats. It preys on vertebrates,
caves in Thailand and Myanmar. such as frogs and lizards.
 S H E AT H -TA I L E D
BATS 567
The Emballonuridae are more commonly 1 1/2–2 in
known as sheath-tailed bats because just 3.5–5 cm

M A MM ALS • BATS
the tip of their tail protrudes through
the tail membrane, giving it the
appearance of being sheathed.
Many have glandular odor- 2–21/4 in
storing sacs on their wings. 4.5–6 cm

3–31/4 in 1 1/2–2 in
7.5–8.5 cm 4–5 cm

HILDEGARDE’S LESSER SHEATH-

TOMB BAT TAILED BAT
Taphozous hildegardeae Emballonura monticola PROBOSCIS BAT GREATER SAC-WINGED BAT
Dependent on caves for The short tail of this bat Rhynchonycteris naso Saccopteryx bilineata
roosting, Hildegarde’s appears to retract into a sheath These bats spend the day roosting Male greater sac-winged bats attract
tomb bat feeds on insects when it stretches its legs. It in groups on the underside of females by a pungent secretion in
in the coastal forests of lives in Indonesia, Malaysia, branches. They live in tropical their wing sacs. The species is found
Kenya and Tanzania. Myanmar, and Thailand. forests in C. and S. America. in C. and S. America.

N E W WOR L D
L E A F -NOSE D BATS 2–23/4 in
4.5–7 cm
The Phyllostomidae contains species that are
distributed from southwestern US to northern
Argentina. Most have large ears and a noseleaf,
shaped like a spearhead, to enhance echolocation.
2 1/2– 4 in 2 1/4–3 in
6.5–10 cm 5.5–7.5 cm

2–21/4 in
4.5–6 cm

GERVAIS’ SEBA’S SHORT-TAILED BAT

FRUIT-EATING BAT Carollia perspicillata
Artibeus cinereus Inhabiting moist evergreen and dry
Preferring to roost in palm deciduous forests across much of
trees, Gervais’ fruit-eating C. and S. America, this species is
bat lives in S. America, a generalist fruit eater.
including Venezuela, Brazil,
COMMON and the Guianas.
TENT-MAKING BAT
PALE SPEAR-NOSED BAT Uroderma bilobatum
Phyllostomus discolor Found in lowland forests
This species emits echolocation from Mexico to central
calls through its nose. It is S. America, this bat creates
found in C. and northern shelters by biting palm and
S. America. banana leaves.
31/4–4 1/4 in 2–21/4 in
8–10.5 cm 4.5–6 cm

GEOFFROY’S SILKY
TAILLESS BAT SHORT-TAILED BAT
Anoura geoffroyi Carollia brevicaudum
A specialized nectar feeder, This leaf-nosed bat is widespread
2 1/4–3 in
this species has a long muzzle, in C. America and the Amazon
6–7.5 cm elongate cheek teeth, and a region. It helps to restore
brush-tipped tongue. It lives degraded forests by dispersing
in C. and S. America. the seeds of fruit trees.

2 3/4–3 3/4 in
7–9.5 cm COMMON VAMPIRE BAT
2 3/4–3 3/4 in 21/4–31/4 in
Desmodus rotundus
7–9.5 cm 6–8 cm CALIFORNIAN Famous for feeding on other
LEAF-NOSED BAT mammals’ blood, the common
FRINGE-LIPPED BAT WHITE-LINED Macrotus californicus vampire bat occurs in a variety
Trachops cirrhosus BROAD-NOSED BAT Primarily hunting for moths of habitats in Mexico and
The fringe-lipped bat catches Platyrrhinus lineatus by sight rather than C. and S. America.
frogs by listening for their Named for the white lines on its echolocation, the Californian
calls. It lives in tropical face and back, this broad-nosed leaf-nosed bat lives
forests in C. and bat roosts in damp forests of in N. Mexico and
northern S. America. central S. America. the S.E. US.
 MOR MOOPIDA E NOCTILIONIDA E NATA L I DA E
568 The family Mormoopidae contains the naked- The two species of bulldog bats from the family The funnel-eared bats of the family Natalidae
backed and moustache bats. Several species have Noctilionidae have long legs and large feet and are small and slender with large ears. Adult
wings that join across their back and they have claws. They have full lips and cheek pouches for males have a sensory structure, called a natalid
M A MM ALS • BATS

a fringe of stiff hairs around the muzzle. storing food while flying. organ, on their forehead.

11/2–13/4 in
3.8–4.3 cm

31/4–4 in MEXICAN FUNNEL-

8–10 cm EARED BAT
DAVY’S NAKED- Natalus mexicanus
BACKED BAT An insectivorous species,
Pteronotus davyi GREATER BULLDOG BAT the Mexican funnel-eared
2–21/4 in Noctilio leporinus
4.5–6 cm Unlike most other bats, this bat lives in C. and S.
bat’s wing membranes meet Specialized for snatching fishes from the water’s America. It roosts in caves
across its back. It is distributed surface with its claws, the greater bulldog or and, sometimes, in
from Mexico to S. America. fishing bat lives in tropical C. and S. America. abandoned mines.

S L I T- FAC E D M YSTAC I N I DA E T H Y ROP T E R I DA E

BATS The one extant member of the Mystacinidae The family Thyropteridae contains five species
The slit-faced bats of the family has extra talons on its thumb and toe claws. of disk-winged bats. Suction cups on the wrists
Nycteridae have a furrow from Its tough, leathery wings can be folded against and ankles allow them to attach to smooth-
their nostrils to a pit between the body when moving on the ground. surfaced tropical leaves when roosting.
their eyes. The cartilage at the
end of their tail ends in a
“Y” shape. 21/4–31/4 in 11/2–2 in
6–8 cm 4–5 cm
2 1/2 –3 in
6.5–7.5 cm

SPIX’S DISK-
WINGED BAT
Thyroptera tricolor
An insectivorous
MALAYAN SLIT-FACED BAT LESSER NEW ZEALAND inhabitant of lowland
Nycteris tragata SHORT-TAILED BAT forests from S.
The facial furrow of this slit-faced bat Mystacina tuberculata Mexico to S.E. Brazil,
helps to direct its echolocation calls. It An agile mover on the ground, the Spix’s disk-winged