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classification and
nomenclature
an introduction —
Marc S.M. Sosef
Jérôme Degreef
Henry Engledow
Pierre Meerts
Botanical
classification and
nomenclature
an introduction —
Marc S.M. Sosef
Jérôme Degreef
Henry Engledow
Pierre Meerts
by Marc S.M. Sosef1, Jérôme Degreef1,2, Henry Engledow1 & Pierre Meerts3
1
Meise Botanic Garden, Nieuwelaan 38, B-1860 Meise, Belgium
2
Service Général de l’Enseignement supérieur et de la Recherche scientifique,
Fédération Wallonie-Bruxelles, Rue A. Lavallée 1, B-1080 Brussels, Belgium
3
Herbarium et bibliothèque de botanique africaine, Université Libre de
Bruxelles, Av. F.D. Roosevelt 50, CP 265, B-1050 Brussels, Belgium
Copyright © 2020 Meise Botanic Garden, Nieuwelaan 38, 1860 Meise, Belgium.
Printed in Belgium by Gewadrupo, Arendonk.
This publication is published and distributed in Open Access under the Creative
Commons Attribution 4.0 International license (CC-BY 4.0), which permits
use, distribution, and reproduction in any medium, provided the original work
is properly cited. A PDF file of this publication can be ordered, free of charge
(send an email to webshop@plantentuinmeise.be), or downloaded from the
webshop of Meise Botanic Garden at http://shopbotanicgarden.weezbe.com.
DOI: 10.5281/zenodo.3706707
ISBN 9789492663207
Subject: Botany
D/2020/0325/002
Content
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
2. Species concepts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
2.1 What is a species? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
2.2 Speciation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
2.3 Infraspecific taxa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Introduction
5
6
Biology is the science that explores the living world around us. To
communicate the wonders of nature, names are needed to describe the
variety of forms we encounter. This wildly diverse nature may be represented
through a hierarchical structure where names are used to indicate groups
of organisms at different levels. The classification and naming of organisms is
an essential tool for scientific communication. It forms the foundation upon
which biological research is based and the discipline is called “Taxonomy”.
Taxonomists explore, describe, name, and classify all living organisms on Earth.
Although the information is general, most examples are drawn from tropical
African plant and fungal diversity. Each chapter is followed by an overview of
literature and web-based sources related to the subjects dealt with. This is by
no means exhaustive, and again focusses on the taxonomy of African plants
and fungi.
The authors hope that this publication will contribute to the development 7
of taxonomic expertise, notably in the Central African region. The booklet is
produced in English and French, and will be available free of charge (under the
CC-BY license) to high schools and universities (both teachers and students),
courtesy of Meise Botanic Garden.
8
1.
The history of
classification
9
Any biological study starts with the simple question “What is this?”. Whether
it concerns a manager of a nature reserve who needs to know which species
grow within the park’s boundaries for setting up a management plan; a
primatologist studying the food eaten by chimpanzees; or a plant breeder
studying the close wild relatives of the potato in search of a disease resistant
gene, all need to be able to identify and name their material. It is preferable
that these names are uniformly used and accepted throughout the world.
The need for a uniform system of naming the living world was already
recognized by the ancient Greeks and Romans. Names were given to ‘entities’
we now call species that had specific morphological characters and uses.
Some produced, for example, edible fruits, or a yellow dye, others had
medicinal properties, or were useful for making musical instruments, etc.
In this chapter (largely based on Magnin-Gonze 2009 and Rouhan & Gaudeul
2014), we highlight the major historical phases in the development of plant
naming and classification. The name for this field of science was coined for
the first time in 1813 by the Swiss botanist Augustin Pyramus De Candolle
(1778–1841) in his book “Théorie élémentaire de la Botanique”. He created
the neologism “taxonomy” by combining the words Greek ταξις (order) and
νόμος (law, rule).
Even before the invention of written language, c. 5600 years ago, it is likely
that an oral plant classification system existed. Initially, names and organisms
were not placed into a hierarchical system since the plants were all named
following their use such as food, medicines, poisons, or materials (Raven 2004).
The Greeks probably did not just consider plants as being only useful but also
as beautiful; the murals in Knossos (1900 BC) not only have useful plants like
barley, fig, and olive, but also narcissus, roses, and lilies.The Greek Theophrastus
(372–287 BC; figure 1), successor of the great philosopher Aristotle, is espe-
cially well known as the first true botanist. Interested in naming plants and
finding an order in the diversity of plants, he is the first one to provide us
with a philosophical overview of plants. He pointed out some of the impor-
tant questions that would later define taxonomy, such as “What have we
got?” or “How do we differentiate between these things?” Moreover, he was
the first to discuss relationships among plant species and to suggest ways to
group them. Theophrastus described ca. 500 plants — probably representing
all known plants at that time – and classified them as trees, shrubs, subshrubs,
and herbs. He also made a distinction between flowering and non-flowering
plants, deciduous and evergreen trees, and be-
Figure 1. Statue of tween terrestrial and aquatic plants. Even if 80% 11
Theophrastus in the of the plants included in his works were cultivated,
botanic garden at he had realized that “most of the wild kinds have
Palermo, Italy. no names, and few know about them,” highlighting
the need to recognize, describe, and name plants
Figure 2.
Plinius the Elder.
Figure 3.
Dioscorides.
The Renaissance (late 14th to 17th century) marked a new era for science.
Europeans were exploring and discovering America, Africa, Asia and Australia,
bringing back many unknown plants to Europe. These were housed in a
rapidly increasing number of gardens, the first being created in the early
16th century in Italy. At first, these were called medicinal gardens, later, when
the interest shifted towards the study of plants themselves rather than their
useful properties, they transformed into the botanical gardens that we
know today. Moreover, with the invention of the
Figure 4. Page from printing press (1450-1455), information was more
Dioscorides’ Materia easily shared and distributed, boosting exchange
Medica showing and discussions in scientific knowledge. People
Cassia fistula. became curious about the world surrounding
them. Around 1530, in the botanic garden of Pisa,
13
14
the Italian Luca Ghini (1490–1556) invented a revolutionary method for
preserving plants by drying and pressing them so they could be studied at any
time of the year. The resulting plant specimens were stored in books known
as a “hortus siccus” (dried garden), later the term “herbarium” was adopted,
and were valuable possessions afforded only by the wealthy (Ghorbanie et
al. 2018; figure 5).
This was followed by the era of the great western European herbals, or
books describing the plants and their uses. These works were no longer
produced only in Latin (the scientific language of the time), but also in the
common languages German, English, Dutch and French. This opened up the
information on plants to an even wider public. From this period, the herbals
of Dodoneus (Cruydeboeck, 1554; figure 6), Fuchs
Figure 5. Frontpiece of (New Kreüterbuch, 1543) and Gerard (Herball, or
the Rauwolf herbarium Generall Historie of Plantes, 1597) are the most
1573-1575 kept at famous. Advances in art led to numerous new
Naturalis Biodiversity plant illustrations. These were way superior to
Center, Leiden. those copied over and over from the books of
Dioscorides and Plinius, from which the actual
Figure 6. Dodonaeus
species could often hardly be deduced.
and two pages from his
famous illustrated herbal A student of Ghini, Andrea Cesalpino (1519–
(Cruydeboeck) printed 1603), was the first to discuss the work of
in 1554. Theophrastus since the Ancient Greeks. He
pointed out that plants should be classified in a
15
more natural and rational way. His De Plantis Figure 7.
In the first half of the 18th century, the bright young Swedish botanist, Carl
von Linné (figure 7), or Linnaeus in Latin, brought order where there was
chaos. While working in The Netherlands, he met famous professors such as
Hermann Boerhaave, Adriaan van Royen and Johannes Burmann with whom
he discussed several of his new ideas.
17
composed of only two words. It was the start of the binomial (two words)
nomenclature we still use, a system where a species name is composed of
the genus name followed by a word indicating the species, called the epithet.
Soon, other botanists appreciated the simplicity and genius of the new
naming system and adopted it in their own work. Shortly after the success of
his Species plantarum, Linnaeus, being a keen zoologist as well, introduced the
same system for animals in his famous Systema naturae (1758).
Linnaeus travelled to England to meet Sir Hans Sloane and Johann Jacob
Dillenius, who were both sceptical about his new naming and classification
ideas at first, but they came around after several years. In Paris, he met
Bernard de Jussieu, who would together with his nephew Antoine Laurent de
Jussieu publish their Genera plantarum. In that work, they stated that a species,
genus, or any other class in the hierarchical classification, which we now call a
taxon (plural taxa), should group plants showing character constancy within
the given taxon, as opposed to the character variability observed among taxa.
Since not all characters are useful at the same level of the classification, their
principle of subordination led to a character hierarchy: characters displaying
high variability should be given less weight than more conserved ones in plant
classifications.
In this period, classification and the study of nature also had a religious
implication. Biologists were seen as scientists studying the living things which
God has created and placed on Earth. Linnaeus, in his Introduction to his
Species plantarum, wrote: “In his omnipotent omniscience, God created
the theatre of all living beings on earth, and it is our divine task to explore
that great creation, served to us as tasty treats, unworthy as we are, and to
recognize His hand in it” [freely translated from the Latin]. One can imagine
that in this context Darwin’s introduction of the novel idea that species were
not created by God Almighty, but had evolved from others over a very long
period of time, had a tremendous impact on broader society.
At the start of the 19th century, new questions arose in the mind of taxonomists.
They were not only interested in naming, describing, and classifying organisms,
but also in the origin of the observed diversity. In 1809, in his Philosophie
zoologique, the zoologist Jean-Baptiste de Lamarck proposed a theory where
species could evolve and change through time.
It took another 50 years before Charles Darwin (1809–1882; figure 10) pub-
lished his famous theory of evolution and survival of the fittest in On the
Origin of Species (1859). Independently, Alfred Russel Wallace (1823–1913)
18 had come to the same conclusion while working in Asia. [In fact, the theory
was published already in 1858, in a paper authored by Darwin and Wallace
in the Journal of the Proceedings of the Linnean Society: Zoology.] Darwin
introduced the central concept of descent with modification that later re-
ceived extensive support and is still generally accepted today. The concept
Figure 10.
Charles Darwin.
Figure 11. Example In the early 1960s, a new technique called ‘nu-
of a phenogram, with merical taxonomy’ arose to produce a tree-like
a measure of similarity output, or phenogram (figure 11), on which one
(Manhattan Distance) could subsequently base a classification. Notably
along the x-axis (from the work of Sokal & Sneath (1963, and later edi-
Pometti et al. 2007). tions), Principles of numerical taxonomy, laid the
19
foundation. The technique, also called phenetics, was based on a quantitative
cluster analysis of overall similarities between taxa, using a characters-by-taxa
data matrix - with a mixture of binary characters (stipules present, yes/no),
multi-state characters (for example flower colour, with states 1=white, 2=yel-
low, 3=blue), or continuous characters (for example calyx length in mm)
- and resulting in pairwise distances among the individuals or taxa, called
OTUs (Operational Taxonomic Units). However, it was soon realized that
overall similarity does not necessarily indicate
an evolutionary relationship. For example, spe-
cies may have developed similar features be-
cause they adapted to the same environmental
stress. As this method was not based on the
evolution theory, it could not interpret the
observed variation in an evolutionary sense
with respect to ancestors and descendants or
observed character state changes. Despite the
fact it produces tree-like phenograms, these do
not represent a natural and evolutionary clas-
sification. Nevertheless, this theory flourished
for a while, greatly benefiting from rapid ad-
Figure 12.
vances in informatics.
Willi Hennig.
It was the German zoologist Willi Hennig
Figure 13. Example
(1913–1976; figure 12) who fundamentally
of a cladogram, showing
changed the way biologists reconstruct the
numbered characters
evolutionary pathway of a taxonomic group.
where one may recognize
In 1960 he published his cladistic theory in
apomorphies (black dots),
Grundzüge einer Theorie der Phylogenetischen
parallellisms (open dots)
Systematik, but it remained relatively unknown
and reversals (hatched
until the English translation Phylogenetic Sys-
dots) as well as their
tematics was published in 1966. The primary
character state changes
principle is not to use the overall similarity
(below each dot).
20
among taxa to reconstruct the phylogeny, but to make a distinction between
primitive character states and those that are derived from them. Only de-
rived character states, called apomorphies, shared by several taxa then indi-
cate a shared common ancestry, while primitive ones, called plesiomorphies,
do not. A group derived from a single common ancestor is called a clade and
the theory behind it cladistics. The result of a cladistic analysis is a tree-like
figure called a cladogram (figure 13), where the branches actually represent
one or more character state changes. When, for example, in a group of plants
with red flowers an evolutionary change gave rise to blue flowers, ‘blue’ then
is the derived state of the character ‘flower colour’ and any species having
that state is likely to have evolved from the same common ancestor. Having a
red flower does not indicate such common ancestry and hence cannot serve
as a criterion on which to base a taxonomic group. When, further down the
evolutionary history it turns out that a red colour was derived from white
flowers, red can still be regarded as a derived character state but at a differ-
ent level in the phylogeny. And going from white to blue then requires two
evolutionary steps rather than one. When a derived state evolves back to the
primitive state again, this is called a reversal, while the independent evolve-
ment of the same character state in two or more different branches of the
evolutionary tree is called a parallelism.
In this new setting, it was felt that a new definition of this biological research
field was needed and the term ‘systematic biology’ or simply ‘systematics’ was
coined (Michener et al. 1970). It embraced the entire field from describing,
naming, classifying, studying the distribution patterns (biogeography),
evolutionary relationships, character evolution and adaptations. The term
‘taxonomy’ was then restricted to describing, naming and classification. Some,
however, treat these two words as synonymous.
The discovery of the double helical structure of the DNA molecule in 1953,
by James Watson and Francis Crick, greatly improved our understanding of
the evolutionary processes. But it was only after it became possible to tar-
get specific fragments of the genome (nuclear, mitochondrial or chloroplast 21
DNA) by selectively amplifying the DNA through the polymerase chain reac-
tion (PCR) (Karry Mullis 1986) that it started to have a dramatic impact on
taxonomy and classification. The introduction of DNA sequence data (Meier
2008) offered access to numerous new characters and statistical approaches.
Thus, at the turn of the 21st century, the use of molecular data and new tree
building algorithms such as Maximum Likelihood and Bayesian statistics led
to a distinct improvement of our abilities to produce phylogenetic hypothe-
ses. The ‘strength’ or reliability of each branch in a cladogram can be assessed
by using other techniques such as bootstrapping (a statistical resampling
technique; Holmes 2003) and, again, Bayesian statistics. All these develop-
ments led to improved insight into the delimitation of orders and families of
flowering plants (Angiosperm Phylogeny Group 2016), as well as a greater
understanding of the classification based on evolutionary relationships.
It logically follows from the above that classifying the natural world into spe-
cies, genera and higher groups has become a search for our best hypothesis
on the structure of the evolutionary tree in order to be able to distinguish
natural groups. In other words, the cladogram produced by one of various
analyses needs to be split into natural parts. However, there are many ways
to do this, and requires informed choices.
The majority of taxonomists will tell you that a classification can only be
natural when it is composed exclusively of so-called monophyletic units;
that is a group of species that includes an ancestral species and all members
derived from that one ancestral species (figure 15), also named the most
recent common ancestor (MRCA). [Note: a single species can represent a
‘group’ comprising a single element.] When some, but not all, of the species
derived from the most recent common ancestor are included, that group is
called paraphyletic (figure 15). The problem is that while mathematically a
cladogram, with taxa represented only at its tips (see figure 13), might be fully
chopped up into monophyletic groups (the nodes of the cladogram are said
After having decided which rules one wants to follow to chop up a phyloge-
netic tree (or a cladogram) into taxa, there are still many choices one can
make that renders the process of classification and naming of taxa partly
subjective. “Which part of the tree will I recognize as a genus?” “Or, would it
be better to call it a subgenus?” etc. are true questions one needs to answer.
Making choices that will cause least perturbation in the existing system is also
a valid argument thereby promoting name stability.
• Stace C.A. (1991) Plant Taxonomy and Biosystematics, 2nd ed.. London,
Edward Arnold. ISBN 071.3129557
• Stuessy T.F., Crawford D.J., Soltis D.E., Soltis P.L. (2015) Plant
Systematics. The Origin, Interpretation, and Ordering of Plant
Biodiversity. ISBN 978-3874294522
2.
Species
concepts
25
2.1 What is a species?
Biologists generally agree that the species is a fundamental natural unit. How-
ever, it has proven incredibly difficult to define what exactly a species is! This
controversy occurs notably on a theoretical rather than a practical level and
has come to be known as ‘the species problem’.
One of the most fundamental aspects of the problem is variation. Most, if
not all, animal and plant species show variation, every individual often be-
ing demonstrably unique. Within a population variation can be continuous
(e.g. height or weight) or discontinuous (e.g. sex; having right- or left-spiralled
corolla lobes), environmental in origin (e.g. flower colour influenced by the
composition of the soil) or genetic (e.g. blood type). Variation can also oc-
cur in space between populations (geographical variation). Even when two
individuals share exactly the same DNA (clones or twins) they may develop
morphological differences under the influence of environment factors; this
is called phenotypic plasticity. The species problem is, in part, a history of
how biologists have tried to address variation. Often, species are thought to
represent a natural unit. The most extreme opposing view to this idea, states
that only individuals exist in nature. Taxonomic groups, including species, are
then seen as man-made abstractions that allow us to conveniently group
large numbers of individuals. Few scientists accept this nominalist approach
with respect to species, but many believe it does apply to higher taxa (World
Conservation Monitoring Centre 1992).
Multiple definitions and species concepts have been proposed. These usually
follow the discipline of the author: the taxonomic species concept, the evolu-
tionary species concept, the ecological species concept, the historical species
concept, and many more. Species concepts can be divided into two main
groups, those concerned with process (evolution, interbreeding) and those
concerned with pattern (morphology, ecological preferences). Below are the
three most widely known ones:
The biological species concept. This concept defines species in terms
of interbreeding. Its biggest advocate was undoubtedly Ernst Mayr, an
ornithologist. He defined species as “groups of interbreeding natural populations
that are reproductively isolated from other such groups”. Later, it was refined
to “a population or group of populations whose members have the potential to
interbreed in nature and produce viable, fertile offspring, but do not produce viable,
fertile offspring with members of other such groups”. It remains the most widely
accepted species concept today. It explains why the members of a species
resemble one another, and differ from other species. The members exchange
genetic material and pass it on to their offspring, but not to other species.
Thus, the evolutionary process involves random mutations that remain inside
a gene pool that has acquired some form of isolation. Over time these
novelties will start to differentiate these populations from other similar gene
pools (or populations). Finally, these differences may lead to reproductive
27
isolation, where the isolated gene pools will start acting as species.
In general, zoologists embrace this species concept, however it poses some
complications for plants. While hybrids in animals are rare, in plants many spe-
cies are known to hybridize and produce fertile offspring (Grant 1981, Stace
et al. 2015). Only if such events are rare and the offspring are less viable, can
the parent species maintain their unique genetic identity and hence be recog-
nized as distinct species. Furthermore, the concept does not apply to asexual
organisms, and in plants for example the occurrence of apomixis would not
allow defining a species according to the biological species concept.
The morphological species concept. This concept characterizes a species by
morphological distinctiveness and is applied to both asexual and sexual or-
ganisms. It can be applied when information on gene flow is unknown e.g.
when only herbarium specimens are available. Researchers may disagree on
which characters to use to differentiate species which leads to subjectivity.
The evolutionary species concept. This concept stresses the importance of a
species being an evolutionary unit. It defines them as “a lineage of interbreed-
ing organisms, reproductively isolated from other lineages, that has a beginning,
an end, and a distinct evolutionary trajectory and historical fate” (Wiley 1978). It
is definitely the least practical concept, but does include time as an essential
element.
Whatever concept a scientist uses to distinguish a species, the delimitation
actually represents a hypothesis about the relationships among the individual
organisms belonging to the species. Such a hypothesis about which group of
individuals forms a species may be tested using morphological, genetic, behav-
ioural or other types of evidence.
2.2 Speciation
Species concepts
Rules of botanical
nomenclature
31
32
Once the character variation within a group has been studied and conclusions
have been drawn about which entities, or taxa, need to be distinguished, the
question arises as to what the correct names for these entities should be.
That is where one enters the realm of botanical nomenclature.
After 1753, when Linnaeus introduced his binomial system, only some ele-
mentary rules for naming plants were developed. Later, in 1813, Augustin de
Candolle in his Théorie élémentaire de la Botanique provided a detailed set of
rules regarding plant nomenclature. However, over time it became apparent
that an internationally recognized and accepted system and rules for nam-
ing plants was necessary. It was Alphonse de Candolle, son of Augustin de
Candolle, who convened an assembly of botanists from several countries
to present a new set of nomenclatural rules. In 1867, he organized the First
International Botanical Congress (IBC) in Paris, which led to the publication of
the so-called Paris Code. Subsequent meetings of the IBC were held in 1892
(Rochester Code), 1905 (Vienna Code), 1907 (American Code) and 1912
(Brussels Code). A general agreement regarding internationally acceptable
rules for plant nomenclature was however only reached in 1930 at the IBC
meeting in Cambridge. Here, for the first time in botanical history, a Code
of nomenclature came into being that was both international in function and
name: the International Code of Botanical Nomenclature (ICBN). Today,
it is composed of a number of Principles, Rules and Recommendations laid
out in 61 Articles, as well as the Provisions for governance of the Code. It
looks quite similar to a book of law. Since 1930 many updates of the ICBN
have been produced. In 2011, the name changed to “International Code of
Nomenclature for algae, fungi, and plants (ICN)”. It also covers the fossils of
these groups (see Turland et al. 2018).
Proposals to amend the Code are published in the journal Taxon. Every 6
years, at the start of the International Botanical Congress, during the ‘Nomen-
clatural Session’ that may take a full week, taxonomists from all over the world
meet to discuss all proposals published during the interim period between
conferences. Each institute has a number of votes, depending on their number
of research staff. Basically, changes to the rules of botanical nomenclature are
decided through a democratic process.
Kingdom/Regnum -tae
Division/Phylum -phyta -mycota
Subdivision/Subphylum -phytina -mycotina
Class -opsida -phyceae -mycetes
Subclass -idae -phycidae -mycetidae
Superorder -anae
Order -ales
Suborder -ineae
Superfamily -acea
Family -aceae
Subfamily -oideae
Tribe -eae
Subtribe -inae
Below the level of Subtribe, taxon names do not have a specific ending. The
most important ones are (mandatory ones in bold):
Supergenus
Genus
Subgenus
Section
Species
Subspecies
Variety
Subvariety
Forma
34 Subforma
The names of non-mandatory taxa are composed of a single word that fol-
lows the nearest mandatory name above it. The name of a species is com-
posed of the genus name plus the species indication, hence a second word,
called the epithet. The name of an infraspecific rank is also composed of a
single word.The species epithet and all infraspecific names are always starting
with a lowercase letter, those of ranks above species start with a capital. Some
examples:
The application of names of taxa at ranks above the family may be deter-
mined by the name of an included genus (e.g. Order Asparagales is derived
from the generic name Asparagus), or may be descriptive names (such as
Division Spermatophyta). The application of names of taxa at the rank of
family or below however is determined by means of nomenclatural types. A
nomenclatural type is the element to which a name is permanently attached,
whether that name is the accepted name or a synonym of another name
does not matter.
- effectively published
In botany, the name of the species epithet cannot be the same as the genus
name. In zoology this is allowed (Bufo bufo for the Common toad, or Giraffa
giraffa for the Southern giraffe). Such a name is called a tautonym and is inva-
lid under the botanical Code.
3.5 Types
Having the correct type specimen linked to a name is essential to plant (and
animal) nomenclature. A number of rules have been put in place to deal with
situations where the type may be uncertain.
Before 1958, one was not obliged to indicate the type specimen for a new
name. As a consequence publications preceding this date regularly have pro-
tologues that do not mention a type but rather cite several collections con-
sulted by the author for the new taxon.These are then all regarded as ‘original
material’ and are called syntypes. As a name may only have a single type, one
must choose a type from amongst this original material (the collections cited
and all their duplicates). Such a chosen type is then called the lectotype.
Duplicates from the lectotype are then called isolectotypes. When someone
publishes a lectotypification, adding the phrase “designated here” is obligatory.
Once a type collection is assigned to the name, the remaining syntype mate-
rial automatically become paratypes.
In a situation where all original material, including any relevant illustrations, has
been lost (after proof of an exhaustive search), one is allowed to select a new
type that is then called the neotype. Duplicates of the neotype then become
isoneotypes. When creating a neotype, one often tries to select material that
was collected at or close to the original type locality, but this is not obligatory.
In general, one aims to select a neotype where nomenclatural stability is guar-
anteed, hence which does not lead to necessary name changes.
Lectotype designation:
Anthephora elegans Schreb. var. africana Pilg. (Pilger 1901: 119). – Type:
D.R. Congo, Stanley-Pool, June 1899, Schlechter 12508 (lectotype: B
[B 10 0168252], designated here; isolectotypes: B [B 10 0168251], BR
[BR0000013591571], K [K000281098], P).
Explanation: Antephora elegans var. africana was published citing
four specimens, Buchholz 1875, Dinklage 464, Dewèvre 120 and
Schlechter 12508 which are to be regarded as syntypes and
comprise the original material. Since the author worked at Berlin
(B), the lectotype should preferably be located there. All except the
Schlechter specimen are not present at B and were presumably lost
during the 1943 fire. At B, there are two sheets of Schlechter 12508,
one of which has no spikelets left, the other with a few spikelets in an
envelope glued onto the sheet. The latter is selected as the lectotype,
with duplicates located at Meise Botanic Garden, Belgium (BR), Royal
Botanic Gardens, Kew (K) and Muséum national d’Histoire naturelle,
Paris (P). The barcodes are added to the specimens for which these
were available.
The person publishing a new taxon name is the author of that name, and in
formal or official documentation is placed after the taxon name concerned.
The author name is often abbreviated, for which a standard abbreviation was
published by Brummitt & Powell (1992) and for which an
online database is now maintained by IPNI (at http://www.
ipni.org).
The fact that an author intends to publish a new taxon name is often indi-
cated by adding the abbreviation spec. nov. or genus nov. or subsp. nov., etc.,
behind the name.
When an author moves a species from one genus to another, the epithet
is transferred to the new genus while the original author is placed between
brackets after it, followed by the transferring author e.g. Cenchrus purpureus
(Schumach.) Morrone. This species was originally named Pennisetum purpure-
um by Schumacher (1827) and transferred to the genus Cenchrus by Morrone
(2010). Note that the gender of the epithet has changed in accordance with
Latin grammar. The name Cenchrus purpureus (Schumach.) Morrone is called
a new combination (often abbreviated as comb. nov.) since it combines the
original (protologue) epithet with the name of another genus. The name that
provided the epithet for the new combination is called the basionym; in this
case Pennisetum purpureum Schumach.
When studying a certain group of taxa, an author may consider two or more
names to represent the same taxonomic unit. Following the type concept,
this basically means that this author is of the opinion that the type specimens
of both names belong to the same taxon. For example, Clayton & Renvoize
(1982) considered the following species names, in alphabetical order, to rep-
resent a single, variable species of grass:
This implies that all five names are synonyms, but the nomenclatural rules
stipulate that only a single name can be the accepted name; so, which one do
we choose? Here we should apply the priority rule (Principle III of the Code),
which tells us that the oldest synonym has priority over the others. In this
case, the correct and accepted name for this species is Pennisetum macrourum
Trin., since it was published in 1826.
The priority rule applies to all taxonomic levels. For example, in 2010, Mor-
rone published an article in which he merged the genus Pennisetum Rich.
(Richard in Persoon 1805: 72) with Cenchrus L. (Linnaeus 1753: 1049). The
priority rule shows the latter genus has priority over the first, and so the
genus in its new circumscription should be called Cenchrus.
When we follow the view of Morrone (2010), the accepted name for the
species Pennisetum macrourum Trin. becomes Cenchrus macrourus (Trin.) Mor-
rone. Note, that when another author does not agree with this hypothesis
and advocates to maintain the genus Pennisetum, there are two accepted
40 names for the same species, depending on the scientific point of view.
Further, it is important to know that the priority rule only applies to names
of the same taxonomic rank! In the previous example, if the name Pennisetum
polystachion (L.) Schult. var. africana Thunb. (Thunberg 1794: 101) would have
been a synonym of the five other names in Pennisetum mentioned above,
then it would have been the oldest name available. However, as it is a name at
the rank of variety, it has no priority over names at species level. When author
Xxx would want to upgrade that variety to the species level (as Pennisetum
africanum (Thunb.) Xxx) the publication date of that name would be the date
this new combination was published. It logically follows that when the taxon
Ixora aneimenodesma K.Schum. subsp. kizuensis De Block has no synonyms
and an author Xxx wants to raise it to species level, they have two options:
1) to publish the name Ixora kizuensis (De Block) Xxx, or 2) to publish a new
species name (e.g. Ixora congoensis Xxx), with the subspecies name as a syn-
onym. Option 2 is not considered ‘polite’ since it removes the original author
from the name. However, the name Ixora kizuensis may already exist for a dif-
ferent species. In that case, the necessary new combination is ‘occupied’ and
one has to opt for a new name, such as Ixora deblockiae Xxx to honour the
original author. The necessity to create such a new name for a taxon already
described before is often indicated by the addition nom. nov.
There are two exceptions to the priority rule. The first is that there are eight
family names and one subfamily name where one is allowed to choose be-
tween two alternatives (ICN Art. 18.5, 19.8). Such names are called nomina
alternativa (or nom. alt.). Below is a list of these allowed alternative family
and subfamily names. In a single publication it is advised to use the names of
only one of the columns.
Apiaceae Umbelliferae
Arecaceae Palmae
Asteraceae Compositae
Brassicaceae Cruciferae
Clusiaceae Guttiferae
Fabaceae Leguminosae
incl. subfam. Faboideae incl. subfam. Papilionoideae
Lamiaceae Labiatae
Poaceae Gramineae
Secondly, the strict application of the rules laid down in the ICN may lead to
‘unwanted’ changes and major instability of the nomenclature within a par-
ticular taxonomic group. In that case, one can make a proposal to conserve
or reject a specific name. In case of ambiguity related to the correct type
specimen, a similar proposal to conserve a specific type can me formulated. 41
Such proposals need to be published in the journal Taxon and are then vot-
ed on at the next International Botanical Congress. Conserved or rejected
names or types are generally followed by the indication ‘nom. cons.’, ‘nom.
rej.’ or ‘type cons.’.
3.8 Hybrids
In the Code, names of hybrid taxa are dealt with in a separate chapter. They
can be recognized by the use of the multiplication sign × or by the addition of
the prefix “notho-” to the term denoting the rank of the taxon. A nothospe-
cies name, composed of a genus name (or a nothogenus name, see below)
and an epithet, indicates a hybrid between two individuals of different species.
A nothogenus name, a single word, is used when a hybrid has been formed
between individuals of species belonging to different genera. It is often com-
posed of parts of the names of the two genera involved.
For example, the hybrid between Oenothera biennis L. and Oenothera villosa
Thunb. can be either indicated by the hybrid formula Oenothera biennis L. ×
Oenothera villosa Thunb., or by the nothospecies Oenothera ×drawertii Renner
ex Rostański.
Names of cultivated plants are not regulated by the ICN, but rather by the
International Code of Nomenclature for Cultivated Plants (ICNCP).
Cultivated forms can be indicated by only three categories, the Cultivar, the
Group or Cultivar group, and the grex. The latter is used solely in orchid
cultivation and indicates the combined hybrid offspring of any cross between
the same two entities (taxa or cultivars). A cultivar, abbreviated as cv., is a very
specific form derived from any type of selection process and may even have
been taken directly from the wild. It is a non-Latin name added after the name
of the taxon from which it was derived e.g. Solanum tuberosum L. cv. Gogu
valley, also written as Solanum tuberosum ‘Gogu valley’. When it is unclear to
which species a cultivar belongs, the cultivar name can follow directly after
the genus e.g. Rosa cv. Penelope. A new cultivar name can be formally regis-
tered by an International Cultivar Registration Authority that needs to be ap-
proved by the ISHS Commission for Nomenclature and Cultivar Registration.
Each Authority is assigned a specific taxonomic group. After the Authority
has formally approved the registration of a new cultivar name, the person
who provided the information ‘owns’ the rights to this name. They can then
42 market both the name and the plants, quite similar to a patent. A Cultivar
group, abbreviated as cv. gr., comprises a number of cultivars having a distinct
characteristic. One could, for example, create a Cultivar group for all yellow
roses. Here it becomes clear that names of cultivated plants are not part of a
natural classification, since they need not indicate or reflect common ancestry.
In literature on cultivated plants, one may regularly come across ‘variety’ or
‘form’. Note that these terms should actually not have been used for cultivat-
ed plants, as they erroneously refer to the ICN which does not deal with cul-
tivated plants. When possible, such uses of ‘variety’ or ‘form’ should be treated
as informal descriptions of the variation observed without the intention to
create a new taxon name under the ICN.
• http://www.iapt-taxon.org/nomen/main.php?
The art
of identification
44
In the preceding chapters, we mentioned that the taxonomic science aims
to organize the immense diversity of living organisms on earth into discrete
units. As such, it provides the essential tools for scientific communication in
the form of names and a classification. To be able to perform biological re-
search, protect nature, use plants for medicinal purposes, etc., it is crucial to
have access to the wealth of information accumulated over several centuries.
Online resources are growing exponentially. However, before one can tap
into the information available on certain species or genera, for example, one
has to know the name of the particular taxon concerned. As specialists who
can identify living organisms by heart are rare, especially in the tropics where
diversity is high, taxonomists have developed tools to reliably identify their
material.
In order to use an identification key, one generally has to have at least a basic
understanding of plant or fungal morphology and terminology. Having a good
glossary may be useful. Various good and extensive botanical glossaries exist
(see the text box at the end of this chapter).
In the example above, each question is called a couplet of the key, having
two leads. It is obvious that the two options need to be mutually exclusive,
not showing overlap. After having correctly answered a certain number of
questions, the user will end up with the name of the plant (or fungus, or 45
animal) at hand.
Generally, the user is given the choice between two options. In such a case,
the key is dichotomous. Some keys allow the choice between three or even
more options (in the example above, one could add ‘Flowers blue’, ‘Flowers
red’, for example to arrive at four leads. These are called polytomous keys.
In general, this structure is deemed less practical and more prone to identifi-
cation errors. One can easily avoid such choices by combining several states
into one lead, such as:
Note that question #3 provides the name of the plant, and that questions
6 and 10 show which previous question pointed to it. The latter is simply
assisting the users in keeping track of where they came from, and is generally
added only when one has arrived after having made a comparatively large
‘leap’ in the key.
Basically, there are two forms of dichotomous keys. The form shown above
where both leads directly follow each other is called a bracketed key. The
second form is called an indented key and separates the two leads in space.
Here is an example of an indented key (adjusted from a key to the species of
Solanum in Africa, Vorontsova & Knapp 2016):
When the first lead of couplet 1 corresponds with the plant to be identified,
one should carry on to the next question formulated immediately below it
(number 2). However, when the second lead of couplet 1 is correct, one con-
tinues with the question below, which is number 4. As can be seen, there are
no numbers to the right-hand side of the key pointing to the next question.
The advantage of an indented key is that the user may more easily derive the
way the species are grouped from the structure of the key. A disadvantage
is that one has to search for the second lead of the question/couplet con-
cerned. This can be quite far down in larger groups, and in longer keys there
will be a lot of unused space on the left side of the page, resulting in a key
that needs more printed pages.
Also, note that geographical information may be used in the key. Despite not
being a morphological character, it may be deemed helpful. Similarly, ecologi-
cal or phenological information (flowering/fruiting period) may be added. It is
however advised that these types of characters be used only as supplemen-
tary data to morphological features.
Start by choosing which type of key to construct (see above). Then, think of
several clear subgroups that can be recognized within the group concerned.
Next, select those groups that can be defined by character states that are
well distinct and can be easily observed with the naked eye or when using a
10× hand lens. If the first question of a key is about pollen grains, for example,
a fair number of users will be immediately stuck and unable to continue. It is
also important that any character mentioned in one lead of a couplet, should
also be present in the other lead(s)! A couplet like the one below is not to
be recommended:
A user having a plant that carries white flowers but also has leaves longer
than 10 cm will become uncertain as to what to choose. This brings us to
another practical issue. When making a key, one should always try to imagine
what a user may have on hand! This is often a single plant, so the key needs
to provide exact information. If a couplet states “Flowers big” against “Flow-
ers small”, this is a relative concept and the user is unable to judge whether
flowers of 1 cm in diameter may be considered ‘big’ or ‘small’.
47
A single taxon may key out more than once. That may occur when a specific
taxon is variable for a certain character. For example, a species may have
white as well as occasionally yellow flowers, while most species are constant
in that respect.
Figure 18. Example of Creating a key for a larger group of species
a species/character data (or other taxa) is often greatly facilitated by
matrix (with fictional taxa the preparation of a taxon/character data ma-
Aus a, Bus x, etc.). trix (see also figure 18). It often helps to get a
better overview of the distribution of charac-
ters and their correlation. (See also the next
paragraph.)
2. In a fully dichotomous key, when all taxa key out only once, the number
of couplets will always be equal to the number of taxa minus 1. Hence,
one cannot influence the number of couplets. However, one can influ-
ence the number of questions to be answered before a species is keyed
out. The best strategy is to strive for questions/couplets that divide the
remaining group of taxa into more or less equal parts.
3. If the species in a group generally do not have flowers and fruits at the
same time, it may be wise to present two different keys, one for flowering
material and one for fruiting material.
The keys discussed above, even when prepared with the utmost care, have
a serious flaw. The user cannot choose the sequence in which characters are
observed. It could be that having red fruits would already greatly reduce the
number of potential species, but that fruit colour is only asked in question
number 5. In other words, there should be easier ways to identify a plant!
A different kind of multi-entry key is the so-called diagnostic key (also called
a synoptic key). This comprises a list of diagnostic or spot characters (typical,
remarkable) occurring within a certain group of taxa. Each character is then
49
followed by a list of all taxa within the group that possess that character. Be-
low is an example of part of a diagnostic key to taxa of Rubiaceae in Central
Africa. Note that the taxa indicated are mostly genera, but also tribes and
even species.
- Leaf
- Flower
unisexual: Anthospermum
heterostylous: Colletoecema, Craterispermum, Gaertnera, Knoxieae,
Lasianthus, Morinda, Mussaendeae, Pauridiantha, Psychotrieae,
Sabicea, Schizocolea, Spermacoceae, Tricalysia
4-merous: Anthospermum, Corynanthe, Eumachia, Galium, Heinsia, Ixora,
Keetia, Knoxia, Lasianthus, Nauclea, Otiophora, Paraknoxia, Pavetta,
Polysphaeria, Pouchetia, Psychotria, Rutidea, Spermacoceae, Tricalysia
pleiomerous (with more elements than usual): Coffeeae, Gardenia,
Rothmannia octomera, Schumanniophyton
calyx tube long (> 1 cm): Adenorandia, Gardenia, Rothmannia,
Schumanniophyton hirsutum
calyx tube with a lateral slit: Calycosiphonia, Gardenia, Polysphaeria,
Rothmannia, Sericanthe, Tricalysia
calyx asymmetrical, with highly unequal lobes, or only 1 lateral lobe:
Knoxieae
The DNA barcode database needs to be based on a sound and stable tax-
onomic framework of genera and species. Even for a comparatively well-
known group like plants, this framework still has many weak spots. In their
turn, the results of DNA barcoding efforts may well assist in making better
taxonomic decisions, thereby strengthening the framework.
While identifying living plants in the field can be difficult, trying to correctly
identify dried and flattened herbarium specimens is often challenging. Not all
characters needed may be directly visible, even with the help of a good 10×
hand lens or a stereo microscope (see also paragraph 5.B). The simple ques-
tion whether the flowers are white or yellow may remain unanswered when
the collector did not record this information in the field. Similarly, one may 51
wonder whether this twig with few leaves and nice flowers or fruits comes
from a big tree, a liana, a shrub or even a perennial herb? The 3-dimension-
al shape (notably of flowers and fruits) could be important, but impossible
to reconstruct, as is information about underground tubers, rhizomes, smell,
taste, etc.These kinds of information should be noted down in the field by the
collector so that the information can be transferred to the label accompany-
ing the specimen. Various publications (Fish 1999, Victor et al. 2004, Bridson
& Forman 2010) offer good advice on how to collect plants and correctly
prepare valuable dried plant specimens.
When a herbarium specimen has been identified, whether this is at the level
of family, genus, species or other, the taxon name is written on a small piece
of paper, called an identification or determination slip. This also includes the
name of the researcher (and affiliation if possible) and the date. It is glued
onto the herbarium sheet (when not gummed, use special glue provided by
the herbarium curator), preferably somewhere in the lower right corner and
always above any previous identifications slips. Make sure only a small part of
the slip is glued, so that the remainder can be flipped back to examine any
material or written text it may cover. Some herbaria will only allow the use of
needles to attach labels and slips to the sheet.
Glossaries
• Beentje H. (2015) The Kew plant glossary. 2nd edition. Richmond, Royal
Botanic Gardens, Kew. EAN: 9781842466049
• Josserand M. (1983). La description des champignons supérieurs. 2e éd.
Paris, Lechevalier.
• Jouy A., Foucault B. de (2016) Dictionnaire illustré de botanique. Mèze,
Biotope.
• Missouri Botanical Garden Glossary:
http://www.mobot.org/mobot/glossary
• Wikipedia: https://en.wikipedia.org/wiki/Glossary_of_botanical_terms
[in French: https://fr.wikipedia.org/wiki/Glossaire_de_botanique]
DNA barcoding
53
54
Here, we focus on a taxonomic revision largely based on herbarium speci-
mens. When studying the taxonomy of a plant or fungal group for which field
observations are not easily obtained, e.g. tropical species, or simply those
from remote places, herbarium specimens are often the only source of infor-
mation available. Adding field observations to a revision based on herbarium
specimens is a big advantage, but it is not a necessity.
Let’s assume that it has been decided to prepare a taxonomic revision for
a specific group. This is generally because specialists have indicated that the
taxonomic framework for a particular group is considered ‘weak’. This could
be the result of the available identification keys being of poor quality, of un-
clear boundaries between taxa/species, or of doubts about the correctness
of the names applied. It is not uncommon that part of the “weakness” may
be due to the presence of undescribed species. Each year, well over 2000
new species of vascular plants are described, as well as some 75 to 100 new
genera. This trend has not decreased over the past 15 years (numbers have
even risen slightly over the past four years), indicating there is still plenty to
be discovered!
Referring to the depth, thoroughness and practicality of the revision, one can
identify four categories:
All scientific studies start with gathering information and data. For a taxonom-
ic revision, it is necessary to gather all the protologues (original publications)
for the names concerned.This is a crucial step, especially in phase E when type
specimens need to be identified, selected and assigned. The service provided
by IPNI (International Plant Name Index; http://www.ipni.org) for vascular
plants and Index Fungorum (http://www.indexfungorum.org) for fungi can
generally provide a list of all names within the relevant genus, although some
further filtering will be needed for regional studies. Note that IPNI did not
collect data on infraspecific taxa until 1971! Some may have been entered
now, but in general these can only be found through extensive searching
(internet and libraries).
From the protologue information, one can now track down the relevant pub-
lications by:
Scan or download the relevant pages and file these in an easily traceable sys-
tem. Also, be sure to note the full reference (see below)! This will be needed
when publishing the study.
56
Next, gather all relevant books and articles that deal with the systematics
of the chosen genus. It is also useful to study relevant papers dealing with
phylogenetics, biogeography, ecology, etc., as this will give greater insight into
the genus and its relatives, especially the evolutionary importance of some of
the characters. Use available internet search engines as well as portals of spe-
cialized libraries using the name of your taxon as keyword, along with others
such as ‘taxonomy’, ‘revision’, ‘systematics’ etc. Start with the most recent pub-
lications, and study the publications cited in their reference lists. Scroll through
their paragraph on the taxonomic and systematic history of the genus when
available. Also, consult Floras relevant to the study area.
Studying relevant literature should give you a fairly good idea of the position
of the genus within the family, which genera are closely related, and which
characters are deemed distinctive and informative for species delimitation.
Going through identification keys to see which characters have been used to
distinguish the species will probably prove to be extremely worthwhile. Nev-
ertheless, it is always important to develop a personal view on the variation
of the group studied.
Do not forget to record the full reference when taking notes from publi-
cations! It can be very frustrating and time consuming when one cannot
remember where one read something.
Here are examples of references for publication types that you are most likely
to encounter:
1. Article in a journal:
Soreng R.J., Peterson P.M., Davidse G., Zuloaga F.O., Judziewicz E.J., Filgue-
iras T.S., Davis J.I., Morrone O. (2015) A worldwide phylogenetic classifi-
cation of the Poaceae (Gramineae). Journal of Systematics and Evolution
53(2): 117--137. http://dx.doi.org/10.1111/jse.12150
2. Book:
Patil J.V. (2016) Millets and Sorghum: Biology and Genetic Improvement.
Chichester, John Wiley & Sons Ltd. 504 pp.
Finally, set up a documentation system or data base, where you indicate for
each taxonomic name, where the protologue can be found and what its type
specimen is. Other authors may have already indicated which specimen is the
type, however this should always be checked. The protologue will generally
provide the necessary information concerning the type(s), but certainly not 57
in all cases. Searching for type specimens can be a time-consuming activity!
This is particularly true for older literature, as it was not obliged to indi-
cate a type. The majority of major Herbaria have scanned and digitized their
type specimens. These images are available at the JSTOR Global Plants portal:
https://plants.jstor.org, and generally also through the web-portals of the in-
dividual institutes. Note that this facility only shows known type material for
the various Herbaria. A fair number of types still have to be identified as such
by taxonomists. In the course of a revision, it is common to encounter type
specimens that had previously gone unnoticed.
B. Herbarium observations
It should be noted that herbarium specimens are valuable and often irre-
placeable scientific objects that are brittle and easily damaged! They must be
handled with the utmost care. Always ask the appropriate Herbarium Cura-
tor about the specific handling instructions.
When material, like pollen or a piece of leaf for DNA extraction, is not
returned to the sheet (called destructive sampling), the researcher is
supposed to add a label to the specimen stating what was removed,
for which purpose, by whom and when. Again, ask permission from the
58 Curator before you do!
One will need to develop a strategy on how to obtain or consult the majority
of the herbarium specimens available elsewhere. Phase A (taxonomic names
and literature) has provided you with a fairly good idea of the distribution
of the genus, and where most of the species occur. One might consult col-
leagues to find out which Herbaria contain the vast majority of the relevant
material. One can either visit these institutes or have the material to be on
loan to your home institute. Herbaria addresses and contact persons can
be found on Index Herbariorum (http://sweetgum.nybg.org/science/ih). Note
that it may take several months for a loan shipment to arrive. Herbaria will
generally not send more than several hundred specimens as a loan, and they
may have restrictions related to the countries requesting material. An option
will then be to visit the institution where the material is housed.
When starting your study, you will need a fair amount of herbarium spec-
imens, so ask for loans as soon as possible. In some cases (i.e. when one
already has a fair knowledge of the group concerned), loans can be requested
months before the actual start of the revision. Herbaria that house a lot of
your material are best visited physically, but this is often expensive. It will gen-
erally be far more efficient to visit them once you already have a good knowl-
edge of the species/taxa within your group. So, a good work plan is essential.
Some herbaria have their collections available on-line (see textbox at the end
of this chapter). These services often provide some label data and high-res-
olution scans of available material. These on-line services are extremely
helpful, but experience
shows that a fair num-
ber of sheets need to be
loaned to perform more
detailed observations on
the actual specimens.
Figure 20.
Example of a simple
laboratory device to boil
water in a small cup.
59
C. Data basing
- Barcode
- Main collector (preferably surname and initials in separate fields)
- Additional collector(s)
- Prefix (some collectors add a code or number before the collection
number which refers to a research mission, the collecting year or desig-
nates a project by its acronym)
- Collection number
- Suffix (any code given after the collection number, see prefix)
- Collecting date
- Country
- Locality
- Latitude
- Longitude
- Habitat
- Altitude
- Uses
- Vernacular names
- Family
- Genus
- Species
- Author(s)
- Infraspecific level (subspecies, variety, forma)
- Infraspecific name
- Infraspecific author(s)
- Identified by
- Identification date
- Herbarium code
- Type of
- Notes
Often, Herbaria that have their specimens digitized will be willing to send
their data in a digital format, which can be uploaded into your own data base
after some necessary adjustments.
A species distribution area can vary from small to large, this is generally re-
lated to the species having a narrow or broad ecological tolerance. A species
restricted to a defined region is said to be endemic to that region. A species
can be endemic to a mountain, national park, province, country, continent, etc.
(and all species are endemic to the planet Earth!). So, simply stating that a
species is endemic without reference to the region is meaningless.
D.1 Mapping
Note that there are various coordinate systems! These are related to different
‘projections’, or ways in which the globe has been transposed onto a map.
There are also different ways of recording coordinates. The most commonly
used coordinate format used by taxonomists is Degrees, Minutes and Seconds,
but in some regions the UTM (Universal Transverse Mercator) coordinate
system is preferred. Also note that some may use a normal DMS (Degrees,
Minutes, Seconds) format, e.g. 15°12’55”N 30°21’32”E, while others prefer
DD (Decimal Degrees), e.g. 1.247°N 25.873°E, or DM (Decimal Minutes), e.g.
11°34.75’N 25°21.30’E. A number of online tools (for example http://www.
synnatschke.de/geo-tools/coordinate-converter.php) allow to easily convert
coordinates from the different systems.
Google Earth is also a useful tool to find the collecting locality, but older col- 61
lections may have place names no longer in use. For some regions, a published
index of plant collecting localities is available, and some websites provide
historical maps (see textbox at the end of this chapter). The libraries of many
natural history institutes will often have a good collection of historical maps,
collecting registers (collecting notebooks of the collector) or Gazetteers (a
book, usually per country, with all place names, including rivers, mountains,
etc., with their lat./long. information). On-line gazetteers, like GeoNames (see
textbox at the end of this chapter), are also a good way to search for loca-
tions as they often have historic names of places as well as allowing “fuzzy”
search options. Furthermore, a collector will often have collected several
specimens at the same locality and so the specific locality may already have
been georeferenced by someone else. Check web-portals providing such
data on-line, notably from an institute where you know duplicates from a par-
ticular collector were deposited. Some specimen data bases offer the option
to create an itinerary for a specific number range of a collector which may
assist one in finding the right information.
Plotting your specimen data on a map can be done using Google Earth, but
when you want to prepare a high-quality distribution map for publication, you
should look for other software (e.g. DivaGis, QGIS or ArcView).
It is often stated that this way of taxon delimitation applies the morphological
species concept (see Chapter 2). However, when we think about what mod-
ern taxonomists actually do, we may conclude that they are in fact trying to
interpret morphological, geographical and ecological data, in terms of a bio-
62 logical species concept based on non-interbreeding populations. When a ‘pile’
of specimens has several distinct morphological characters, one may assume
they have arisen from a distinct genetic basis. This can only remain distinct if
there is no interbreeding. The same applies to geographical and ecological
information. A taxonomist will generally interpret such data within the light of
potential interbreeding. In conclusion, one might state that herbarium-based
plant and fungal taxonomy (but also natural history specimen-based animal
taxonomy) attempts to apply a biological species concept through interpreta-
tion of morphological and other patterns observed. Incorporating the results
of molecular studies to determine whether the various species are reproduc-
tively isolated is encouraged when available, but that is beyond the scope of
this chapter.
After one has finalized the taxonomic framework, the next step is to establish
the correct scientific name for each pile. This is where nomenclatorial deci-
sions have to be made.
Locate all type specimens of all names, species as well as infraspecific taxa,
and verify in which pile they are (even those that may not be physically
present!). The type specimens in each pile represent the potential names to
be considered for that particular taxon. Following the rules of the ICN, as
described in Chapter 3, should then lead to identifying the accepted name
and its synonyms. Any pile that has no type specimen is a new taxon that will
need to be formally described.
When creating a new taxon, one must meet all requirements of the ICN in
order to validly publish the name (see Chapter 3).
One should avoid vague or relative terminology like “rather densely hairy”
or “fairly long”. For 2-dimensional shapes, it is advisable to use the stand-
ardized set of terms provided by the Systematics Association Committee
for Descriptive Biological Terminology (1962). Descriptions follow a logical
format, with the various elements being dealt with in a specific sequence:
plant – root/stem – leaves – inflorescence – flower – fruit – seed. Within
these elements, the sequence is from bottom to the top and from outside to
inside. For any organ, a good sequence for descriptive characters is: number
of elements – position – overall shape – overall dimensions – base/top/mar-
gins – texture – colour and lustre – surface (smooth, rough) – indumentum
and/or appendages. For more detail, see the text box below.
For fungi, a description guideline, similar to the one presented in the text
box, has been recently published in French (Eyi et al. 2011), and can be freely
downloaded from http://www.abctaxa.be/volumes/volume-10-les-champi-
gnons-comestibles-de-l-afrique-centrale
Then, notes may be added providing the arguments for the taxonomic deci-
sions and/or the choices made related to the typification of names, etc.
64
It is advisable to add botanical illustrations to the revision. These are inval-
uable in the identification process and assist the user in understanding the
diagnostic elements of the taxa. One can make these illustrations oneself,
but it is usually better to ask a skilled botanical illustrator. Becoming a skilled
Guide to a well-structured plant description
Finally, one will need to prepare an identification key to the taxa studied.
The key should be practical and assist the user in the identification process.
Preferably use characters that are easily observed. For other suggestions, see
Chapter 4.
When the taxonomic part of the study (Genus treatment, Key to the spe-
cies, Species treatments, maps, illustrations, etc.) is finished, one will need to
prepare the manuscript for publication. When the revision is meant to be
incorporated in a Flora, the taxonomic part is often all that is needed. When,
however, one wants to publish the results as an article in a scientific journal,
various other paragraphs will need to be added. While some are general, e.g.
Introduction, Materials and Methods, others are typical for a taxonomic revi-
sion, e.g. the History of the Genus, providing a historical overview of previous
studies and their contributions to the taxonomic framework of the genus.
66
Plant collecting
Index Herbariorum
• http://sweetgum.nybg.org/science/ih
68
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