Arctic
Biodiversity
Assessment
Status and trends in Arctic biodiversity
ARCTIC COUNCIL
i
Chief scientist and executive editor:
Hans Meltofte
Assistant editors on species chapters:
Alf B. Josefson and David Payer
Lead authors:
Tom Barry
Dominique Berteaux
Helga Bültmann
Jørgen S. Christiansen
Joseph A. Cook
Anders Dahlberg
Fred J.A. Daniëls
Dorothee Ehrich
Jon Fjeldså
Finnur Friðriksson
Barbara Ganter
Anthony J. Gaston
Lynn J. Gillespie
Lenore Grenoble
Eric P. Hoberg
Ian D. Hodkinson
Henry P. Huntington
Rolf A. Ims
Alf B. Josefson
Susan J. Kutz
Sergius L. Kuzmin
Kristin L. Laidre
Dennis R. Lassuy
Patrick N. Lewis
Connie Lovejoy
Hans Meltofte
Christine Michel
Vadim Mokievsky
Tero Mustonen
David C. Payer
Michel Poulin
Donald G. Reid
James D. Reist
David F. Tessler
Frederick J. Wrona
Arctic Biodiversity Assessment
Status and trends in Arctic biodiversity
ARCTIC COUNCIL
ARCTIC COUNCIL
ii
Arctic Biodiversity Assessment
Status and trends in Arctic biodiversity
Conservation of Arctic Flora and Fauna (CAFF),
Arctic Council, 2013
Chief scientist and executive editor:
Hans Meltofte, Aarhus University
Suggested referencing:
CAFF 2013. Arctic Biodiversity Assessment.
Status and trends in Arctic biodiversity.
Conservation of Arctic Flora and Fauna, Akureyri.
or
Meltofte, H. (ed.) 2013. Arctic Biodiversity Assessment.
Status and trends in Arctic biodiversity.
Conservation of Arctic Flora and Fauna, Akureyri.
Linguistic editor:
Henry P. Huntington, Huntington Consulting
Graphics and layout:
Juana Jacobsen, Aarhus University
The report and associated materials can be downloaded for free at www.arcticbiodiversity.is
Disclaimer:
The views expressed in this peer-reviewed report are the responsibility of the authors of the
report and do not necessarily relect the views of the Arctic Council, its members or its observers,
contributing institutions or funding institutions.
Funding and Support:
The Arctic Biodiversity Assessment has received inancial support from the following sources:
Canada, Denmark/Greenland, the Danish Environmental Protection Agency as part of the Danish
environmental support programme Dancea – the Danish Cooperation for Environment in the Arctic,
Finland, Norway, Sweden, United States of America, and the Nordic Council of Ministers.
Educational use:
This report (in part or in its entirety) and other Arctic Biodiversity Assessment products available from
www.arcticbiodiversity.is can be used freely as teaching materials and for other educational purposes.
The only condition of such use is acknowledgement of CAFF/Arctic Biodiversity Assessment as
the source of the material according to the recommended citation. In case of questions regarding
educational use, please contact the CAFF Secretariat: caf@caf.is
Note:
This report may contain images for which permission for use will need to be obtained from original
copyright holders.
Print: Narayana Press, Denmark
Impression: 1,300
ISBN: 978-9935-431-22-6
Cover photo: Muskoxen are hardy animals that had a circumpolar distribution in the Pleistocene, but
Holocene climate changes along with heavy hunting may have contributed to its disappearance in
the Palearctic and from Alaska and Yukon. In modern times, humans have reintroduced muskoxen to
Alaska and the Taymyr Peninsula together with a number of places where the species did not occur in
the Holocene. Photo: Lars Holst Hansen.
iii
Contents
Preface by CAFF and Steering Committee Chairmen
................................................................. v
Foreword by the Chief Scientist . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Indigenous peoples and biodiversity in the Arctic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
1. Synthesis: Implications for Conservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
2. Species Diversity in the Arctic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
3. Mammals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
4. Birds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142
5. Amphibians and Reptiles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182
6. Fishes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 192
7. Terrestrial and Freshwater Invertebrates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 246
8. Marine Invertebrates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 276
9. Plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 310
10. Fungi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 354
11. Microorganisms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 372
12. Terrestrial Ecosystems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 384
13. Freshwater Ecosystems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 442
14. Marine Ecosystems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 486
15. Parasites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 528
16. Invasive Species: Human-Induced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 558
17. Genetics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 566
18. Provisioning and Cultural Services . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 592
19. Disturbance, Feedbacks and Conservations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 628
20. Linguistic Diversity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 652
Lead author biographies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 664
Author ailiations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 670
iv
v
Preface by CAFF and Steering Committee Chairmen
The eyes of the world are turning northwards. In recent years, interest in the Arctic
has increased dramatically within and outside of Arctic countries. This is relected
in the amount of attention given to Arctic biodiversity. While the landscapes and
wildlife have been the subject of explorers, scientists, artists and photographers as
well as the home of a variety of peoples for a long time, until recently Arctic biodiversity did not feature very prominently in national or international policy work.
This, however, is changing, as the unique values of Arctic nature are increasingly
discussed at high levels. At the same time, more and more attention has been paid
to the interface between science and policy to ensure that policy is built on the best
science available.
We are therefore very happy and proud to present the Arctic Biodiversity Assessment (ABA), which has been seven years in the making. It is the result of the
contributions from 253 scientists together with holders of traditional knowledge.
The chapters in the main document, which you are holding now, have been peerreviewed by over 100 scientists from all over the Arctic and the rest of the world. We
are very grateful for the eforts they have made to ensure the quality of this assessment. We would especially like to thank Chief Scientist Hans Meltofte and the lead
authors of the chapters.
In order to communicate the indings presented in this scientiic work and to inform
policy makers, the board of the Arctic Council’s working group on the Conservation
of Arctic Flora and Fauna (CAFF) has prepared a summary of the key indings and
developed policy recommendations. The key indings and recommendations have
been provided in a separate document, which we trust will be useful for all those
who make decisions that may afect Arctic biodiversity.
The Arctic is home to a vast array of biodiversity, including many globally signiicant
populations. Included among these are 30% of the world’s shorebird species, two
thirds of the global numbers of geese, several million reindeer and caribou, and
many unique mammals, such as the polar bear. During the short summer breeding
season, almost 200 species of birds arrive from almost all parts of the world, connecting the Arctic with the rest of the globe. We therefore hope that the ABA will be
consulted frequently within as well as outside of the Arctic.
Biodiversity is life. It is the very foundation of our existence on Earth. In the Arctic,
links between biodiversity and traditional ways of life are often seen more clearly
than in many other parts of the world. These are examples of ecosystem services,
the beneits that we receive from nature. Many ecosystems and ecosystem functions
in the Arctic remain largely unstudied and involve little-known organisms, especially
microbes. The ABA presents current knowledge also on these processes and organisms and thus provides a base for further work.
But biodiversity is more than a means for humankind to survive. The unique nature
of the Arctic is not just an asset for us to use. It is also a source of wonder, enjoyment
and inspiration to people living in the Arctic and across the globe. It has intrinsic
values that cannot be measured. We sincerely hope that the ABA will not only create
the baseline reference for scientiic understanding about Arctic biodiversity, but that
it also may inspire people to take efective actions on the conservation of Arctic lora
and fauna. We hope it gives people reasons to love Arctic nature as much as we do.
Yakutsk, 17 February 2013
Evgeny Syroechkovskiy, Chair of CAFF
Mark Marissink, Chair of the ABA Steering Committee
vi
The king eider is one of the fascinating species endemic to the Arctic.
Photo: Patrick J. Endres.
vii
Foreword by the Chief Scientist
Until recently, most Arctic biodiversity was relatively unafected by negative impacts
from human activities. Only over-exploitation of certain animal populations posed
serious threats, such as the extermination of Steller’s sea cow, the great auk, the
Eskimo curlew and a number of whale populations in recent centuries, in addition
to the contribution that humans may have made to the extermination of terrestrial
mega-fauna in prehistoric times.
Human impacts, however, have increased in modern times with increasing human populations in much of the Arctic, modern means of rapid transport, modern
hunting and ishing technology, increasing exploration and exploitation of mineral
resources, impacts from contaminants and, most importantly, with climate change,
which is more pronounced in the Arctic than elsewhere on the globe.
There is no inherited capacity in human nature to safeguard the Earth’s biological assets – moral and intellectual strength are needed to achieve conservation and wise
use of living resources through cultural and personal ethics and practices. Sustainability is a prerequisite for such balance, but it does not come without restraint and
concerted eforts by all stakeholders, supported by mutual social pressure, legislation and law enforcement.
The Arctic is changing rapidly with shorter winters, rapidly melting sea ice, retreating glaciers and expanding sub-Arctic vegetation from the south. If greenhouse gas
emissions are not reduced, Arctic biodiversity will be forever changed, and much
may disappear completely.
On 18 May 2011, 50 prominent thinkers, among them 15 Nobel Prize winners, issued
The Stockholm Memorandum, which among other things states that:
Science indicates that we are transgressing planetary boundaries that have kept civilization safe for the past 10,000 years. Evidence is growing that human pressures are starting to overwhelm the Earth’s bufering capacity. Humans are now the most signiicant
driver of global change, propelling the planet into a new geological epoch, the Anthropocene. We can no longer exclude the possibility that our collective actions will trigger
tipping points, risking abrupt and irreversible consequences for human communities
and ecological systems. We cannot continue on our current path. The time for procrastination is over. We cannot aford the luxury of denial. We must respond rationally,
equipped with scientiic evidence.
Among the many current and projected stressors on Arctic biodiversity addressed
in this report is that of invasive species. However, if we want to do something about
the many problems facing nature and biodiversity in the Arctic, we need to focus on
the impacts of the most globally ‘invasive species’ of all: Homo sapiens.
Hans Meltofte
Copenhagen, 8 February 2013
8
As opposite to the huge and almost totally ice covered Antarctica, the Arctic is an ocean of
pack ice surrounded by a relatively limited fringe of tundra on the adjacent islands and continents. Bowhead whales surfacing amongst melting ice with black guillemots resting on ice.
Foxe Basin, Nunavut, Canada, July. Photo: Eric Baccega.
Arctic Biodiversity Assessment
310
Arctic Biodiversity Assessment
Purple saxifrage Saxifraga oppositifolia is a very common plant in poorly vegetated areas all over the high Arctic. It even
grows on Kafeklubben Island in N Greenland, at 83°40’ N, the most northerly plant locality in the world. It is one of the irst
plants to lower in spring and serves as the territorial lower of Nunavut in Canada. Zackenberg 2003. Photo: Erik Thomsen.
311
Chapter 9
Plants
Lead Authors
Fred J.A. Daniëls, Lynn J. Gillespie and Michel Poulin
Contributing Authors
Olga M. Afonina, Inger Greve Alsos, Mora Aronsson, Helga Bültmann, Stefanie Ickert-Bond, Nadya A. Konstantinova,
Connie Lovejoy, Henry Väre and Kristine Bakke Westergaard
Contents
Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 312
9.1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 313
9.2. Vascular plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 314
9.2.1. Taxonomic categories and species groups . . . . . . . . . . . . . . . . . . . . 314
9.2.2. The Arctic territory and its subdivision . . . . . . . . . . . . . . . . . . . . . . . 315
9.2.3. The lora of the Arctic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 316
9.2.3.1. Taxonomic structure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 317
9.2.3.2. Endemic species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 318
9.2.3.3. Borderline species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 318
9.2.3.4. Non-native species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 318
9.2.3.5. Origin and integrity of the Arctic lora . . . . . . . . . . . . . . . . 319
9.2.3.6. Species richness in Arctic subzones . . . . . . . . . . . . . . . . . . 322
9.2.3.7. Species richness in loristic provinces . . . . . . . . . . . . . . . . 322
9.2.3.8. Hotspots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 326
9.2.4. Traditional use of vascular plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 326
9.2.5. Rare and threatened Arctic endemic species . . . . . . . . . . . . . . . . . 326
9.2.5.1. Rare Arctic endemic species . . . . . . . . . . . . . . . . . . . . . . . . . 326
9.2.5.2. Threatened Arctic species . . . . . . . . . . . . . . . . . . . . . . . . . . . . 327
9.2.6. Trends and monitoring eforts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 328
9.2.7. Conclusions and recommendations . . . . . . . . . . . . . . . . . . . . . . . . . . 329
9.3. Bryophytes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 329
9.3.1. Bryophytes in the Arctic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 330
9.3.2. Arctic bryoloristic studies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 332
9.3.3. Regional surveys of Arctic bryodiversity . . . . . . . . . . . . . . . . . . . . . 332
9.3.3.1. Russia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 334
9.3.3.2. Svalbard . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 335
9.3.3.3. Greenland . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 335
9.3.3.4. Canada . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 335
9.3.3.5. Alaska . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 336
9.3.4. Taxonomic structure of the Arctic bryolora . . . . . . . . . . . . . . . . . . 336
9.3.5. Large scale variation of species richness . . . . . . . . . . . . . . . . . . . . . 336
9.3.5.1. Longitudinal variation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 336
9.3.5.2. Latitudinal variation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 337
9.3.6. Origin of Arctic bryoloras and distribution types . . . . . . . . . . . . 338
9.3.7. Trends . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 339
9.3.8. Conclusions and recommendations . . . . . . . . . . . . . . . . . . . . . . . . . . 339
9.4. Algae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 339
9.4.1. Major algal groups . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 341
9.4.2. Arctic algal taxonomic diversity and regionality . . . . . . . . . . . . . . 342
9.4.2.1. Russia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 343
9.4.2.2. Svalbard . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 344
9.4.2.3. Greenland . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 344
9.4.2.4. Canada . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 344
9.4.2.5. Alaska . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 345
9.4.3. Pan-Arctic surveys . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 345
9.4.4. Trends . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 346
9.4.5. Conclusions and recommendations . . . . . . . . . . . . . . . . . . . . . . . . . . 346
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 347
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 347
»
Willows grow much faster now on the banks of Kolyma.
As well in the summer pasture areas along the Arctic
Ocean tundra willows are more plentiful and more now.
On River Suharnaya the willow bushes are much bigger.
Reindeer herders of the Chukchi community of Nutendli,
northeastern Sakha-Yakutia, Siberia; Mustonen 2007.
312
SUMMARY
Based on published scientific literature, the diversity of
plants in the Arctic is reviewed. The plants are divided
into three main groups according to essential differences in anatomy, morphology and reproduction. These
are vascular plants, bryophytes (mosses and liverworts)
and algae (micro- and macroalgae). As a whole, these
three plant groups have the ability to perform photosynthesis. As primary producers they play a key role in the
environment, since photosynthesis provides resources
for all other organisms. Vascular plants and bryophytes
(together with the lichenized fungi, the lichens) are the
main structural components of terrestrial vegetation
and ecosystems, while algae are more abundant in fresh
water and marine ecosystems.
Our knowledge of the taxonomic diversity of these three
main groups is very uneven. Although serious knowledge
gaps still exist, our understanding of vascular plant diversity in the Arctic was recently improved considerably
by the publication of the Annotated Checklist of the Panarctic
Flora (PAF) Vascular plants (Elven 2011), a result of many
years of laborious research by taxonomists associated
with the Panarctic Flora Project. The Arctic bryoflora is
relatively well known, but a circumpolar Arctic checklist of mosses and liverworts has not yet been finalized.
Knowledge of the circumpolar Arctic taxonomic diversity
of algae is still rather fragmentary. Preliminary biodiversity assessments have been made for Arctic marine algae,
but there has been no attempt yet to synthesize knowledge of the diversity of Arctic freshwater algae. Knowledge of the biodiversity of terrestrial algae in the Arctic is
also very fragmentary.
The main difficulties in assessing biological diversity at
subgeneric levels are the dissimilarities that exist in the
taxonomic species concept and classification between
the Arctic countries. Moreover, current species concepts
from traditional morphological assessments are challenged by the latest molecular phylogenetic analyses,
which sometimes conflict with traditional classification.
The vascular plant flora of the Arctic is relatively poor.
Approximately 2,218 vascular plant species (including subspecies, apomictic aggregates and some collective species)
are recognized. This is less than 1% of the known vascular
plant species in the world (c. 0.85% based on an estimated
total of 260,000 species; Raven et al. 2005). Arctic vascular plants belong to 430 genera and 91 families, almost all
within the flowering plants (angiosperms). Gymnosperms
are rare and species diversity per genus and family is low.
Species-rich families with more than 100 species include
Asteraceae (composite family), Poaceae (grass family),
Cyperaceae (sedge family), Brassicaceae (mustard family),
Rosaceae (rose family), Fabaceae (pea family), Ranunculaceae (buttercup family) and Caryophyllaceae (pink
family). The genera Carex (sedge), Salix (willow), Oxytropis
(oxytrope) and Potentilla (cinquefoil) are well represented,
with each having more than 50 species. The majority of
the Arctic species have a circumpolar distribution.
Arctic Biodiversity Assessment
The Arctic territory is divided into 21 floristic provinces
and five subzones. These strongly differ in species richness and composition. There is a pronounced increase
in species numbers from the northernmost high Arctic
subzone A (102 species) to the southernmost low Arctic
subzone E (2,180 species). A comparison of species numbers per floristic province showed a range from approximately 200 species for the rather heavily glaciated and
northern floristic province Ellesmere Land-N Greenland
to more than 800 species for Beringian W Alaska.
Polyploidy1 (allopolyploidy) levels are high in Arctic
plants. Endemism is well developed. One hundred six
species (and subspecies), or c. 5% of the Arctic vascular
plant flora, are endemic to the Arctic. The genera Papaver (poppy), Puccinellia (salt marsh-grass, goose grass),
Oxytropis, Potentilla and Draba (draba, whitlow-grass)
are particularly rich in endemic species, and almost all
endemic species are forbs and grasses, whereas there are
no endemic woody species. Though the absolute number
of Arctic endemic species increases from north to south,
i.e. from the high Arctic to the low Arctic, the relative
percentage of endemic species decreases.
The floras of the northern floristic provinces Ellesmere
Land-N Greenland, Svalbard-Franz Joseph Land and
Wrangel Island are relatively rich in Arctic endemic
species. Ten Arctic endemic species are restricted to
Wrangel Island and underline the hotspot character of
this high Arctic island. Twenty Arctic endemic species
are very rare, and as such are possibly threatened.
Borderline species are primarily non-Arctic species just
reaching the southernmost extent of the Arctic (subzone
E). Taxonomically this is a rather diverse group of 136
vascular plant species in 91 genera and 45 families.
Non-native species that occur as persisting stabilized
introductions in the Arctic account for 5% of the flora
(101 species). In addition there are 89 species native to
the Arctic that also occur as stabilized introductions in
other parts of the Arctic. In addition, more than 205
non-native species have been recorded in the Arctic only
as casual introductions that do not persist. Non-native
species mainly occur in and around settlements and
towns, in particular in climatologically favorable parts of
the Euro-Siberian Arctic.
No single, predominantly Arctic vascular plant species
is known to have gone extinct due to human activities in the last 250 years. There are no species in the
Arctic that are considered to be seriously invasive, but
some are at risk of becoming so with increasing human
traffic combined with climate change. The Arctic flora
1 Polyploidy: variations in chromosome number involving more
than the diploid number of complete chromosome sets; allopolyploidy: polyploidy resulting from hybridization of species in which there was no effective pairing of the different
sets of chromosomes preceding spontaneous doubling of the
chromosome number.
313
Chapter 9 • Plants
is considered taxonomically, ecologically, biologically
and genetically a coherent and distinctive complex of
young and dynamic species that occupy a vast natural
area characterized by a cold climate. The present Arctic
vegetation shows climate change related changes such as
greening, shrub expansion and floristical changes.
Local plants always played an essential role in the lives
and cultures of Arctic indigenous peoples. The most
useful plants have indigenous names, including not only
vascular plants, but bryophytes and algae as well.
There are an estimated 900 species of Arctic bryophytes
(mosses and liverworts). Distributional types are similar
to those observed for vascular plants. Arctic endemism is
known among bryophyte species, whereas many widely
distributed species in the Arctic show considerable morphological plasticity representing subspecies, variants
or forms. The bryoflora is in general rather uniform.
Almost 80% of the species have a circumboreal distribution. In rather stable, moist to wet sites, bryophytes
contribute substantially to vegetation biomass, and they
contribute significantly to species richness of many
vegetation types in other habitats. Very few vegetation types in the Arctic occur without bryophytes, and
single shoots occur almost everywhere, in particular in
the high Arctic. The ecosystem function of bryophytes
is poorly studied, and the bryofloras of several Arctic
regions are still incompletely known. The most speciesrich families include Bryaceae (threadmoss family),
Dicranaceae (forkmoss family), Amblystegiaceae (feathermoss family), Pottiaceae (tuftmoss family), Grimmiaceae (grimmia family), Sphagnaceae (bogmoss family),
Hypnaceae (feathermoss family), Mniaceae (thyme-moss
family), Brachytheciaceae (feathermoss family), Polytrichaceae (haircap family) and Splachnaceae (dung
moss family), which collectively account for 70% of the
total moss flora. Bryum (bryum moss), Sphagnum (bogmoss, peatmoss), Pohlia (nodding moss) and Dicranum
(forkmoss) are among the most species-rich genera.
Species-rich liverwort families include the leafy liverworts Scapaniaceae (earwort family), Jungermanniaceae
(flapwort family), Gymnomitriaceae (frostwort family),
Cephaloziaceae (pincerwort family) and Cephaloziellaceae (threadwort family), whereas Scapania (earthwort) and
Lophozia (notchwort) are prominent genera. The use of
bryophytes by indigenous peoples is very limited. There
are no known threatened bryophyte species.
Algae are ubiquitous, ecologically important and constitute the first layer of marine and freshwater food
webs. They occur either free floating in the upper water
column (pelagic), associated with sea ice (sympagic),
or attached to bottom substrates (benthic). Phaeophyta
(brown algae) range in size from less than 2 µm to more
than 100 m long in giant kelps. Pelagic algae, known as
phytoplankton, and sea ice algae are autotrophic, singlecelled eukaryotes ranging in size from 0.2 to 200 µm.
Benthic algae mainly refer to marine macroalgae characteristic of coastal regions, but also include microalgae
attached to various substrates along the seashore. Algae,
including the autotrophic prokaryote cyanobacteria
(blue-green algae), are classified into different groups or
phyla, depending on the classification system used.
The following groups have been recognized in this
review: (1) Archaeplastida, including Chlorophyta
(green algae), Streptophyta, Glaucophyta, Rhodophyta
(red algae), (2) Chromalveolata, with Cryptophyta,
Haptophyta, Dinophyta, Stramenopiles (including
Dictyochophyceae, Eustigmatophyceae, Pelagophyceae,
Bacillariophyta (diatoms), Phaeophyceae (brown algae),
Xanthophyceae, Chrysophyceae (yellow-green algae),
Rhaphidophyceae), (3) Excavata (Euglenophyta), (4)
Opisthokonta (Choanoflagellida), (5) Rhizaria (Chlorarachniophyta) and (6) Cyanophyceae (blue-green algae).
There is a conservative estimate of 4,000 algal species
reported from the circumpolar Arctic, including both
freshwater and marine habitats. The species diversity of
microalgae and cyanobacteria for the Arctic is still largely
unknown, especially in terrestrial and freshwater environments, but it is assumed to be much lower than in warmer
regions of comparable size. In Arctic regions, marine
diatoms are very diverse and abundant in annual sea ice,
pelagic waters and benthic environments. Recent molecular studies reported a high diversity in the smallest-sized
fraction of the phytoplankton in polar regions, frequently
contributing to more than 50% of total phytoplankton
biomass and production. In the western Canadian Arctic
alone, 10,000 species of single-celled phytoplankton species were documented through molecular analyses, at least
half of which are likely autotrophic. There are c. 200-215
seaweed (macroalgae) taxa in the Arctic, with endemism
poorly developed. A major challenge facing biodiversity
assessments will be matching morphology of a singlecelled alga to a given gene sequence, which will require
development of better sampling strategies and culture
techniques for these small-sized microalgae.
9.1. INTRODUCTION
This plant chapter deals with the taxonomical biodiversity of organisms that are able to perform photosynthesis.
They use light energy for conversion of carbon dioxide
and water into chemical energy in the form of sugar and
other organic substances under release of oxygen. Most
of them are autotrophic, using carbon dioxide as their
carbon source.
They include three main groups based on differences in
anatomy, morphology, physiology and reproduction, and
phylogenetic relationships.
The kingdom Plantae of the eukaryotic life domain
comprises the green land plants. These are the vascular plants – Tracheophyta (Section 9.2) and the bryophytes – Bryophyta (Section 9.3). The vascular plants
are subdivided into spore-producing plants (clubmosses
– Lycopodiophyta and ferns – Pteridophyta) and seedproducing plants (Gymnospermae with uncovered seeds,
314
Angiospermae with covered seeds). The bryophytes are
divided into the hornworts (Anthocerophyta), liverworts
(Hepatophyta) and mosses sensu stricto (Bryophyta) (Raven et al. 2005).
The autotrophic algae (Section 9.4) of the kingdom
Protista comprise eukaryotic organisms which cannot
be attributed to the kingdoms Fungi, Plantae or Animalia. The green algae (Chlorophyta) are ancestral to the
algal Streptophyta and hence to the kingdom Plantae,
the bryophytes – Bryophyta and vascular plants – Tracheophyta. Some other algae are both autotrophic and
heterotrophic (Poulin et al. 2011).
The blue-green algae belong to the prokaryotic life domain Bacteria and are classified as Cyanobacteria (Raven
et al. 2005).
As primary producers, all groups play a key role in the
environment, since photosynthesis provides resources
for all other organisms. Vascular plants and bryophytes
(together with the lichenized fungi, the lichens; see
Dahlberg & Bültmann, Chapter 10) are the main structural components of terrestrial vegetation and ecosystems, while algae are more abundant in freshwater and
marine ecosystems.
The state of knowledge of Arctic vascular plants, bryophytes, and algae differs among countries, regions, and
floristic provinces, and there remain many differences
in taxonomic opinions among botanists on different
continents. The data presented here should be viewed as
a preliminary assessment.
Scientific names are used throughout the manuscript since
there are no standardized common or vernacular names
for plants, and many species (e.g. algae) lack common
names altogether. For taxa with common names, these are
provided in parentheses following the scientific names the
first time a taxon is mentioned. These names are derived
from several sources (among others Clapham et al. 1962,
Böcher et al. 1968, Hultén 1968, Porsild & Cody 1980,
Rønning 1996, Smith 2004 and Edwards 2012).
The total land surface of the Arctic is estimated at 7.11
million km2, with an estimated 5 million km2 covered by
vegetation; the remainder is ice-covered (Walker et al.
2005). The Arctic territory has been and still is sparsely
populated. While there was almost no impact by human
populations on Arctic flora and vegetation prior to the
1960s, human impacts now pose an increasing threat in
certain areas. Nevertheless, these impacts are minor compared with human impacts in the adjacent boreal zone.
9.2. VASCULAR PLANTS
The assessment presented here is largely based on the
Checklist of Panarctic Flora (PAF) Vascular Plants compiled
and edited by Reidar Elven and colleagues, who provided
the first checklist and taxonomic assessment of all Arctic
Arctic Biodiversity Assessment
vascular plant species. The draft version from May 2007
was made available to us in March 2009 and is cited here
as Elven (2007) and PAF. It is basically congruent with
the more recent (and slightly updated) on-line version,
the Annotated Checklist of the Panarctic Flora (PAF) Vascular
Plants (Elven 2011). This first checklist covering all Arctic vascular plants is a working list that will continue to
undergo modifications as new knowledge about species
accumulates and taxonomic problems are resolved.
9.2.1. Taxonomic categories and species
groups
The present assessment of Arctic vascular plant biodiversity uses the definitions of the Arctic area and the
taxonomic concept presented in the PAF (Elven 2007).
It includes 2,218 recognized species and subspecies, as
well as several ‘apomictic aggregates’ (used for Taraxacum, dandelion and Hieracium, hawkweed) and a few
‘collective’ species (for convenience these will all be
referred to as species here) (Appendix 9.1). In addition,
some hybridogenous taxa that are reproductively isolated
from their parents mostly by ploidy level are treated as
species. More than 600 named entities (at the species
level or below) are not accepted as distinct entities in the
PAF, thus they are not considered here. The assessment
of Arctic vascular plant diversity presented here should
be regarded as preliminary; it is based largely on the PAF
table of species distribution and frequency of occurrence
within floristic provinces and subzones. The exact number of vascular plant species in the Arctic has not been
settled mostly due to the difficulty of determining species boundaries in taxa that frequently hybridize and/or
are primarily clonal, but also due to lack of knowledge
and taxonomic expertise for some groups and due to the
occurrence of cryptic species.
All species considered in this chapter occur somewhere
in the Arctic territory – the definition used here approximates but is not completely congruent with its
delineation on the Circumpolar Arctic Vegetation Map
(CAVM) (compare Fig. 9.1 and 9.2; see also Section 2 in
Meltofte et al., Chapter 1). They are classified as native
or non-native (Elven 2007). Native species include all
those that are naturally established in areas within the
Arctic. Native species include Arctic endemics which
are restricted in their distribution to areas within the
Arctic territory (floristic provinces and subzones A-E
of the PAF), species that also occur outside the Arctic
(typically Arctic-alpine species, which are also found in
alpine areas of the neighboring non-Arctic regions to the
south, subzone N of the PAF, which is defined as boreal
and/or boreal-alpine), and ‘borderline species’ (mainly
non-Arctic species just reaching the southernmost extent
of the Arctic in subzone E). We also include non-native
species with stabilized populations within the Arctic
territory. Most of these species reached the Arctic as a
result of human activities after approximately 1700 A.D.
We follow the taxonomic nomenclature as provided
in the PAF. Numbers and calculations are based on all
Chapter 9 • Plants
accepted 2,218 species. For our calculations of species
richness for Arctic floristic provinces and subzones we
do not include species with uncertain occurrence, which
are indicated with “?” in these areas in the PAF. While
the PAF made a concerted effort to reconcile the different taxonomic concepts across the Arctic, there remain
some regional differences. How well a flora is known
also varies among Arctic regions. Both problems likely
affect the statistics presented here, in particular comparisons of species diversity across floristic provinces.
9.2.2. The Arctic territory and its subdivision
The delineation of the Arctic has been interpreted differently by various authors (Yurtsev 1994, Elvebakk et
al. 1999, Nordal & Razzhivin 1999, CAVM Team 2003,
Walker et al. 2005 and PAF). We follow the PAF delineation, which is largely derived from the Circumpolar
Vegetation Map (CAVM Team 2003 and Walker et al.
2005). In areas with continental climate, the Arctic is
considered to be the area north of treeline, just as in the
CAVM. However, areas with extra-zonal occurrences
of small pockets of trees in places with a distinct winter
frost climate and mean July temperature above 10°C,
such as small areas in the inland of S Greenland, are also
included in the PAF. On the CAVM, this small area in S
Greenland is excluded from the Arctic. Further, many
areas with an oceanic climate in the North Pacific and
North Atlantic are lacking trees, but have higher mean
Figure 9.1. Bioclimatic subzones
of the Arctic territory according
to the Circumpolar Arctic Vegetation Map (CAVM Team 2003,
Walker et al. 2005).
315
July temperatures and a less pronounced winter frost period. They are excluded from the Arctic in the CAVM,
as are the vegetation belts of mountains in the neighboring boreal zone to the south. Thus we follow here the
concept of including all of Greenland as belonging to the
Arctic as proposed in PAF, and excluding those treeless
areas in the North Pacific and North Atlantic regions
that are included in the non-Arctic boreal zone.
The Arctic territory is roughly subdivided along two
main axes in latitudinal subzones (Fig. 9.1) and longitudinal floristic provinces (Fig. 9.2). The latitudinal northsouth axis mainly reflects the present climate gradient
divided into five different subzones, which are separated
according to climate and vegetation in the lowlands of
each zone.
There is not consensus regarding a uniform nomenclature of the subzones, and in the following assessment
we use the letters: A, B and C (for the high Arctic) and
D and E (for the low Arctic) (CAVM Team 2003, Walker et al. 2005). However, subzone A might be appropriately named the Arctic herb subzone; B, the northern
Arctic dwarf shrub subzone; C, the middle Arctic
dwarf shrub subzone; D, the southern Arctic dwarf
shrub subzone and E, the Arctic shrub subzone (Daniëls
et al. 2000, CAVM Team 2003, Walker et al. 2005).
The latitudinal extent of the subzones corresponds
approximately with altitudinal vegetation belts in the
316
Arctic Biodiversity Assessment
Figure 9.2. Map of species richness and endemicity of Arctic vascular plant loras in loristic provinces of the Arctic. Species richness of the loristic provinces is expressed as percentage of the total species richness of the Arctic (2,218 species), and species endemicity of the loristic provinces as rounded of percentage of the total number of Arctic endemic species (106). Floristic provinces and subzones according to Elven (2007).
Arctic mountains. Accordingly, the lower mountain belt
(named d) in subzone E corresponds with subzone D,
the middle belt (named c) with subzone C, the upper
belt (b) with subzone B, and the highest belt (a) with
subzone A (Walker et al. 2005, Sieg et al. 2006).
The longitudinal east-west axis reflects different conditions in the past such as different plant chorological
histories (i.e. history of spatial distributions) related
to glaciations, land bridges and north-south-trending
mountain ranges, resulting in the delineation of 21
floristic provinces (Fig. 9.2; Yurtsev 1994, PAF, see also
CAVM Team 2003 and Walker et al. 2005).
9.2.3. The lora of the Arctic
The vascular plant flora of the Arctic is relatively poor
in species (see Fig. 1.1 in Meltofte et al., Chapter 1). Our
review suggests that 2,218 species occur in one or more
subzones and floristic provinces of the entire Arctic territory. This is less than 1% of the estimated number of
all vascular plant species of the world (Chapman 2009).
The mean diversity of Arctic vascular plant flora is estimated to be 100-200 species per 10,000 km2 (Barthlott
et al. 1996).
317
Chapter 9 • Plants
Figure 9.3. Characteristics of the Arctic vascular plant lora.
Fabaceae (109)
Cyperaceae (190)
Figure 9.4. Distribution types and plant functional types of the
106 Arctic endemic vascular plant species.
European-west Siberian (3)
Central Siberian (7)
Canadian (7)
Caryophyllaceae (100)
Ranunculaceae (102)
Misc. families
(978 species)
Rosaceae (128)
Beringian (39)
North Atlantic
(11)
Brassicaceae (133)
Scattered (16)
Asteraceae (254)
Poaceae (224)
Circumpolar (23)
Climatological distribution types
• Low diversity, species number less than 1% of the world lora
• 91 families, 430 genera, 2,218 species
High Arctic (8)
• Few gymnosperms, angiosperms represent 96% of the lora
• Low species diversity per family, with 8 species-rich families
accounting for > 50% of the lora (see above)
Arctic (59)
Low Arctic (39)
• Low species diversity per genus, with most species-rich genera
Carex (152), Salix (72), Oxytropis (58), Potentilla (50), Draba (41),
Ranunculus (40), Papaver (39), Poa (36), Saxifraga (35), Artemisia
(33), Puccinellia (31), Juncus (30), Pedicularis (29), Rumex (28), Astragalus (28), Silene (24), Erigeron (24), Luzula (22) and Eriophorum (18)
• Around 5% endemic species, mainly Poaceae (24), Papaveraceae
(15), Brassicaceae (15), Fabaceae (14) and Asteraceae (6),
accounting for 70% of the endemic species lora
• Genera with relative high numbers of endemic species include
Papaver (15), Puccinellia (13), Oxytropis (11), Potentilla (8) and Draba
(8), accounting for 52% of the endemic species lora. Almost all are
forbs or grasses, a few sedges, woody species absent
Plant functional types
Sedges (2)
Grasses (24)
• Coherent with high percentage of circumpolar species
• High polyploidy (allopolyploidy) levels, in particular in the northern
and longer glaciated areas
• Young with distinct phytogeographic history
• No strong inluence by aliens
Forbs (80)
• Still native and with intact historical plant geography
9.2.3.1. Taxonomic structure
The 2,218 Arctic species included in the PAF (Appendix
9.1) are assigned to 430 genera in 91 families. The mean
number of species per family is 24.4 and per genus 5.2;
the mean number of genera per family is 4.7. Of the
2,218 recognized Arctic species, 106 are Arctic endemics (5%, Fig. 9.3 and 9.4, Appendix 9.2), 136 (6%) are
‘borderline species’ (Appendix 9.3), and 101 (4.8%) are
introduced stabilized non-native species (Appendix 9.4).
Spore-bearing vascular plants play a very minor role in
the Arctic vascular plant flora. They represent ferns and
fern allies (Pteridophyta) and lycophytes (Lycopodiophyta) and comprise only 12 families with 21 genera and
72 species. They represent 13, 5 and 3%, respectively,
of these taxonomic categories in the entire Arctic flora.
Included here are the Lycopodiaceae (clubmosses, 14
species), Selaginaceae (spikemosses, 3), Isoëtes (quillworts, 3), Equisetum (horsetails, 12) and ferns (various
families, 40). In contrast, seed-plants (Spermatophyta)
are represented by 79 families, 409 genera and 2,146
species. They comprise gymnosperms (Gymnospermae,
seed-plants with naked seeds not enclosed in an ovary)
and flowering plants (angiosperms, Angiospermae,
seed-plants with covered seeds enclosed in an ovary).
The gymnosperms are represented in the Arctic flora by
two families, five genera and 16 species representing 2,
1 and 1%, respectively, of these categories in the total
flora. Gymnosperms are thus a minor contributor to the
diversity of the Arctic flora (Fig. 9.3), a fact considered a
prominent feature of the Arctic flora (Yurtsev 1994).
Flowering plants comprise 77 families, 404 genera and
2,130 species, representing 85, 94 and 96% respectively
of the total flora. The basal angiosperms (including the
318
magnoliids and Ceratophyllaceae, horn wort family) are
very poorly represented with only two families (Nymphaeaceae, water lily family and Ceratophyllaceae, hornwort family), three genera and six species. The monocots
(Monocotyledoneae) include 18 families, 102 genera and
566 species, representing 20, 24 and 26%, respectively,
of these taxonomic categories in the flora. The eudicots
(sensu Angiosperm Phylogeny Group 2009) are represented by 57 families with 299 genera and 1,558 species.
They represent 63, 69 and 70%, respectively, of the
diversity of these taxonomic categories in the entire flora.
The most prominent families in terms of species numbers are the Asteraceae (composite family, 254 species),
the Poaceae (grass family, 224), the Cyperaceae (sedge
family, 190), the Brassicaceae (mustard family, 133), the
Rosaceae (rose family, 128), the Fabaceae (pea family,
109), the Ranunculaceae (crowfoot or buttercup family,
102) and Caryophyllaceae (pink family, 100). With 1,240
of 2,218 species, these eight families represent 56% of
the species of the entire Arctic flora (Fig. 9.3). Only 21
families (23% of 430) have more than 24 species, which is
the mean species number per family. Sixty percent of the
families have fewer than 10 species. The relatively low
species diversity per family is considered another characteristic feature of the Arctic flora (Yurtsev 1994).
The most species-rich genera are Carex (sedge, with 152
species), Salix (willow, 72), Oxytropis (58), Potentilla (50),
Draba (41), Ranunculus (crowfoot, buttercup, 40), Papaver
(poppy, 39), Poa (bluegrass, meadow-grass, 36), Saxifraga
(saxifrage, 35), Artemisia (wormwood, 33), Puccinellia
(alkali grass, 31), Juncus (rush, 30), Pedicularis (lousewort, 29), Rumex (sorrel, dock, 28), Astragalus (milk
vetch, 28), Silene (campion, 24), Erigeron (fleabane, 24),
Luzula, (wood rush, 22) and Eriophorum (cotton grass,
18). Together they account for 38% of the total species
diversity (Fig. 9.3). Overall species diversity of genera
is rather low: 337 genera (78%) have 1-5 species. Only
93 genera (22%) have more species, accounting for 70%
of the total species number. The relatively low species
diversity per genus is considered another typical feature
of the Arctic flora (Yurtsev 1994).
9.2.3.2. Endemic species
The Arctic flora includes 106 species and subspecies endemic to the Arctic. Several of them are ‘restricted range
species’, i.e. species with a distribution in an area of less
than 50,000 km2 (e.g. Puccinellia svalbardensis Svalbard
alkali-grass). These 106 species account for about 5% of
the entire flora (Appendix 9.2, Fig. 9.4). They mainly
belong to Poaceae (24), Papaveraceae, the poppy family
(15), Brassicaceae (15), Fabaceae (14) and Asteraceae (6),
accounting for 70% of the entire endemic species flora.
Genera with relatively high numbers of endemic species
include Papaver (with 15 species), Puccinellia (13), Oxytropis
(11), Potentilla (8) and Draba (8). Together these genera
account for 52% of the endemic species flora. The genera
Poa and Braya (braya, rose cress) each has four endemic
species, and Festuca (fescue grass), Ranunculus and Saxifraga each has three endemic species in the Arctic. Almost
Arctic Biodiversity Assessment
all of these species are forbs (non-graminoid herbs) or
grasses (Fig. 9.4). There are few sedges, and woody species are absent among Arctic endemics. The Beringian
and circumpolar distribution types dominate (Fig. 9.4).
The majority of species occur in both the low and high
Arctic, with few restricted to the high Arctic (Fig. 9.4).
The majority of endemic species show high polyploidy (allopolyploidy) levels; of the 75 species with known ploidy
level, 31% are tetraploid and 45% are higher polyploids
(Appendix 9.2). Polyploidy is a prominent feature of the
entire Arctic flora, in particular in the northern and
longer-glaciated North Atlantic region (e.g. Brochmann et
al. 2004, Solstadt 2008).
9.2.3.3. Borderline species
Borderline species (Appendix 9.3) are primarily nonArctic species that just reach the southernmost extent
of the Arctic (subzone E). This group of species is rather
diverse, and includes 190 species representing 6% of all
Arctic species, 91 genera and 45 families. Many ‘borderline’ species belong to the Asteraceae (16 species;
12%) or Cyperaceae (15 species; 11%). Hydrophytes and
other species associated with wet habitats are frequent in
this group. Seven aquatic genera (Nymphaea, water lily;
Nuphar, pond lily; Alisma, water-plantain; Sagittaria, arrowhead; Butomus, flowering rush; Scheuchzeria, rannochrush; and Potamogeton, pondweed) account for 14 species
(10%), with the genus Potamogeton represented by six
species (4%). The borderline species group also includes
some shrub and tree species of the genera Salix, Alnus
(alder) and Abies (fir) that are common in boreal and
temperate regions.
9.2.3.4. Non-native species
Introduced non-native plants may be divided into two
groups, stabilized introductions and casual introductions. Stabilized introductions are considered to be
self-sustaining somewhere in the Arctic for at least one
generation by generative or vegetative reproduction,
whereas casual introductions are species that are present
for short periods of time but do not persist (Elven 2007,
2011). One hundred and one non-native species (5% of
the flora) are considered to be stabilized introductions
in the Arctic (Appendix 9.4). In addition, there are
89 species (4%) that are native to one or more Arctic
floristic provinces and subzones, but that are also found
as non-native introduced species elsewhere in the Arctic.
Of these 89 native species, 45 are found as stabilized
introductions, 20 as casual introductions, and 24 as both
stabilized and casual introductions in at least one floristic
province or subzone. A total of 170 (8%) native and nonnative species are considered as stabilized introductions
somewhere in the Arctic. Including the native Arctic
species also present as casual introductions in some parts
of the Arctic increases this number to 190 species.
The group of non-native stabilized introductions is
taxonomically diverse; however, Poaceae and Asteraceae
account together for 33% of the stabilized introduced
319
Chapter 9 • Plants
flora (Appendix 9.4). The most widespread non-native
stabilized introduced species are Lepidotheca suaveolens
(pineapple weed, stabilized introduction in 10 floristic
provinces), Plantago major ssp. major (common plantain,
stabilized in nine, casual in two) and Trifolium pratense
(red clover, stabilized in eight). The native Arctic species that occur most widely as stabilized introductions
include Trifolium repens (white clover, stabilized in 10,
casual in one, native in two), Puccinellia hauptiana (european alkali grass, stabilized in nine, native in three),
Poa pratensis ssp. pratensis (meadow-grass or Kentucky
bluegrass, stabilized in eight, native in 11), Stellaria media
(common chickweed, stabilized in eight, casual in one,
native in five) and Draba nemorosa (woodland draba, stabilized in eight, native in three).
Elven (2011) lists an additional 205 non-native species
that are known to occur in the Arctic only as casual
introductions. As these species are not listed according to floristic regions in the main checklist, they are
not included in the analyses presented here. They range
from species reported only once to those that may appear regularly around settlements. The majority of these
species are annual or biennial plants that are unable to
reproduce in the Arctic. Others are perennials that are
unable to survive the harsh Arctic climate. The number
of casual introductions listed in PAF may be highly underestimated: a throughout review of casuals in Svalbard
revealed about 100 species (I.G. Alsos & R. Elven, unpubl.) whereas PAF lists 53 species for this region.
Although there is little information about non-native
invasive species in the Arctic, the majority of introduced
species appear not to be invasive. Several native plants
have become weedy in disturbed habitats, either in their
native range or elsewhere in the Arctic, but are not
considered to be a threat to the native vegetation. Lupinus
nootkatensis (Nootka lupine), native to NW North America, was introduced as an ornamental in Greenland and
has become a stabilized weedy species in SW Greenland,
but is not considered to be seriously invasive since it is
mostly restricted to disturbed sites (C. Bay and K. Høegn,
pers. com. 2012). In Iceland it is a threat, including in
the Arctic area of Iceland (Magnusson 2010). Hordeum
jubatum (foxtail barley) is a troublesome native weed in the
western Arctic of North America, but again is primarily restricted to disturbed sites and thus not considered a
threat to native vegetation. Invasive species were not noticed in many areas in Greenland (F.J.A. Daniëls, unpubl.)
and the Canadian Arctic (L.J. Gillespie, unpubl.). Daniëls
& de Molenaar (2011) and Daniëls et al. (2011) did not
observe such species in the tundra near the town of Ammassalik, SE Greenland during their fieldwork in the last
40 years, although in the town, a few casual non-invasive
introductions were occasionally recorded. In Svalbard,
the majority of introduced species have been considered as
no risk, but recently several species were evaluated as low
risk (watch list) and one as high risk (black list) of becoming invasive in Svalbard (Gederaas et al. 2012). Anthriscus
sylvestris (cow parsley) was rather recently established
(after 1988) (Liška & Soldán 2004), but the population
is now large with fertile individuals about to 2 m high
(Alsos et al. 2012b). Any spread of this species to bird cliffs
would pose a threat both to the many redlisted species
found there and to the birds as the foxes could hide during
hunting. Although not currently a problem in the Arctic,
invasive species are likely to increase in the Arctic due
to the expanding visitation rates combined with climate
warming. Each visitor to Svalbard transport on average
a minimum of four seeds, many of them from species
known to be invasive in other regions, and 26% of them
are able to germinate under current Arctic climate (Ware
et al. 2012).
9.2.3.5. Origin and integrity of the Arctic lora
The present-day Arctic flora is of relatively recent origin
(Murray 1995) and has been shaped through numerous
large-scale climate changes resulting in cycles of fragmentation, range expansion and reunion of previously
isolated populations (Stebbins 1984, 1985). During most
of the Tertiary (65.2 million years ago) forests grew at
high latitudes, e.g. in Canada and Greenland (McIver &
Basinger 1999), and tundra did not appear until the late
Pliocene when global temperatures dropped (Matthews
& Ovenden 1990). Initially, tundra was distributed
discontinuously, but became continuous by three million
years ago (Matthews 1979). The early Quaternary flora
was likely recruited from the Arcto-Tertiary forest and
immigrants of ancestral stocks from temperate high
mountain ranges in Asia and North America (Hultén
1937, Hedberg 1992, Murray 1995, Ickert-Bond et al.
2009). This floristic mixture has since repeatedly been
re-arranged and re-mixed spatially by more than 20
cycles of glacials and interglacials during the Quaternary
period (reviewed in Birks 2008; see also Payer et al.,
Chapter 2).
During the last two decades, comparative molecular research on Arctic plant populations has contributed significantly to a better understanding of patterns and processes
in the present Arctic flora (cf. Abbott et al. 2000, Abbott
& Brochmann 2003, Abbott & Comes 2003, Brochmann
et al. 2004, Alsos et al. 2005, 2007, Grundt et al. 2006,
Solstadt 2008, Tkach et al. 2008, Consaul et al. 2010,
Hoffmann et al. 2010, Westergaard et al. 2010, 2011a,
2011b, Hoffmann 2011). Abbott & Brochmann (2003) reviewed fossil, molecular and phytogeographical evidence
for the existence of Beringia as a major glacial refugium
for Arctic plants as previously proposed by Hultén
(1937), and concluded that the evidence is excellent to
support his proposal (e.g. for Dryas integrifolia, entire-leaf
mountain avens and Saxifraga oppositifolia, purple saxifrage; Tremblay & Schoen 1999, Abbott et al. 2000).
Recently, molecular evidence supporting in situ glacial
survival in the North Atlantic region has been reported
for Saxifraga rivularis (brook saxifrage), Arenaria humifusa
(low sandwort), Sagina caespitosa (tufted pearlwort) and
Carex ruina (reddish sedge) (Westergaard et al. 2010,
2011a, 2011b). However, molecular evidence supports
several long-distance dispersal events across great dis-
320
tances in North America, and between North America,
Greenland and Europe, thus not corroborating Hultén’s
(1937) hypothesis that the North Atlantic was a strong
barrier for plant dispersal in the Holocene.
The extreme Beringian/Atlantic disjunction in Saxifraga
rivularis has evidently formed at least twice, with expansions out of Beringia to the Atlantic regions both before
and after the last glaciation (Westergaard et al. 2010). A
long-distance dispersal event of similar magnitude was
also reported for the Beringian species Arenaria longipedunculata (long-stemmed sandwort), in which molecular
and morphological data revealed a highly disjunct occurrence in W Greenland (Westergaard et al. 2011b). Postglacial, trans-Atlantic dispersal has been reported for
an increasing number of Arctic and Arctic-alpine plant
species, even for species lacking obvious morphological
adaptations to long-distance dispersal (see Abbott & Brochmann 2003, Brochmann et al. 2004, Alsos et al. 2007,
Westergaard et al. 2011a, 2011b). Although the Arctic
flora has long been viewed as depauperate and speciespoor, Grundt et al. (2006) showed how three circumpolar species of recent origin, Draba ladnizensis (white
Arctic draba), D. nivalis (snow draba) and D. subcapitata
(hemispherical draba), actually consist of many cryptic
species separated by genetically based crossing barriers,
but not morphologically and ecologically differentiated.
Frequency of polyploids is particularly high in the Arctic, and both frequency and ploidy level strongly increase
northwards within the Arctic (Brochmann et al. 2004).
A large number of species show high ploidy levels (mostly allopolyploids), particularly in northern areas and
areas more recently glaciated such as the North Atlantic
(Abbott & Brochmann 2003, Brochmann et al. 2004).
A majority of plants are of hybrid origin, many of them
between plants which themselves are, or were, of hybrid
origin (reticulate evolution). Most of these hybrids have
been stabilized via polyploidy. However, not all Arctic polyploids have been formed in the Arctic (Murray
1995). Successive cycles of divergent evolution among
populations isolated in different glacial refugia, migration into deglaciated terrain, hybridization and polyploidy have built up increasingly intricate, high-ploidy
complexes (Abbott & Brochmann 2003). Brochmann et
al. (2004) found that for ‘Arctic specialist taxa’ (mainly
Arctic and exclusively Arctic species with limited distribution) the frequency of polyploids appears much lower
in Beringia, which was largely unglaciated during the
last ice age, than in the heavily glaciated North Atlantic
area. This was interpreted as an indication that polyploids are more successful in colonizing ice-free areas
after deglaciation than diploids are. The evolutionary
success of polyploids in the Arctic may be based on their
fixed-heterozygous genomes, which may buffer against
inbreeding and genetic drift through periods of dramatic
climate change (Brochmann et al. 2004). Another aspect
of the success of polyploids is their broader ecological
amplitude and thus greater ability to cope with a changing climate and adapt to more diverse ecological niches
than a diploid could (Brochmann et al. 2004).
Arctic Biodiversity Assessment
Asexual reproduction commonly occurs in the Arctic
flora and is demonstrated through apomictic seed production (e.g. dandelion Taraxacum; hawkweed Hieracium),
pseudovivipary (e.g. viviparous alpine bluegrass Poa alpina
ssp. vivipara), bulbils formation (e.g. nodding saxifrage
Saxifraga cernua, leafy saxifrage Micranthes foliolosa) and
vegetative spread through stolons and tillers (e.g. Bigelow’s sedge Carex bigelowii, creeping alkaligrass Puccinellia phryganodes, spider saxifrage Saxifraga platysepala).
However, sporadic sexual reproduction is still maintained, likely to avoid loss of genetic diversity. Seed banks
are persistent and in several cases long-term viability and
genetic differentiation among seeds could be demonstrated (summarized from Crawford 2008a; see also Section
2.3.5, Ims et al., Chapter 7 and Cook, Chapter 17).
Annual herbs and trees are almost absent from the Arctic
biome. The cold, dark and long winter and short summer
with mean July temperature < 10 °C allow only very
few annual species to complete their life-cycles within
the year (e.g. Iceland purslane Koenigia islandica and the
mastodon plant Tephroseris palustris var. congesta). Most
species are perennial without specific Arctic life-history
traits (Jónsdóttir 2011). Tall aerial plants such as trees
(phanerophyte life-form) are absent due to harsh climatological conditions. The low hemicryptophyte life-form
(half-earth plants, e.g. grasses, many forbs) strongly
dominates the life-form spectrum, followed at a distance
by representatives of the chamaephyte life-form (surface
plants, e.g. dwarf shrubs) and geophytes (earth plants,
e.g. orchids and several other forbs). All these survive
harsh winter conditions near or in the soil, often under
snow cover. Hence, compared with the other Earth
biomes the Arctic flora is characterized by the absence of
trees (phanerophytes), a few annual plants (therophytes)
and the predominance of hemicryptophytes (forbs and
graminoids) (cf. Polunin 1967). In terms of strategy types
(Grime 2001), stress-tolerators (which are adapted to resist all conditions that restrict photosynthesis production)
predominate, whereas ruderals, plants adapted to habitats
where disturbance (destruction of biomass) frequently
occurs, are almost absent.
Despite the slightly different concepts of the Arctic
flora and territory used here, the results of our taxonomic examination of the Arctic vascular plant flora still
support Yurtsev’s (1994) view that the integrity of the
Arctic flora is high, and therefore the Arctic deserves
the status of its own floristic region. The identity of the
Arctic vascular plant flora is based on several distinct
features pertaining to taxonomic structure, distribution
(e.g. endemism, circumpolar distribution), ecology and
morphology (growth forms) and flora genesis and speciation (cf. Yurtsev 1994). There are relatively few species
per genus and family in the Arctic, respectively five and
24 on average (Fig. 9.3). Several unique floristic characteristics support the Arctic flora: 60% of the families
have less than 10 species, gymnosperms are poorly
represented, percentage of endemism is relatively high
(5%) and the percentage of non-native stabilized introduced species is relatively low (5%). The proportion of
321
Chapter 9 • Plants
Frequent
species
Present,
abundance unknown
Present,
abundance likely rare
Present, frequency likely
scattered or sparse
Species for which presence
is uncertain
Total species number
without uncertain occurrencies
52
18
239
165
14
151
3
0
0
22
642
Polar Ural-Novaya Zemlya
UN
32
8
209
228
5
137
6
0
0
24
625
Yamal-Gydan
YG
16
4
238
129
23
96
7
0
0
29
513
TM
23
16
310
121
35
136
4
0
0
38
645
Rare species
Mean species number
per loristic province group
Borderline
species
KP
Floristic province
Casual introduced species
Kanin-Pechora
Number of stabilized
introduced species
Species with a scattered
distribution
Table 9.1. Summary of Arctic vascular plant species and distribution by Arctic lora province and subzone based on Elven (2007). Arctic
loristic provinces, subzones (A-E), neighbouring boreal or boreal-alpine zone (N) and distribution derived from Elven (2007).
European Russian-W Siberian
593
E Siberian
Taimyr-Severnaya Zemlya
Anabar-Olenyok
AO
1
0
200
144
14
67
2
0
1
37
429
Kharaulakh
Kh
14
3
184
118
13
223
3
0
0
20
558
Yana-Kolyma
YK
3
0
280
80
12
53
3
0
1
46
432
516
Beringian
W Chukotka
CW
8
4
219
151
2
238
14
0
0
28
636
Wrangel Island
WI
0
1
89
76
0
145
4
0
0
16
315
S Chukotka
CS
7
2
236
146
9
136
13
0
0
33
549
E Chukotka
CE
20
4
225
140
0
265
14
0
0
36
668
W Alaska
AW
20
14
316
212
50
210
3
0
0
27
825
N Alaska-Yukon Territory
AN
11
3
247
172
40
256
3
0
0
26
732
Central Canada
CC
8
3
303
157
30
137
2
0
0
36
640
Hudson Bay-Labrador
HL
30
9
355
174
58
141
1
1
0
27
769
Ellesmere Land-N Greenland
EP
0
0
77
46
0
76
0
0
0
5
199
GW
50
26
144
174
0
159
1
0
0
18
554
621
Canadian
536
N Atlantic
W Greenland
E Greenland
GE
5
4
136
105
0
141
0
0
0
11
391
N Iceland-Jan Mayen
Ic
52
2
80
74
17
211
0
0
0
16
436
N Fennoscandia
FN
63
13
216
157
28
171
0
1
0
23
649
Svalbard-Franz Joseph Land
SF
4
32
53
47
0
79
0
0
0
10
215
A
0
0
41
23
0
38
0
0
0
11
102
449
Subzone
Arctic herb subzone
N Arctic dwarf shrub subzone
B
0
0
91
39
0
90
0
0
0
18
220
Middle Arctic dwarf shrub subzone
C
3
27
204
85
0
188
0
0
0
91
507
S Arctic dwarf shrub subzone
D
18
10
349
213
0
389
1
0
0
65
980
Arctic shrub subzone
E
101
0
868
392
136
681
2
0
0
4
2,180
N
34
0
195
563
0
1,304
1
0
0
13
2,097
Non-Arctic-Boreal or Boreal-alpine
endemic species is very high considering the short period
since the latest glaciation. The PAF analysis also shows
that 14 species are found in all five latitudinal subzones
and 21 floristic provinces around the Arctic, whereas
many others have a distinctive circumpolar distribution,
occurring in all geographical floristic province groups
(Appendix 9.1). In addition, 76 Arctic non-endemic
species, nearly 4% of all Arctic species, occur in all five
subzones. Although not analyzed here, there is no reason
to doubt that circumpolar species account for 35% to
over 80% of the species in local floras (Yurtsev 1994).
The Arctic flora is young and has its own distinct natural
phytogeographic history (Elven 2011). No single, predominantly Arctic vascular plant species is known to
have gone extinct due to human activities in the last 250
322
Arctic Biodiversity Assessment
years (Elven 2011), nor is the Arctic strongly influenced
by invasive species. Unlike much of the rest of the world,
the Arctic’s native flora and plant communities are still
intact (Elven 2011).
(small-flowered draba), D. simmonsii (Simmons’ draba),
D. oblongata (Canada Arctic draba), Cerastium arcticum
(Arctic chickweed), Minuartia rossii (Ross’ stitchwort or
cushioned sandwort), Silene uralensis ssp. arctica (polar
campion) and S. sorensensis (Sorensen’s campion).
9.2.3.6. Species richness in Arctic subzones
The 136 borderline species are – not unexpectedly – all
confined to subzone E, with their main distribution
remaining outside the Arctic (Appendix 9.3).
The five Arctic subzones strongly differ in species richness
and species composition. There is a pronounced increase
in the number of vascular plant species from the northernmost subzone A (102 or 4.6% of known Arctic species) to
the southernmost subzone E (2,180 or 98.2%; Tab. 9.1).
The increase in species numbers from north to south (subzones A-E) in the Arctic is strongly correlated with the
increase of the mean July temperature (e.g. Young 1971,
Edlund & Alt 1989, Daniëls et al. 2000). Seventy-six
Arctic species, all non-endemic, occur in all five subzones;
fourteen of these species also occur in all floristic provinces (see Tab. 9.3) due to their very broad ecological and
chorological amplitude. An interesting feature of subzone
A is the absence of sedges (Carex) and woody plants (cf.
Edlund & Alt 1989, Walker et al. 2011) and the high biodiversity on the sample plot scale of 25 m2. In such plots up
to 100 species can be found: vascular plants, bryophytes
and lichens (cf. Vonlanthen 2008).
It is noteworthy that the number of Arctic endemic
species also increases from north to south, i.e. from subzone A to subzone E, with 24 species present in subzone
A, 34 in B, 66 in C, 76 in D and 71 in subzone E (Tab.
9.2, Appendix 9.2). However, the percentage of all species per subzone that are endemic is highest in subzone A
(23.5%) and decreases from subzone B through E (15.4,
13, 7.8 and 3.3%, respectively). There is only one endemic species, Saxifraga nathorstii (Nathorst’s saxifrage),
restricted to a single subzone (subzone C) within the
high Arctic, whereas 26 species are restricted to a single
subzone in the low Arctic. Four species are confined to
the low Arctic subzone D (Oxytropis beringensis, O. kateninii, O. sverdrupii and Puccinellia banksiana, Banks Island
alkali-grass) and 22 species to subzone E (Appendix 9.2).
A group of eight high Arctic endemics with exclusive or
main distribution in one, two or three of the high Arctic
subzones (A, B and C) include Puccinellia svalbardensis
(Svalbard saltmarsh grass), P. gorodkovii, xPucciphippsia vacillans (sterile hybrid between Phippsia algida and
Puccinellia vahliana), Saxifraga nathorstii, S. svalbardensis,
Braya humilis ssp. ellesmerensis (Ellesmere Island braya), B.
glabella ssp. prostrata (prostrate braya) and Draba arctica
ssp. ostenfeldii (Ostenfeld’s braya). Thirty-nine endemic
species are confined to the low Arctic subzones (D
and E), and this group includes several species of the
genus Poa and of the families Ranunculaceae, Fabaceae
and Asteraceae. Fifty-nine Arctic endemic species
occur both in the high Arctic and low Arctic. Common endemic species with wide distribution across the
subzones include the grasses Puccinellia angustata (narrow
alkali-grass) and P. vahliana (Vahl’s alkaligrass) and the
forbs Potentilla hyparctica ssp. hyparctica (Arctic cinquefoil), Draba paucilora (few-flowered draba), D. micropetala
As expected, the species considered stabilized introductions somewhere in the Arctic are mainly confined
to the two southernmost subzones of the Arctic, with
the majority confined to the warmest subzone (E). No
introduced non-native species, either stabilized or casual,
have been reported from high Arctic subzones A and B,
whereas only one non-native species, Barbarea vulgaris
(winter cress), has been reported as a stabilized introduction in subzone C (Appendix 9.4). In addition, two native
Arctic species (Sisyrinchium montanum, blue-eyed grass,
Rumex acetosa ssp. acetosa, common sorrel) have been
reported as stabilized introductions in subzone C. Hence,
the number of stabilized introduced species decreases
considerably from south to north with 101 in subzone E,
18 in D, three in C, and – as mentioned – none recorded
from the high Arctic subzones A and B. In contrast,
casual introduced species are more numerous in the high
Arctic, and temporarily occur primarily in settlements
and towns. Of the species that are native or stabilized in
the Arctic, 27 have been reported as casual introductions
from subzone C. Diversity of species that are only known
as casual introductions in the Arctic is expected to be
higher than that of stabilized introductions for all subzones, but distributional data have not yet been compiled.
9.2.3.7. Species richness in loristic provinces
A comparison of species numbers per floristic region
showed a range from 199 species for the rather heavily glaciated and northern floristic province Ellesmere
Land-N Greenland to 825 species for Beringian W
Alaska (Tab. 9.1 and 9.2). The mean number for a floristic region is 543 species or 24.5% of the total number of
species occurring in the Arctic.
Comparatively species rich are the provinces W Alaska
(825; 37.2% of the total number of species occurring in
the Arctic), Hudson-Labrador (769; 34.7%), N AlaskaYukon (732, 33%), E Chukotka (668, 30.1%), TaimyrSevernaya Zemlya (645; 29.1%) and Central Canada
(640; 28.9%) (Tab. 9.1 and 9.2, Fig. 9.2). Provinces
with a comparatively low species number, far below
the mean value of 543 (24%) per floristic province,
include E Greenland (391; 17.6%), the small Wrangel
Island province (315; 14.2%), and the two high Arctic
provinces Svalbard-Franz Joseph Land (215; 9.7%) and
Ellesmere Land-N Greenland (199; 9%).
The mean species number for the three European
Russian-W Siberian flora provinces is 593, for the four
E Siberian flora provinces 516 and for the six Beringian
323
Chapter 9 • Plants
Number of Arctic species conined to
one loristic province/subzone
Percentage of Arctic endemic species
in total Arctic endemic lora (106)
Percentage Arctic endemics in lora
of loristic province/subzone
Number of Arctic
endemic species
Percentage of number of species/
total species number (2,218)
Floristic province
Total number of vascular
plant species
Mean number of vascular
plant species
Table 9.2. Species numbers in loristic provinces and subzones. Islands indicated by •.
1 Subzone constitutes < 20% of loristic province, 2 20-50% and 3 > 50%.
Subzone
A
B
C
1
2
2
2
2
D
E
2
3
2
2
2
2
European Russian-W Siberian
Kanin-Pechora
KP
Polar Ural-Novaya Zemlya
UN
Yamal-Gydan
YG
593
642
28.9
7
1.1
6.6
0
625
28.2
16
2.6
15.1
2
513
23.1
7
1.4
6.6
0
645
29.1
16
2.5
15.1
4
429
19.3
9
2.1
8.5
0
E Siberian
Taimyr-Severnaya Zemlya
TM
Anabar-Olenyok
AO
516
2
2
2
2
2
2
3
2
1
3
2
2
1
3
2
Kharaulakh
Kh
558
25.2
14
2.5
13.2
2
Yana-Kolyma
YK
432
19.5
11
2.5
10.4
0
CW
636
28.7
22
3.5
20.8
5
2
315
14.2
35
11.1
10
3
549
24.8
4
0.7
2
Beringian
W Chukotka
• Wrangel Island
WI
S Chukotka
CS
621
33
3.8
0
1
2
3
E Chukotka
CE
668
30.1
24
3.6
22.6
6
3
3
W Alaska
AW
825
37.2
13
1.6
12.3
4
2
3
N Alaska-Yukon Territory
AN
732
33
26
3.6
24.5
3
2
3
640
28.9
34
5.3
32.1
2
769
34.7
20
2.6
18.9
0
EP
199
9
28
14.1
26.4
2
• W Greenland
GW
554
25
29
5.2
27.4
• E Greenland
GE
391
17.6
28
7.2
• N Iceland-Jan Mayen
Ic
436
19.7
1
0.2
FN
649
29.3
1
SF
215
9.7
22
A
102
4.6
24
23.5
1
Canadian
Central Canada
CC
Hudson Bay-Labrador
HL
• Ellesmere Land-N Greenland
536
1
2
2
2
2
2
1
2
3
2
2
2
3
2
2
2
26.4
1
2
2
2
0.9
0
0.2
0.9
0
10.2
20.8
3
22.6
0
N Atlantic
N Fennoscandia
• Svalbard-Franz Joseph Land
449
1
2
3
3
2
2
2
Subzone
Arctic herb subzone
N Arctic dwarf shrub subzone
B
220
9.9
34
15.5
32.1
0
Middle Arctic dwarf shrub subzone
C
507
22.9
66
13
62.3
1
S Arctic dwarf shrub subzone
D
980
44.2
76
7.8
71.7
Arctic shrub subzone
E
2,180
98.3
71
3.3
67
N
2,097
Non-Arctic-Boreal or Boreal-alpine
flora provinces 621. The three Canadian flora provinces
have a mean of 536 species, whereas the five North Atlantic flora provinces have the lowest mean value at 449.
A comparison of mean species richness of the six Beringian floristic provinces with the 15 non-Beringian prov-
4
24
inces shows that the Beringian floristic provinces have
more species (mean 621) compared with a mean value of
524 for the non-Beringian provinces. One hundred and
four species are both widespread in Beringia (i.e. found
in at least three of the six provinces) and restricted to
the Beringian region (i.e. the six Arctic Beringian floris-
324
Arctic Biodiversity Assessment
Number of scattered species
Number of frequent species
Floristic province
Number of rare species
Trisetum spicatum ssp. spicatum
Saxifraga hirculus
Saxifraga cespitosa ssp. cespitosa
Saxifraga cernua
Ranunculus pygmaeus
Poa pratensis ssp. alpigena
Phippsia algida
Oxyria digyna
Micranthes nivalis
Juncus biglumis
Equisetum variegatum ssp. variegatum
Equisetum arvense ssp. alpestre
Cardamine pratensis ssp. angustifolia
Bistorta vivipara
Species
Table 9.3. Fourteen species and subspecies distributed in all 21 loristic provinces and ive subzones. Presence of indigenous/native species
is indicated by a frequency value r = rare, s = scattered and f = frequent. Derived from Elven (2007).
r
s
f
European Russian-W Siberian
Kanin-Pechora
KP
f
f
f
s
f
s
s
r
f
f
Polar Ural-Novaya Zemlya
UN
f
f
Yamal-Gydan
YG
f
f
f
f
s
r
f
f
f
s
Taimyr-Severnaya Zemlya
TM
s
f
s
s
f
Anabar-Olenyok
AO
s
f
Kharaulakh
Kh
f
f
s
r
s
s
Yana-Kolyma
YK
s
f
r
W Chukotka
CW
Wrangel Island
WI
f
f
f
f
f
s
f
f
f
f
f
f
f
s
f
f
s
r
f
f
f
f
f
f
f
f
s
f
s
f
s
s
f
f
f
s
f
f
f
r
f
f
f
f
f
f
f
r
f
f
f
f
f
f
f
f
f
f
f
f
s
f
1
5
8
f
f
s
f
0
0
14
2
5
7
f
f
0
3
11
s
s
s
f
1
8
5
f
f
0
4
10
s
s
f
r
3
3
8
f
f
f
s
f
f
1
1
12
f
f
f
f
0
0
14
E Siberian
Beringian
S Chukotka
CS
s
r
f
f
s
f
r
f
f
s
f
r
f
f
3
3
8
E Chukotka
CE
f
f
f
f
f
f
f
f
f
f
f
s
f
f
0
1
13
W Alaska
AW
f
f
f
s
s
s
f
s
f
f
f
s
f
f
0
5
9
N Alaska-Yukon Territory
AN
f
f
s
f
f
s
f
f
f
f
f
f
f
f
0
2
12
Central Canada
CC
f
f
s
f
f
f
f
f
s
f
f
f
f
f
0
2
12
Hudson Bay-Labrador
HL
f
f
f
f
f
f
f
f
f
f
f
f
f
f
0
0
14
Ellesmere Land-N Greenland
EP
f
s
s
f
f
f
f
f
r
s
f
f
f
f
1
3
10
W Greenland
GW
f
f
f
f
f
f
f
f
f
f
f
f
r
f
1
0
13
E Greenland
GE
f
f
f
f
f
f
f
f
f
f
f
f
s
f
0
1
13
N Iceland-Jan Mayen
Ic
f
f
f
f
f
f
f
r
f
f
f
f
s
f
1
1
12
Canadian
N Atlantic
N Fennoscandia
FN
f
f
s
s
f
f
f
r
f
f
s
f
r
s
2
4
8
Svalbard-Franz Joseph Land
SF
f
f
f
f
f
f
f
f
f
f
f
f
f
f
0
0
14
Arctic herb subzone
A
f
f
r
r
f
s
f
f
r
f
f
f
s
r
4
2
8
N Arctic dwarf shrub subzone
B
f
f
f
f
f
f
f
f
f
f
f
f
f
f
0
0
14
Middle Arctic dwarf shrub subzone
C
f
f
f
f
f
f
f
f
f
f
f
f
f
f
0
0
14
S Arctic dwarf shrub subzone
D
f
f
f
f
f
f
f
s
f
f
f
f
f
f
0
1
13
Arctic shrub subzone
E
f
f
r
f
f
f
f
r
f
f
f
f
f
f
2
0
12
N
f
s
r
f
s
f
f
r
f
s
f
f
f
f
2
3
9
Subzone
Non-Arctic-Boreal or Boreal-alpine
tic provinces plus adjacent Beringian areas to the south).
These data clearly demonstrate the prominent position
of Beringia in the Arctic flora when species richness is
considered. However, in terms of species composition,
this region also stands out with a much higher number of
species either confined to or with their main distribution
in the Beringian floristic provinces, such as Selaginella
sibirica (Siberian spike-moss), Carex podocarpa (graceful mountain sedge), Ranunculus grayi, Salix phlebophylla
(skeleton-leaved willow), Oxytropis czukotica, Potentilla
elegans (elegant cinquefoil), Phlox pumila, Douglasia ochotensis (Arctic montane dwarf primrose) and Tephroseris
Chapter 9 • Plants
frigida (Arctic groundsel). Most of the more common
strictly Beringian species are confined to the low Arctic
(subzones E and D), whereas almost 20% are known
from the southernmost subzone C of the high Arctic,
mostly from Wrangel Island.
Fourteen species are common to all 21 Arctic floristic
provinces and all five Arctic subzones: Cardamine pratense ssp. angustifolia (cuckoo flower), Micranthes nivalis
(snow saxifrage), Saxifraga cernua (nodding saxifrage),
S. cespitosa ssp. cespitosa (tufted saxifrage), S. hirculus
(yellow marsh saxifrage), Oxyria digyna (mountain
sorrel), Bistorta vivipara (alpine bistort), Ranunculus
pygmaeus (pygmy buttercup), Phippsia algida (ice-grass or
spiked snow-grass), Poa pratensis ssp. alpigena (northern
meadow-grass), Trisetum spicatum ssp. spicatum (northern
oat-grass), Juncus biglumis (two-flowered rush), Equisetum
arvense ssp. alpestre (polar horsetail) and E. variegatum
ssp. variegatum (variegated horsetail, Tab. 9.3). Several
other species have a pronounced circumpolar distribution, e.g. Hippuris vulgaris (common mare’s-tail), Stellaria
humifusa (saltmarsh starwort), Koenigia islandica (Iceland
purslane), Ranunculus hyperboreus ssp. hyperboreus (Arctic
buttercup), Carex rupestris (rock sedge), C. lachenalii (Arctic hare’s-foot sedge), Cystopteris fragilis (fragile fern) and
Luzula confusa (northern wood-rush).
The number of Arctic endemic species per floristic
province varies from only one (N Iceland-Jan Mayen and
N Fennoscandia) to 35 (Wrangel Island) (Fig. 9.2, Tab.
9.2, Appendix 9.2). One-third of the 106 Arctic endemic species (33%) occur in the small Wrangel Island
floristic province, and an almost equal number occur in
the vast floristic province of Central Canada (34 species, 32%). Arctic endemic species are also well represented in the provinces of W Greenland (29, 27%), E
Greenland (28, 26%), Ellesmere Land-N Greenland (28,
26%), N Alaska-Yukon (26, 25%) and E Chukotka (24,
23%). However, if the percentage of Arctic endemics in
relation to the total flora of all the floristic provinces is
considered, different conclusions are reached. The flora
of the province Ellesmere Land-N Greenland consists
now of 14% Arctic endemics, whereas the floras of the
small provinces Wrangel Island and Svalbard-Franz
Joseph Land consist of 11 and 10% Arctic endemic species, respectively, and Central Canada only 5% Arctic
endemic species.
The paramount position of Wrangel Island is also shown
by its 10 Arctic endemic species that are restricted to
this island: Poa hartzii ssp. wrangelica, Puccinellia wrightii
ssp. colpodioides, Potentilla wrangelii, Papaver uschakovii, P.
multiradiatum, P. chionophilum, P. nudicaulis ssp. insulare,
Oxytropis uschakovii, O.unilora and Packera hyperborealis
ssp. wrangelica (Tab. 9.2, Appendix 9.2). E Chukotka has
six endemics (Carex norvegica ssp. conicorostrata, Puccinellia beringensis, xPucciphippsia czuckzorum (hybrid between
Phippsia algida and Puccinellia probably wrightii), Oxytropis
beringensis, O. katenii and Cardamine sphenophylla), W Chukotka has five of its own (Smelowskia czukotica, Papaver anjuicum, P. hypsipetes, Oxytropis sverdrupii and Plantago canes-
325
cens ssp. jurtzevii), and four endemic species are confined
to W Alaska (Ranunculus glacialis ssp. alaskensis (glacier
buttercup), Parrya nauraq, Primula anvilensis (primrose)
and Douglasia beringensis). The provinces N Iceland-Jan
Mayen, N Fennoscandia, Kanin-Pechora, Yamal-Gydan,
Anabar-Olenyok, Yana-Kolyma, S Chukotka and Hudson
Bay-Labrador lack their own Arctic endemic species.
Borderline species are absent in the remote floristic
provinces of Svalbard-Franz Joseph Land, Wrangel
Island, Ellesmere Land-N Greenland and E Greenland.
All of these provinces represent islands isolated from the
mainland to the south, and are mainly entirely high Arctic
(Tab. 9.2). They are also unknown from E Chukotka and
W Greenland, whereas Hudson Bay-Labrador has the
highest reported number of borderline species (44 species), followed by W Alaska (25) and Central Canada (19).
Numbers of stabilized introduced species are highest
for the floristic provinces N Fennoscandia (63 species),
Kanin-Pechora (52), W Greenland (50) and N IcelandJan Mayen (52), followed by Polar Ural-Novaya Zemlya
(32), Hudson Bay-Labrador (30), Taimyr-Severnaya
Zemlya (23) and W Alaska (20) (Tab. 9.1). Some parts
of the Arctic, such as the floristic provinces Ellesmere
Land-N Greenland and Wrangel Island, are noteworthy
for the complete absence of stabilized introduced species. Such species have also not been reported for the
province Anabar-Olenyok, whereas only a few species
are reported for E Greenland (2), S Chukotka (5) and W
Chukotka (6). Several non-native species (Plantago major
(great plantain), Chenopodium album (common lamb’squarters), Thlaspi arvense (field penny-cress), Brassica rapa
ssp. campestris (turnip) and Trifolium hybridum (alsike clover)) are widespread in the Arctic, occurring in all five
floristic province groups (see further Appendix 9.4).
Stabilized introductions can often be attributed to continuous human activities when they occur in and around
settlements and towns, in particular in climatologically
favorable parts of the Euro-Siberian Arctic. In Iceland
and N Fennoscandia, stabilized introductions strongly
dominate in old cultural landscapes. In S and W Greenland, extensive small-scale agriculture using fields, hay
meadows and pastures was introduced by Norse settlers
more than 1,000 years ago, and nowadays agriculture,
sheep breeding and forestry are common practices
there (cf. Pedersen 1972, Fredskild 1988). The group
of stabilized introductions is taxonomically very heterogeneous, as expected. They mainly include common
species of synanthropic European vegetation (Appendix 9.4). Species of ruderal and arable weed vegetation
include Descurainia sophia (tansy-mustard), Thlaspi arvense
(penny-cress), Capsella bursa-pastoris (shepard’s-purse)
and Lamium purpureum (purple dead-nettle). The grasses
Elytrigia repens (couch), Anthoxanthum odoratum (sweetvernal grass), Lolium perenne (rye-grass), Poa pratensis
ssp. angustifolia (narrow-leaved meadow-grass), P. supina
(supine bluegrass), Dactylis glomerata (cock’s foot), Phleum
pratense (timothy) and Alopecurus geniculatus (marshfoxtail), and the forbs Trifolium pratense (red clover),
326
Primula elatior (oxlip) and Veronica chamaedrys (germander
apeedwell) are indicators for nutrient-rich mesic and wet
grasslands (Mucina 1997, Jarolímek & Šibík 2008).
9.2.3.8. Hotspots
Seen from large-scale perspectives such as subzones and
floristic provinces, the taxonomic diversity of vascular plants in the Arctic is low in comparison with the
vascular plant flora of non-Arctic biomes (Barthlott et al.
1996; see Fig. 1.1 in Meltofte et al., Chapter 1). The same
often applies for regional Arctic floras. However, there
are several areas of enhanced taxonomic diversity in the
Arctic due to strong abiotic factors, such as a heterogeneous climate. These areas of enhanced taxonomic diversity
are often associated with dramatic topography, such as
mountainous areas, and have been referred to as ‘Arctic hotspots’ (Elvebakk 2005) or polar oases (Crawford
2008a). They are extrazonal, locally warm areas with
biodiversity elements not found in their surroundings. In
Arctic hotspot complexes, topographic complexity leads
to high climatic diversity and correspondingly higher
biodiversity in flora and vegetation (Elvebakk 2005).
Four such hotspot complexes are found in Svalbard,
where several thermophilous, low Arctic or southern
species occur locally in high Arctic environments (Elvebakk 2005). Hotspots in high Arctic Canada include the
Fosheim Peninsula on west-central Ellesmere Island (e.g.
Hot Water Creek; Edlund et al. 1989) and Lake Hazen
(Crawford 2008a) in the northernmost part of Ellesmere
Island with a rich thermophilous vascular flora of 117
species (this is about 57% of the total flora of the Ellesmere Land-N Greenland floristic province (see Section
9.2.3.7), in the world’s northernmost extension of high
Arctic subzone C. Other hotspots in Canada include the
Minto Inlet area of NE Victoria Island, where scattered
tall willow riparian thickets with a diverse understory
occur in Carex dwarf shrub tundra (Edlund 1983).
Hotspots in Greenland are among others the continental inland of W Greenland around Søndre Strømfjord/
Kangerlussuaq (Böcher 1954, Sieg et al. 2006) (subzone
E of the low Arctic) with south-facing slopes of boreallow Arctic steppe vegetation (Saxifrago-Calamagrostietea
purpurascentis; Drees & Daniëls 2009) and the inland of S
Greenland (Feilberg 1984) with Qinguadalen (Fredskild
& Odum 1990) as a core area of the sub-Arctic forest enclave in low Arctic Greenland. The central part of Beringian Wrangel Island is certainly the most pronounced
hotspot complex of the Russian Arctic (Kholod 2007).
Due to the small-scale climatic and biotic diversity, the
Arctic hotspot complexes are strongly recommended
as Arctic field laboratories for climate change-related
research (see Elvebakk 2005 and see Section 9.2.7).
9.2.4. Traditional use of vascular plants
Local fauna and flora have always played an essential role
in the life and culture of the indigenous peoples of the
Arctic (e.g. Robbé 1994, Ainana & Zagrebin 1997, Garibaldi 1999, Jones 2010). Traditionally, many plant spe-
Arctic Biodiversity Assessment
cies were collected for consumption of flowers, berries,
stems, leaves or roots, and for other uses. Preservation
for later consumption followed traditional customs. The
use of plants as food by Inupiat and use as medicines by
indigenous peoples in Alaska are described in Garibaldi
(1999) and Jones (2010). Plant use by Chukotkan indigenous peoples is reviewed in Ainana & Zagrebin (1997).
Traditional uses of plants by Arctic peoples in Canada
are summarized in Aiken et al. (2007). Recent publications synthesize the knowledge of elders on the traditional use of plants by Gwich’in in sub-Arctic western
Canada (Andre & Fehr 2000), by Inuit on Baffin Island
(Ootoova et al. 2001), and by Inuvialuit in the western
Canadian Arctic (Inuvialuit elders with Bandringa 2010).
Traditional use of plants in low Arctic SE Greenland is
described by Robbé (1994), who studied the life and culture of Inuit hunters and their families in the small settlement Tiniteqilaq, Ammassalik district, in the 1960s.
The uses of plants are varied in Greenland, as they are
elsewhere in the Arctic, and include the use as vegetables
in particular Angelica archangelica (garden angelica; stem),
Rhodiola rosea (stone crop; flower, leaf, root), Oxyria digyna (mountain sorrel; leaf, root), Taraxacum croceum (leaf)
and Bistorta vivipara (alpine bistort; root). The berries of
other species such as the dwarf shrubs Empetrum nigrum
ssp. hermaphroditum (black crowberry) and Vaccinium
uliginosum ssp. microphyllum (polar bilberry) were eaten
fresh or preserved for consumption. Salix glauca ssp. callicarpaea (grayleaf willow), Betula nana (dwarf birch) and
Juniperus communis ssp. nana (common mountain juniper)
were used as firewood or to make tools, while Thymus
praecox (wild thyme) was used as a substitute for tobacco.
Grasses and sedges, in particular Poa alpina (alpine
meadow-grass), Carex spp. and some others, were used as
insulation between the two sole layers in double-skinned
seal skin boots (kamiks). Many useful plants have Greenlandic names (e.g. Osterman 1938, Böcher et al. 1966,
Robbé 1994, Foersom et al. 1982).
9.2.5. Rare and threatened Arctic endemic
species
9.2.5.1. Rare Arctic endemic species
Among the 106 Arctic endemic species, 69 species show
a very restricted distribution within the Arctic territory,
occurring in only one or two of the 21 floristic provinces.
Forty-seven species are only known from one province
(Appendix 9.2), and 28 of these are known only from one
of the six Beringian provinces. The small island province
Wrangel Island harbors 10 of its own endemics, E Chukotka has six and W Chukotka has five such endemics,
whereas S Chukotka has none. Of the Beringian North
American provinces, W Alaska has four own endemics
as compared with only three in N Alaska (Poa hartzii ssp.
alaskana, Papaver “murrayii” and Potentilla aff. pensylvanica).
Among the Canadian provinces, Central Canada has two
of its own endemic species (Papaver sp. “Banks” and Braya
thorild-wulii ssp. glabrata, smooth Greenland braya). Elles-
327
Chapter 9 • Plants
mere Land-N Greenland has two local endemics (Braya
humilis ssp. ellesmerensis, Ellesmere braya and B. glabella
ssp. prostrata, prostrate braya), whereas Hudson Bay-Labrador lacks its own endemic species (see also Tab. 9.2).
Among the North Atlantic floristic provinces, W Greenland has three of its own rare endemics (Sisyrinchium
groenlandicum, Puccinellia porsildii and P. groenlandica),
whereas E Greenland has only one single endemic of its
own: Saxifraga nathorstii. N Iceland-Jan Mayen and N
Fennoscandia are lacking their own Arctic endemics altogether, whereas Svalbard-Franz Joseph Land is known
to harbor (at least) three local endemics: Puccinellia
svalbardensis (Svalbard alkali-grass), Saxifraga svalbardensis
(Svalbard saxifrage) and Potentilla insularis. Of the three
European Russian-W Siberian floristic provinces, only
Polar Ural-Novaya Zemlya has two endemic species (Astralagus gorodkovii and A. igoshinae), while Kanin-Pechora
and the Yamal-Gydan province are lacking their own
endemic species. Similarly, two of the four E Siberian
flora provinces (Anabar-Olenyok and Yana-Kolyma)
lack their own endemics, whereas the Taimyr-Severnaya
Zemlya province has four endemics (Puccinellia gorodkovii,
Oxytropis tichomirovii, O. middendorfii ssp. schmidtii and
Draba taimyrensis), and two endemics are restricted to the
Kharaulakh province (Oxytropis sordida ssp. arctolensensis
and Papaver leucotrichum).
Twenty rare (r) Arctic endemic species are restricted to
only one floristic province and only one Arctic subzone
(Tab. 9.4, Appendix 9.2) and almost all occur in the
low Arctic (subzones D and E). An exception is Saxifraga
nathorstii, which is found in subzone C of the high Arctic
in the E Greenland floristic province.
9.2.5.2. Threatened Arctic species
Currently, we have no evidence that any Arctic plant
species has become extinct in the last 250 years (Elven
2011). Nevertheless, all species with very low abundance and a restricted distribution might be considered
potentially threatened in the context of future climate
change. Such species may meet the guidelines of IUCN
(2008) for threatened status when climate warming is
considered a threat, however exact information on distribution, population number and size is often lacking.
Other IUCN assessment criteria, such as information
on population trends, are almost completely lacking for
Arctic vascular plants.
In the Atlas of Rare Endemic Vascular Plants of the
Arctic (Talbot et al. 1999), 69 taxa were identified as
rare, although a different species concept from that in
PAF was used. Moreover, 12 ‘micro-species’ (agamospecies) of Taraxacum were included by Talbot et al. (1999)
as well as several species of sub-Arctic or boreal territories such as the treeless Aleutian Islands. Considering
these differences, we present in Tab. 9.4 a reduced list
of 20 potentially threatened rare Arctic endemic species
that occur in only one floristic province and one Arctic
subzone based on our own evaluations from the PAF.
Table 9.4. The 20 Arctic endemic species and subspecies known
only from one Arctic loristic province and one subzone as rare (r),
and as such potentially threatened. Species ordered by family; their
status in Talbot et al. (1999) is also indicated where included: VU =
vulnerable; DD = data deicient; LR = lower risk; nt = near threatened.
Species
Brassicaceae
Status
Parrya nauraq
Smelowskia czukotica
VU
Cyperaceae
Carex norvegica ssp. coniorostrata
Fabaceae
Astralagus gorodkovii
Astralagus igoshinae
Oxytropis beringensis
Oxytropis katenii
Oxytropis middendorii ssp. schmidtii
Oxytropis sordida ssp. arctolenensis
Oxytropis sverdrupii
VU
DD
LR (nt)
DD
Iridaceae
Sisyrinchium groenlandicum
LR (nt)
Papaveraceae
Papaver anjuicum
Papaver hypsipetes
Papaver leucotrichum
LR (nt)
Plantaginaceae
Plantago canescens var. jurtzevii
Poaceae
Poa hartzii ssp. alaskana
Puccinellia beringensis
LR (nt)
LR (nt)
LR (nt)
Primulaceae
Douglasia beringensis
VU
Ranunculaceae
Ranunculus glacialis ssp. alaskensis
VU
Saxifragaceae
Saxifraga nathorstii
While threats to the endemic Arctic species are generally poorly known, they might increase because the Arctic
is at the forefront of experiencing the effects of climate
change, and other more direct human impacts are also
increasing (ACIA 2005).
A new project on Red-listing of Arctic vascular plant
species was recently initiated and should be finalized
during 2014 (CAFF Flora Group). Only the species level
is considered in the project, because of uncertainties
about the status of many subspecies. For the same reason,
Taraxacum and Hieracium micro-species are excluded from
consideration, as they are poorly known in many cases and
have extremely narrow distributions. The Red List for
Arctic vascular plants will include mainly rare endemic
species because of lack of good monitoring data. So far, a
candidate list of 164 species is under discussion. For those
species, all known data will be collected and evaluated
following IUCN criteria (IUCN 2008). If considered
threatened, they will be assigned an IUCN status of Critically Endangered, Endangered or Vulnerable. It is likely
that only very few Arctic vascular plants will be considered threatened according to these criteria. It is critical to
note that in the near absence of any population trend data
on Arctic plants, critical evaluation of these taxa is very
difficult. Trends on declining population size or number
of individuals have so far not been detected and, if they do
exist, would likely be too low to meet the IUCN criteria.
The current status and trends of Arctic plants is based on
328
much fragmentary information, and many assumptions
have to be made in order to make informed decisions on
the status of Red-listed species. For now, the best available
information is the number of sites recorded in herbaria
based on specimen collections or field observations. One
needs to consider that the absence of a taxon in a distributional map is not necessarily a reflection of the situation in
nature, but might represent a knowledge gap. For example, in the recent Red List reassessment of vascular plant
species in Svalbard (which has a relatively well-known
flora), the status of 28 species changed, but the majority of
these changes was due to increased knowledge as a result
of recent fieldwork (K.B. Westergaard, pers. com.).
Genetic diversity, in addition to species diversity, is also
an important consideration when assessing the threatened status of a species. For example, the loss of genetic
diversity is greatest if small genera are lost; given that
mean species diversity within a genus is low in the Arctic, this may be an important consideration. While rare
Arctic endemics are of obvious conservation concern,
Arctic populations of species with an Arctic-alpine distribution should also be considered. Such species often
have a very fragmented distribution with isolated populations likely not cross-breeding; thus, giving greater
weight to these isolated genetically distinct Arctic
populations may be appropriate (Väre et al. 2003). Also,
climate change induced loss of habitat may cause considerable loss of genetic diversity within species, which may
reduce the species ability to adapt to a changing climate
(Alsos et al. 2012a)
9.2.6. Trends and monitoring eforts
Elders of the Kolymskaya village reported in 2006 that
willows are moving to tundra and to river banks. They
said:
»
It tells of the changes which are under way. You should
graze cows and horses, not reindeer on these spots. All of the
tundra is covered with willows and bushes. It grows very fast now.
We do not know how we can herd reindeer in the middle of these
changes.
(Mustonen 2007).
Most vascular plants in the Arctic are long-lived to
very long-lived, and many of them may have the genetic potential to spread into novel niches or persist in a
changing climate. There are few threats that will affect
the entire Arctic today or in the near future, but climate
change certainly will (ACIA 2005). It is hard to predict
what direct effects climate change will have on Arctic
plant species. Many of them have already experienced
pronounced climatic changes in post-glacial times. Climate change could affect the vegetation in several ways,
and this too is hard to predict (Euskirchen et al. 2009).
Northern plants are expected to loose 36-43% of their
current distribution under the A2 climate change scenario and 26-43% under the B2 scenario (Alsos et al. 2012).
It is reasonable to predict that in many cases borderline
Arctic Biodiversity Assessment
species and many others with southern distributions will
increasingly move into the Arctic. Daniëls & de Molenaar (2011) observed a tendency of sub-arctification of
the vascular plant flora near Ammassalik, SE Greenland, during the last 100 years. Other studies across the
Arctic have shown decadal and multidecadal changes in
species composition of plant communities (e.g. Callaghan et al. 2011a, 2011b, Daniëls et al. 2011, Henry et al.
2012, Kapfer et al. 2012, Schmidt et al. 2012). In general,
these changes are minor in dry habitats, however more
pronounced in moist and wet sites, such as snow beds,
mires, fens and shallow ponds. This is likely explained
by substrate drying due to earlier snow melt along with
strong warming in summer. Shrub expansion in the Arctic is reported in several publications (see summary by
Klein et al. 2008 and Ims & Ehrich, Chapter 12). A recent study by Henry et al. (2012) showed that increased
shrub cover all over the Canadian Arctic was supported
by results from experimental warming. The fate of
cold climate plants in a warmer world has been amply
addressed and summarized by for example Callaghan
(2005) and Crawford (2008a, 2008b). Possible changes
in the composition of the vascular plant flora are difficult
to predict. However the heterogeneity of Arctic habitats
together with genotypical and phenotypical variability of
Arctic plants will certainly result in the evolution of adaptations that may benefit from higher temperatures and
longer growing seasons (Crawford 2008b). Callaghan
(2005) suggests that Arctic biodiversity is likely resistant
to variations in climate, but perhaps not to competition
that will come from southern species expanding their
ranges to the north. Crawford (2008b, p. 224) expects
that “botanists in the future may look forward to relaxed
exploration of a diverse and plentiful flora as far as north
as land exists.”
While the CAFF Circumpolar Biodiversity Monitoring
Program (CBMP) is promoting active engagement of
local communities in monitoring programs (Gofman
2010) and has initiated a Terrestrial Expert Group, the
formation of formal networks dealing with plants in the
Arctic as a whole has been rather slow. There are a few
regional initiatives that are working towards providing
baseline information on plant biodiversity in the Arctic.
The most advanced of these initiatives is a detailed monitoring plan in Svalbard to document population trends in
selected rare plant species, a monitoring effort initiated
by the Norwegian Polar Institute.
Monitoring programs have been established in two areas
of Svalbard. A program in Colesdalen is focusing specifically on population trends in five rare vascular plant species (Arnesen et al. 2012), while the program in Endalen
is focusing on monitoring vegetation and possible effects
of climate change and pollution on floristic composition (Aarrestad et al. 2010). In 2008-09, monitoring of
population trends in five vascular plants, of which four
are regionally Red-listed, was initiated in Colesdalen:
the annual herb Euphrasia wettsteinii (EN), the perennial herb Campanula rotundifolia ssp. gieseckiana (common
harebell) (EN), and the dwarf shrubs Vaccinium uliginosum
329
Chapter 9 • Plants
spp. microphyllum (CR), Empetrum nigrum ssp. hermaphroditum and Betula nana ssp. tundrarum (EN) (Alsos 2011).
Occurring on warm south-facing slopes with favourable
climate, these species are at their climatic limit in Svalbard and are expected to be sensitive to climate change.
Action plans are currently being prepared for the four
Red-listed species (Alsos & Arnesen 2009). With their
local distributions very well documented, monitoring is
focusing on population dynamics and development, and
includes monitoring of local climate and physical conditions at the habitats.
In Alaska, the U.S. National Park Service’s Arctic Parks
Network has initiated inventory and monitoring in
remote regions to describe current biodiversity, and to
give land managers and other agencies the information
needed to make more informed decisions about protecting potentially rare and endangered species (Parker
2006, Racine et al. 2006). Similarly, the U.S. Bureau of
Land Management is actively monitoring rare plants in
Alaska (Carroll et al. 2003, Cortés-Burns et al. 2009).
The University of Alaska Fairbanks, which administers
the Toolik Lake Field Station, has several Long-Term
Ecological Research sites in the northern foothills of
the Brooks Range. Cortés-Burns et al. (2009) reviewed
information on 31 rare vascular plants of Alaska’s North
Slope Region, including population number and location, but no information on population size is given. In
response to the increasing threat of the establishment
and spread of non-native plant species (see Lassuy &
Lewis, Chapter 16), the U.S. Forest Service has developed a ranking system to help identify problematic nonnative plants and to prioritize control efforts in Alaska
(Carlson et al. 2008).
While there are numerous programs focused on monitoring vegetation change as a result of climate change in
Arctic Canada and a few in Greenland, there are almost
none focused specifically on monitoring populations of
rare or threatened species. One example is the extensive monitoring at Zackenberg, NE Greenland, which
has taken place over the last 18 years (e.g. Bay 2006,
Meltofte et al. 2008). The Government of the Northwest
Territories in Canada initiated the NWT General Status
Ranking Program in 1999 to collect information on wild
species as a tool for setting conservation priorities. Information is assembled in an on-line database, the NWT
Species Monitoring Infobase, and reports on the general
status ranks are published every five years. The ranks of
all vascular plant species were first published in 2006
using the following categories: ‘May Be at Risk’, ‘Sensitive’, ‘Secure’, ‘Alien’ or ‘Undetermined’ (Working
Group on General Status of NWT Species 2006). The
2011 report documented 89 changes in ranks of vascular
plant species, all due to new information, taxonomic
change or a more rigorous assessment of threats, rather
than a real change in status (Working Group on General
Status of NWT Species 2011).
The necessity of establishing an International Arctic Vegetation Database for circumpolar biodiversity studies was
recently articulated by Walker & Raynolds (2011) and
Walker et al. (2013). Plants, mainly vascular plants and
bryophytes, and lichens are the main structural components of the plant communities in the terrestrial Arctic
landscapes. The floristic composition of these communities reflects the present day local habitat conditions
and the geographical position of the plant communities.
Thus, the distribution of plant communities shed light
on environmental conditions, and they can be regarded
as early warning systems for environmental change.
However, circumpolar knowledge of plant community
types and their classification is still rather poorly developed (cf. Walker et al. 1994, Daniëls et al. 2005), and a
circumpolar floristic classification system is still lacking.
There is thus a strong need to bring together all existing
plot-based vegetation analyses in an International Arctic
Vegetation Database along with intensifying the exploration of vegetation in poorly studied and unstudied areas.
In particular, floristical vegetation plot analyses are very
scarce from the climatologically most extreme, ecologically unique and likely most sensitive and vulnerable part
of the Arctic, its northernmost Subzone A (cf. Walker
et al. 2012). Hence, storage of plot-based vegetation
analyses in an International Arctic Vegetation Database
is fundamental for circumpolar biodiversity studies,
monitoring and predictive modeling efforts (Walker &
Raynolds 2011, Walker et al. 2013).
9.2.7. Conclusions and recommendations
There is a great need for intensifying biodiversity research on Arctic flora with emphasis on molecular phylogenetic taxonomy, vegetation classification, monitoring
and modeling. Coordination and cooperation between
researchers must be improved. Baseline information on
the distribution of Arctic plant species, including population number and size, is essential for accurately determining species status. Given the almost complete lack of
population trend data for Arctic plant species, monitoring programs should be established in order to gather
trend data. The conservation status of Arctic plant
species can only be objectively assessed once information
becomes available on the population status and trends of
individual species and their plant community types. Due
to their small-scale climatic and biotic diversity, Arctic
hotspot complexes are strongly recommended as Arctic
field laboratories for climate change-related research
(see Elvebakk 2005) and for consideration as protected
areas. In particular, monitoring of species ranges along
altitudinal gradients in Arctic mountains is strongly
recommended. Here we might expect above all species
response to climate warming due to the relatively steep
climate gradient (e.g. Elvebakk 2005, Schwarzenbach
2006, Pauli et al. 2007, Jedrzejek et al. 2012, 2013).
9.3. BRYOPHYTES
Bryophytes comprise three monophyletic groups: mosses
(Bryophyta; Bryopsida), liverworts (Hepatophyta;
Hepaticopsida) and hornworts (Anthocerotophyta;
330
Antherocerotopsida) (Raven et al. 2005). The liverworts
are generally considered the oldest group of aquaticterrestrial plants, derived from algal (charophycean)
ancestors in the Ordovician (Graham & Gray 2001).
Earth’s oldest fossil record of bryophytes is the liverwort
Metzgeriothallus sharonae, dating from the Middle Devonian (Givetian) in eastern New York, USA (Hernick et al.
2008). Our current understanding of the phylogeny of
bryophytes and other land plants was reviewed by Groth
& Knoop (2005). Their phylogenetic analysis of molecular sequences from the mitochondrial genome provided
evidence for the status of liverworts as the basal group of
extant land plants and hornworts as sister to the tracheophytes (Gradstein & Heinrichs 2005).
Bryophytes strongly differ from vascular plants in life
cycle, structure and physiology. Both have a diplohaplont
life cycle characterized by the fact that meiosis (reduction of chromosome pairs to one copy) occurs before
syngamy. However in bryophytes meiosis does not occur
immediately before syngamy as in vascular plants. The
haploid (n) gametophyte is the dominant stage (or phase)
in all bryophyte groups, whereas in vascular plants it is
the diploid (2n) sporophytic generation. The gametophyte
is formed after germination of uniform haploid spores
(homoiospory) produced by meiosis in the sporophyte,
which is attached to the gametophyte. Germinated spores
each produce a protonema that develops into a mature
individual. These can be thallose or with leafy stems with
different growth forms. In male sex organs many spermatozoids are produced, whereas only one egg cell is formed
in female sex organs. Bryophytes can be monoecious,
with gametophores possessing both male and female sex
organs on the same plant; or dioecious, with gametophores that bear male and female sex organs on separate
plants. Transport of spermatozoids to the egg cells is
only possible in water. After fertilization, the sporophyte
develops and remains connected with the gametophyte.
Asexual propagation is common and contributes to
short-distance dispersal, whereas spores, which can be
long-lived, also contribute to long-distance dispersal.
Chromosome numbers in liverworts and hornworts are
generally low and stable (9 and 5, respectively) but polyploidy including allopolyploidy is present especially in the
Arctic (Schuster & Damsholt 1974, Vilnet et al. 2010). In
mosses, higher chromosome numbers occur, and autopolyploidy (polyploidy arising from chromosomes of the
same source through spontaneous doubling of chromosome number) is not uncommon (Schofield 1985, During
1992, Damsholt 2002, Raven et al. 2005, Glime 2007).
Bryophytes have a rather simple morphology, anatomy
and physiology. The gametophyte is either thallose (in
liverworts and hornworts) or with leafy stem (liverworts
and mosses) and may be attached to the substrate by
rhizoids. True roots, stems and leaves are not developed.
Lignin synthesis for support of vertical growth is absent.
Most bryophytes are not able to regulate uptake and
release of water, gases and minerals. Contrary to almost
all vascular plants, bryophytes are poikilohydric, thus
physiologically inactive when dry (Schofield 1985). Low
Arctic Biodiversity Assessment
stature and a poikilohydric nature make them mainly
dependent upon conditions of the uppermost soil,
substrate surface and adjoining atmosphere. As a result,
bryophytes are weaker competitors than vascular plants.
However they seem well adapted to their “limited mode
of life, but also liberated, being able to grow where vascular plants cannot” (Proctor 2000a, 2000b).
Bryophyte biodiversity on a worldwide scale in terms of
species number is rather low compared with that of the
vascular plants. Nowadays c. 16,000 bryophyte species are known worldwide; c. 6,000 liverworts, c. 100
hornworts and c. 9,900 mosses (Raven et al. 2005). This
is only c. 6% of the c. 260,000 vascular plant species
(Raven et al. 2005).
As a group, bryophytes have a cosmopolitan distribution
(Herzog 1926, Shaw et al. 2005). They become dominant
where vascular plants meet less optimal growth conditions. They can cope with harsh and special environmental conditions and show various growth forms (Birse
& Gimingham 1957) and life strategies (During 1979,
1992) (see also Schofield 1972 and Longton 1982, 1988).
9.3.1. Bryophytes in the Arctic
Bryophyte vegetation in the Arctic is mainly dominated
by turf and mat growth forms (Schofield 1972). Short
turfs are loose or more frequently compact colonies,
< 1-2 cm tall, formed by sparingly branched acrocarpous
mosses with main shoots parallel and erect. The branches
are erect and of indeterminate growth, such as in e.g.
the moss genera Pottia and Bryum and some liverworts
e.g. Tetralophozia setiformis (monster pawwort), Lophozia
personii (chalk notchwort) and species of the genera Gymnomitrion (frostwort) and Scapania (earwort). Tall turfs
are > 2 cm tall, and are divided into turfs of acrocarpous mosses with erect branches such as in e.g. Dicranum
(forkmoss) and Polytrichum (haircap moss) and turfs of
pleurocarpous mosses with divergent branches forming
more frequently loose colonies such as in e.g. Orthothecium chryseon (golden autumn moss), Drepanocladus (hookmoss) and Sphagnum (bogmoss). Mats are formed by
leafy liverworts, e.g. the genus Cephalozia (pincerwort),
Ptilidium ciliare (ciliated fringewort), Marsupella arctica
(Arctic rustwort), Cephaloziella (threadwort) and Lophozia
(notchwort), or mosses (e.g. Racomitrium, fring-moss)
with determinate branching of prostrate or ascending interweaving shoots in compact colonies. In turfs,
mats and carpets many species of bryophytes are usually
intermingled, so in small patches of c. 3-5 m2 up to 1520 species can be found. Open turfs and thread growth
form (single shoots on lichens and on bare soil) are
characteristic for many bryophytes in the high Arctic,
especially on exposed sites. In cracks in patches of bare
soil in spotted tundras, on bare soil between boulders,
in cracks between polygons, and on wet bare soil on
solifluction slopes, single thalli of liverworts such as e.g.
Athalamia, Sauteria as well as Scapania gymnostomophyla
(narrowlobed earwort), S. cuspiduligera (untidy earwort),
Leiocolea heterocolpos var. harpantoides (ragged notchwort)
Chapter 9 • Plants
and many more occur. Their small size (often less than 1
mm broad and several mm long) allow many species to
persist in microhabitats.
There is little known about life strategy types of bryophytes in Arctic. This topic is discussed in only a few
publications (Mogensen 1987, 2001). The life strategy
types perennial stayer, colonist and fugitive are most
conspicuous in the Arctic as was shown for Greenland by
Mogensen (1987, 2001), however shuttle species occur
as well. Perennials stayers occur in relatively constant or
regularly, moderately fluctuating environments. They
have a long, variable life span, a low sexual and asexual
reproduction effort and small spores (< 20 µm), and can
be divided into competitive and stress-tolerant perennial stayers (During 1992). Their growth form is mainly
tall turf (e.g. Dicranum) and rough mats (Calliergon, spear
moss, Drepanocladus and Sphagnum). Colonists have a
short lifespan (< 5 years), a high asexual and sexual
reproductive effort and very persistent, small spores
(< 20 µm). Their growth form is predominantly short
turf (e.g. Pohlia, nodding moss, Andreaea, rockmoss and
Amphidium, yokemoss) and smooth mat (e.g. Cephaloziella, Anthelia, silverwort and Cephalozia). Fugitives are
331
ephemeral or annual with a high sexual reproductive
effort, frequent sporophyte development and with small
spores (< 20 µm). The spores are very persistent and
can be long-lived. The gametophyte has an open turf
growth form. They are widespread and mainly occur in
small-scale unstable and disturbed habitats (e.g. Funaria,
cord-moss, Desmatodon, screwmoss, Pottia, tuftmoss and
Stegonia, screwmoss). Species of the dung moss family
Splachnaceae (e.g. Splachnum vasculosum, rugged collarmoss and Tetraplodon pallidus, cruet-moss) have a shuttle
strategy. They grow on temporary organic substrates
such as the dung of musk oxen, reindeer, Arctic hares
and lemmings. They have a short turf growth form, a
pauci-pluriennial life span and their sexual reproductive
effort is high. They frequently produce sporophytes and
clumps of spores are distributed by insects.
In small ponds and lakes, mires, bogs, spring areas,
along melt water creeks and snow beds (Fig. 9.5), and
amid dwarf shrub heaths and rocks, mosses and liverworts are locally abundant and dominant, constituting
plant communities of their own (cf. Holmen 1955, Brassard 1971a, 1971b, Steere 1976, Longton 1982, 1988,
Frisvoll & Elvebakk 1996, Dierßen 2001, Dierssen &
Figure 9.5. Moss-rich snow bed near Cape Isachsen, Ellef Ringnes Island, Canada (Arctic subzone A). The vegetation is dominated by the
mosses Bryum cryophilum (red), Aulacomnium turgidum (mountain groove moss) and Orthothecium chryseon. Photo: Fred J.A. Daniёls, July 2005.
332
Dierssen 2005). Beyond that, bryophytes often contribute strongly to the species richness of many other tundra
vegetation types (cf. Hadač 1989, Möller 2000, Sieg et
al. 2006, Kholod 2007, Walker et al. 2011, Jedrzejek
et al. 2012). Very few vegetation types in the Arctic
occur without bryophytes. Single shoots occur almost
everywhere, in particular in the high Arctic. On a fine
scale (up to a few square kilometers), species diversity of
bryophytes (and lichens) is higher than that of vascular
plants. The bryophyte flora of Svalbard counts c. 388
species whereas less than 200 species of vascular plants
are recorded for this archipelago. Nevertheless, the
overall species number of bryophytes in the Arctic is distinctly lower than that of vascular plants (c. 2,218) (this
chapter), while the small group of hornworts (worldwide
c. 100 species) is absent. In general species number of
bryophytes decreases from the taiga zone to the Arctic
but the contribution of bryophytes in plant diversity increases. Particularly in the Murmansk Province situated
in the sub-Arctic the ratio of bryophyte species to vascular plants species (including adventive species) is c. 1:2,
whereas in Svalbard this ratio is c. 2:1 (N.A. Konstantinova, unpubl.). Diversity of bryophytes in the Arctic
(c. 900 species) is 2.5 times lower than the diversity of
vascular plants, whereas the world species number of
bryophytes (16,000 species) is c. 16 times lower than
that of vascular plants (c. 260,000 species).
The use of bryophytes by indigenous peoples of the Arctic is little known and probably very minor. In SE Greenland, three bryophyte species were reported to be commonly used, although not for consumption: Polytrichum
species as lighter of oil lamps, Sanionia uncinata (sickleleaved hookmoss) as wick in oil lamps and Racomitrium
lanuginosum (wooly fringemoss) as sponge and cleaning
tissue (Robbé 1994). Some bryophytes have Greenlandic
names: Dicranum fuscescens (dusky forkmoss, issuatsiaat
illaagutaasat), Marchantia alpestris (mountain livergreen,
issuatsiaat sialussiutillit), Hylocomium splendens (glittering
woodmoss, issuatsiaaat qaleriiaattut), Racomitrium lanuginosum (issuatsiaat qasertut), Sanionia uncinata (issuatsiaat
kukiusallit) and Sphagnum girgensohnii (Girgensohn’s bogmoss, issuatsiaat iparaq) (Foersom et al. 1982).
9.3.2. Arctic bryoloristic studies
Steere (1954, 1971) reviewed the main results of Arctic
bryology research up to the 1950s and 1970s, respectively. The latter publication is an important reference
work comprising more than 150 titles of taxonomic and
floristic studies. One year later, Schofield (1972) published a thorough review of the main results of bryological research in Arctic and boreal North America and
Greenland.
The monographs of the moss floras of the Queen Elizabeth Islands in high Arctic Canada (Brassard 1971a,
1971b) and of Arctic Alaska (Steere 1978), the liverwort
floras of Arctic Alaska (Steere & Inoue 1978), of W
Greenland (Schuster & Damsholt 1974) and S Greenland
(Schuster 1988), the three volumes of the Illustrated
Arctic Biodiversity Assessment
Moss Flora of Arctic North America (Long 1985, Crum
1986, Murray 1987), and the moss checklist of Canada
(Ireland et al. 1987) are other key contributions to the
Arctic bryoflora.
Longton (1988) mentioned species numbers for several
Arctic regions stating that assessment of the size,
geographical affinity and history of the Arctic bryofloras is still hampered by inadequate distribution data
and taxonomic uncertainty. Particularly in boreal and
Arctic mosses and liverworts, taxonomic problems
are considerable due to their reactions to unfavorable
or extreme environmental conditions (e.g. Schuster &
Damsholt 1974, Steere 1979, Schuster 1988). Steere
(1979) stated “The genus Bryum in the high Arctic will
remain an almost impenetrable mystery.” The taxonomic
problems are of biological/physiological and bryogeographical nature. In polar deserts with high alkalinity
and little precipitation many species occur as dwarf
forms. Liverworts of the genera Lophozia and Scapania
show considerable plasticity, which may explain the
lack of consistency in taxonomic approach particularly
in the families Lophoziaceae, Scapaniaceae and Jungermanniaceae (e.g. Schuster & Damsholt 1974, Damsholt
2002, Konstantinova & Vilnet 2009, Konstantinova et
al. 2009, Söderström et al. 2010, Vilnet et al. 2010; see
also Appendix 9.5). Thus, in Russia Scapania is by far
the most species-rich liverwort genus (with 29 species),
whereas Lophozia is split into a number of separate genera
and has only eight species. In Greenland and Alaska,
Lophozia is the most species-rich genus (with 35 and 31
species, respectively), whereas Scapania has 24 and 20
species, respectively. These differences are mainly due
to different taxonomic concepts of these genera and
their families (Tab. 9.6, Appendix 9.5). There is also a
considerable variation in the moss genera Drepanocladus,
Calliergon (spearmoss) and Brachythecium (feathermoss)
(Hedenäs 1992). Other problems are associated with the
lack of knowledge in many areas influencing the results
of biodiversity studies on genus and family level (Afonina
& Czernyadjeva 1995, 1996).
Other important floristic publications in the last three
decades include checklists (e.g. Frisvoll & Elvebakk
1996, Afonina 2004) and regional and local monographs
(e.g. Lewinsky 1977, Schuster 1988, Afonina et al. 2005,
Belkina & Likhachev 2008, Konstantinova & Savchenko
2008, Damsholt 2010). Molecular phylogenetic studies of Arctic bryophytes are becoming more common,
and the results of such studies will certainly influence
taxonomic concepts and classification systems in the
future, and consequently the assessment and interpretation of diversity and origin of the Arctic bryoflora (e.g.
Konstantinova & Vilnet 2009, Konstantinova et al. 2009,
Söderström et al. 2010, Vilnet et al. 2010).
9.3.3. Regional surveys of Arctic bryodiversity
The present assessment of species richness in different regions of the Arctic is derived from heterogeneous sources that differ in age (from 2010 back to 1978)
333
Chapter 9 • Plants
Moss region
Census
Russia
1995/6
Svalbard
1996
Greenland
2003
Canada
1987
Alaska
1978
Number of moss species
530
288
497
343
408
Number of genera
154
103
133
103
136
Number of families
43
22
30
38
Number of species in genus
Bryum
39
18
42
25
26
Sphagnum
36
13
24
18
30
Pohlia
20
11
18
14
11
Dicranum
16
10
13
12
12
Hypnum
11
4
12
11
14
Encalypta
10
8
10
9
12
4
1
15
12
15
Grimmia
15
8
14
10
6
Brachythecium
20
6
19
9
8
8
12
15
6
7
Drepanocladus
Schistidium
Splachnum
Percentage of moss lora
5
5
5
5
5
32
22
36
30
29
65
33
64
44
45
Table 9.5. Species numbers of
species-rich moss genera and
families in the Arctic. Numbers
highlighted in coloured ields are
used in calculating the percentage
of the total moss lora. Listed are
Splachnum, genera with at least 10
species and families with at least
nine species.
Number of species in family
Bryaceae
Dicranaceae
53
31
48
35
37
Amblystegiaceae
46
19
50
40
46
Pottiaceae
43
14
33
31
44
Grimmiaceae
35
27
43
23
17
Sphagnaceae
36
13
24
18
30
Hypnaceae
34
10
25
19
23
Mniaceae
28
13
22
17
15
Brachytheciaceae
27
9
23
12
13
Polytrichaceae
20
13
15
15
15
Splachnaceae
18
9
14
10
17
77
62
73
77
74
Number of liverwort species
Percentage of Arctic moss lora
201
85
173
78
135
Total number of bryophyte species
731
373
670
421
543
and species concepts, classification and nomenclature.
These sources also pertain to regions of unequal size and
intensity of research. Primarily, sources for this assessment include the following: for Arctic Russia, Afonina
& Czernyadjeva (1995, 1996) (mosses) and Konstantinova et al. (2009) (liverworts); for Svalbard, Frisvoll &
Elvebakk (1996) (mosses and liverworts); for Greenland,
Goldberg (2003) (mosses) and Damsholt (2010) (liverworts); for the Canadian Arctic Archipelago, Ireland
et al. (1987) (mosses); and for Alaska, Steere (1978)
(mosses) and Steere & Inoue (1978) with later additions
of Potemkin (1995) (liverworts).
(Walker et al. 2005). The very small Arctic part of
northern Iceland and the Norwegian continent are not
covered, whereas for Canada only the Arctic Archipelago and the eastern Arctic are included. Transition
areas between taiga and tundra in Russia and Alaska are
probably partly included.
In the absence of a detailed circumpolar checklist of
bryophytes of the Arctic, the present comparative analysis and evaluation of these sources allow a global picture
of the variation of the Arctic bryoflora.
Only species are considered; infraspecies categories are
not considered here. Because of the absence of a checklist
of bryophytes of the Arctic, the nomenclature of species
and genera follows the literature used in the regional
checklists. Liverworts are primarily assigned to families
according to Damsholt (2002), otherwise to Konstantinova et al. (2009) and Steere & Inoue (1978). Moss
genera were primarily assigned to families according to
Brotherus (1923), otherwise to Afonina & Czernyadjeva
(1995, 1996).
The area covered by these publications is not completely
compatible with the Arctic territory of the CAVM
The time limitations of preparing the present assessment
did not allow inclusion of all new scattered literature
334
Liverwort region
Census
Number of liverwort species
Arctic Biodiversity Assessment
Russia
2009
Svalbard
1996
Greenland
2010
Canada
1947
Alaska
1978
201
85
173
78
135
Number of genera
73
34
50
28
49
Number of families
29
15
22
30
Scapania
29
16
24
20
Lophozia
8
16
35
31
Jungermannia
5
4
9
4
Cephalozia
9
4
5
5
Number of species in genus
Cephaloziella
11
2
12
4
Marsupella
8
2
10
2
Leiocolea
6
Lophoziopsis
7
Orthocaulis
5
Nardia
5
1
4
2
Barbilophozia
4
5
Gymnomitrion
4
3
5
2
Tritomaria
3
4
5
4
Anastrophyllum
1
1
4
6
46
44
61
46
Scapaniaceae
32
18
28
24
Jungermanniaceae
77
36
64
52
Gymnomitriaceae
14
6
18
4
Cephaloziaceae
13
7
11
8
Cephaloziellaceae
11
2
12
4
73
64
77
56
Percentage of liverwort lora
Table 9.6. Species numbers of
species-rich liverwort genera and
families in the Arctic. Numbers
highlighted in coloured ields are
used in calculating the percentage
of the total liverwort lora. Listed
are liverwort genera with at least
10 species and families with at
least 9.
3
Number of species in family
Percentage of liverwort lora
Number of moss species
530
288
497
343
408
Total number of bryophyte species
731
373
670
421
543
of recent years, such as e.g. on three Sphagnum species
new to the bryoflora of Greenland (Flatberg 2007). The
same applies for some recent molecular-based studies
on phylogeny and systematics of the liverwort families
Lophoziaceae, Scapaniaceae, Gymnomitriaceae and Jungermanniaceae (e.g. Söderström et al. 2010, Vilnet et al.
2010), which resulted into rearrangements of taxa.
9.3.3.1. Russia
About 731 bryophyte species are known from Arctic
Russia including 530 moss species and 201 liverwort species (Tab. 9.5, 9.6, 9.7). In the Russian Arctic, the 530
moss species include 154 genera in 43 families (Afonina
& Czernyadjeva 1995; Tab. 2 in Afonina & Czernyadjeva 1996) (Tab. 9.5). Prominent families (with more
than 10 genera) include Pottiaceae and Dicranaceae
(each 16) and Amblystegiaceae and Hypnaceae (feather
moss family; each 14). Mniaceae (thyme moss family),
Bryaceae and Polytrichaceae have seven genera each, and
Brachytheciaceae and Grimmiaceae each have six (Tab.
1 in Afonina & Czernadjeva 1996). Species diversity is
highest in Bryaceae (65 species), followed by Dicranaceae (forkmoss family; 53), Amblystegiaceae (feather-moss
family; 46), Pottiaceae (43), the monotypic Sphagnaceae
(36), Grimmiaceae (35), Hypnaceae (34) and Mniaceae
(28), Brachytheciaceae (27), Polytrichaceae (20) and
Splachnaceae (18). These families provide 77% of the
species of the moss flora. Species diversity is highest in
the genus Bryum (Bryaceae; 39 species) and Sphagnum
(Sphagnaceae; 36), followed by Pohlia (Bryaceae; 20),
Brachythecium (Brachytheciaceae; 20), Dicranum (Dicranaceae; 16), Grimmia (Grimmiaceae; 15) and Encalypta
(extinguisher moss, Encalyptaceae, extinguisher moss
family; 10). These genera account for 32% of the species
of the total Russian Arctic moss flora (Tab. 9.5).
Two hundred and one species of liverworts in 73 genera
and 33 families are reported from the Russian Arctic
(Konstantinova et al. 2009) (Tab. 9.6, Appendix 9.5),
and are here assigned to 29 families. Species diversity is
highest in Jungermanniaceae (sensu Damsholt 2002; 77
species) and Scapaniaceae (32) followed by Gymnomitriaceae (14), Cephaloziaceae (13) and Cephaloziellaceae
(11) (Tab. 9.6). Together they make up 73% of the total
Russian liverwort flora. Species diversity is highest
in the genus Scapania (Scapaniaceae; 29 species), followed by Cephaloziella (Cephaloziellaceae; 11), Cephalo-
335
Chapter 9 • Plants
zia (Cephaloziaceae; 9), the rustwort genus Marsupella
(Gymnomitriaceae; 8) and the genus Lophozia (Jungermanniaceae; 8). Lophoziopsis has seven species, Leiocolea
six and Jungermannia, Orthocaulis and Nardia (flapwort)
five each. These latter five genera are all classified here
as Jungermanniaceae. These genera comprise 46% of
the liverwort flora in the Russian Arctic. However,
molecular-based studies on phylogeny and systematics of
the Lophoziaceae, Scapaniaceae, Gymnomitriaceae and
Jungermanniaceae (e.g. Söderström et al. 2010, Vilnet et
al. 2010) have resulted in considerable rearrangements of
families and genera. Thus, quite different classification
concepts are certain to be considered in the future.
9.3.3.2. Svalbard
The small high Arctic island archipelago of Svalbard has
been relatively well investigated. About 373 bryophyte
species are known from here (Tab. 9.5 and 9.6). Frisvoll
& Elvebakk (1996) accepted 288 moss species in 103
genera for Svalbard. Genera with 10 or more species include Bryum (18 species), Sphagnum (13), Schistidium (12),
Pohlia (11) and Dicranum (10), collectively contributing
to 22% of the total moss flora of Svalbard. Species-rich
families include Bryaceae (with 33 species), Dicranaceae
(31), Grimmiaceae (27), Amblystegiaceae (19), Pottiaceae (14), Sphagnaceae, Mniaceae and Polytrichaceae each
with 13 species, Hypnaceae (10) and Brachytheciaceae
and Splachnaceae with nine species each, collectively
making up 62% of the Svalbard moss flora (Tab. 9.5).
The 85 liverwort species belong to 34 genera assigned
here to 15 families (Tab. 9.6, Appendix 9.6). Jungermanniaceae shows by far the highest species diversity
(36 species), followed by Scapaniaceae (18). These two
families collectively account for 64% of the liverwort
flora of Svalbard. Species numbers are highest in Lophozia
(including Lophoziopsis, Leiocolea sensu Konstantinova et al.
2009) and Scapania (16 each), followed by Barbilophozia
(pawwort, 5, including Orthocaulis). Recently, 14 species
were added to the bryoflora of the archipelago (Konstantinova & Savchenko 2006, 2008a, 2008b, 2012, Borovichev 2010).
9.3.3.3. Greenland
Around 670 bryophyte species are known from Greenland
(Tab. 9.5 and 9.6). Mogensen (2001) mentioned 478 moss
species (incl. 31 species of Sphagnum and seven of Andreaea)
from Greenland (cf. also Mogensen 1987, Jensen 2003).
However Lange (1984) and Crum (1986) accepted 23 species of Sphagnum. Goldberg (2003) listed 497 moss species
in 133 genera collected from Greenland and housed at the
Museum Botanicum Hauniense (Copenhagen). This is the
only ‘checklist’ currently available. Species-rich genera
include Bryum (42 species), Sphagnum (24), Brachythecium
(19), Pohlia (18), Drepanocladus and Schistidium (15 each),
Grimmia (14), Dicranum (13), Hypnum (12) and Encalypta
(10) (Tab. 9.5). Together, they comprise 36% of the total
number of moss species in Greenland (Tab. 9.5). The species number of Sphagnum is not up to date, since Flatberg
(2007) found three new species in Greenland, Sphagnum
concinnatum, S. tundrae and S. olaii. They are not considered in the present calculations (see Section 9.3.3). The
most species-rich families are Bryaceae (64 species), Amblystegiaceae (50), Dicranaceae (48), Grimmiaceae (43),
Pottiaceae (33), Hypnaceae (25), Brachytheciaceae and
Sphagnaceae (24 each), Mniaceae (22), Polytrichaceae (15)
and Splachnaceae (14). Together they account for 73% of
the total moss flora of Greenland (Tab. 9.5).
The estimated number of liverwort species is 135 (Mogensen (1987, 2001). A recent unpublished checklist of
the hepatics of Greenland by Damsholt (2010) comprises
173 species in 50 genera, assigned here to 22 families
(Damsholt 2002) (Appendix 9.7). Genus diversity is
highest in Jungermanniaceae (10 genera), followed by
Gymnomitriaceae and Cephaloziaceae (5 each) and
Aneuraceae, Aytoniaceae, Cleveaceae and Scapaniaceae
(three each). Together they account for 61% of the species diversity of the liverwort flora of Greenland. Species
diversity is highest in Jungermanniaceae (64 species),
followed by Scapaniaceae (28), Gymnomitriaceae (18),
Cephaloziellaceae (12) and Cephaloziaceae (11), together
accounting for c. 77% of the species diversity of the
entire liverwort flora of Greenland (Tab. 9.6).
9.3.3.4. Canada
At least 421 bryophyte species occur in Arctic Canada
(Tab. 9.5 and 9.6). Steere (1947) provided a first thorough account of the moss flora of the eastern Canadian
Arctic, including 304 species. Brassard (1971a) produced an impressive bryogeographical monograph of the
moss flora of the high Arctic Queen Elizabeth Islands,
NWT. and Nunavut, comprising 233 moss species. The
checklist of the mosses of Canada (Ireland et al. 1987)
accepts 343 species in 103 genera in 30 families for the
Arctic archipelago (Appendix 9.8), which represents
c. 35% of the total Canadian moss flora (965 species).
High generic diversity is found in Pottiaceae (14 genera),
Dicranaceae (13) and Amblystegiaceae (9), whereas
Polytrichaceae, Splachnaceae, Bryaceae, Mniaceae and
Hypnaceae each have five genera. Species diversity is
highest in the genera Bryum (25 species), followed by
Sphagnum (18), Pohlia (14), Drepanocladus and Dicranum
(12 each), Hypnum (11) and Grimmia (10) accounting for
30% of the moss flora, and in the families Bryaceae (44
species), Amblystegiaceae (40), Dicranaceae (35), Pottiaceae (31) and Grimmiaceae (23), followed by Hypnaceae (19), Sphagnaceae (18), Mniaceae (17), Polytrichaceae
(15), Brachytheciaceae (12) and Splachnaceae (10).
These families account for 77% of the total Canadian
Arctic moss flora (Tab. 9.5).
The liverwort flora is likely less well known. The most
comprehensive work to date is that by Polunin (1947)
referring to 72 species in 28 genera from the Canadian
eastern Arctic. Since then, many local studies have been
published (e.g. Steere & Scotter 1979, Scotter & Vitt
1992, Maass et al. 1994). In Schuster’s (1966-1992) comprehensive six-volume hepatic flora of North America,
336
new data on the eastern Canadian Arctic are accumulated; however, there is no published reference work
specifically for Canadian Arctic liverworts.
9.3.3.5. Alaska
Approximately 543 bryophyte species are known from
Alaska (Tab. 9.5 and 9.6), including 408 moss species
north of the Arctic Circle (Steere 1978) (Tab. 9.5).
They are referred to 136 genera and 38 families. The 11
most species-rich families are Amblystegiaceae (46 species), Bryaceae (45), Pottiaceae (44), Dicranaceae (37),
Sphagnaceae (30) and Hypnaceae (23), Grimmiaceae
and Splachnaceae (17 each), Polytrichaceae and Mniaceae (15 each) and Brachytheciaceae (13). Together they
account for 74% of the total species diversity of this part
of Alaska. The most species-rich genus is Sphagnum (with
30 species, Sphagnaceae), followed by Bryum (26, Bryaceae), Drepanocladus (15, Amblystegiaceae), Hypnum (12,
Hypnaceae), Dicranum (12, Dicranaceae), Encalypta (11,
Encalyptaceae) and Pohlia (11, Bryaceae). They contribute 29% of the total moss flora of this part of Alaska.
The liverwort flora of Arctic Alaska comprises at least
135 species in 49 genera and 30 families (Steere & Inoue
1978) (Tab. 9.6). The Lophoziaceae (with 43 species)
and Scapaniaceae (24) are the most species-rich families,
followed by Jungermanniaceae (9), Cephaloziaceae (8)
and Calypogeiaceae (5). The genera Lophozia and Scapania are by far the most species-rich (31 and 20 species,
respectively), followed by Cephalozia (5 species), and
Cephaloziella and Diplophyllum (earwort) (4 each). Assigning the genera and species to families according to Damsholt (2002), Jungermanniaceae and Scapaniaceae are
by far the most species-rich families (52 and 24 species,
respectively), accounting for 56% of the total liverwort
biodiversity of Arctic Alaska (Tab. 9.6).
9.3.4. Taxonomic structure of the Arctic
bryolora
Based on previous bryofloristic surveys (Section 9.3.3
and Tab. 9.5 and 9.6), the estimation of 850 species by
Matveyeva & Chernov (2000) cited in Callaghan (2005)
and several recent new records, we estimate the total
species number of the Arctic bryophyte flora to be c.
900. Arctic Russia is the most species rich (720 species),
followed by Greenland (670), Arctic Alaska (543), the
Canadian Arctic Archipelago (543) and the Svalbard
archipelago (373).
Species number of the mosses varies from 288 (Svalbard)
to 530 (Arctic Russia). Prominent moss families in all
five regions include Bryaceae, Dicranaceae, Pottiaceae
and Amblystegiaceae, Sphagnaceae and, to a lesser degree Grimmiaceae and Hypnaceae. Species numbers of
the ecologically specialized peat moss family Sphagnaceae are distinctly lower in Svalbard, the Canadian Arctic
Archipelago and Greenland (13, 18 and 24, respectively)
than in Arctic Alaska and Russia (30 and 36, respectively). The latter regions are contiguous to or belong to the
Arctic Biodiversity Assessment
northern boreal mainland of the large North American
and Eurasian continents (cf. Afonina 2004), where conditions for peat formation are more favorable than in the
high Arctic. The ecologically specialized dung moss family Splachnaceae is widely distributed in all five regions,
with five genera and 9-18 species. Throughout the Arctic
territory, the genera Bryum, Pohlia, Sphagnum, Dicranum,
Drepanocladus, Brachythecium, Schistidium and Grimmia are
well represented with many species.
Species diversity is distinctly lower for liverworts than
for mosses (78-201 liverwort species per region versus
288-530 moss species, Tab. 9.5 and 9.6). The most
prominent liverwort families with many species in the
Arctic include Jungermanniaceae and Scapaniaceae,
followed at far distance by Gymnomitriaceae, Cephaloziaceae and Cephaloziellaceae, while Scapania and Lophozia
are the most prominent genera with many species.
In Arctic Russia, 65% of the moss families and 95% of
the moss genera have fewer than 10 species, whereas
79% of the genera have fewer than five species (Tab. 1
and 2 in Afonina & Czernyadjeva 1996). For the liverwort families these values are 83, 97 and 85%, respectively (derived from Tab. 9.6).
Comparable percentages occur in the other Arctic
regions showing that, as in vascular plants, high species
diversity is restricted to a relative small group of genera
and families. This is a typical feature of Arctic floras
(Yurtsev 1994).
9.3.5. Large scale variation of species
richness
Bryophyte floras show variation in longitudinal and
latitudinal distribution related to climate and habitat
variation, different glaciations and migration histories
(e.g. Brassard 1971a, 1974, Schofield 1972, Steere
1976, 1979, Afonina & Czernyadjeva 1996). However, a
detailed survey of this variation as has been provided for
vascular plants does not exist at present for bryophytes.
9.3.5.1. Longitudinal variation
The longitudinal phytogeographical variation of the moss
flora is exemplified here for Arctic Russia (Tab. 9.7). The
Russian Arctic is divided from west to east into three
phytogeographical sectors with a total of 15 regions: the
European-W Siberian (EWS) sector with five regions,
Franz Joseph Land (ZF), Kanin-Pechora (KP), Polar
Ural (PU), Novaya Zemlya and Yamal-Gydan (YG); the
E Siberian (ES) sector with six regions, Taimyr (TA),
Severnaya Zemlya (SZ), Anabar-Olenik (AO), Kharaulakh (KH), Yana-Kolyma (YK) and Novosibirskiye Islands
(NS); and the Chukotka (C) sector with four regions,
Continental Chukotka (CC), Wrangel Island (WI), S
Chukotka (SC) and Beringian Chukotka (BC) (Tab. 9.7).
Afonina & Czernyadjeva (1996) pointed out that real,
existing diversity of the Russian Arctic moss flora de-
337
Chapter 9 • Plants
European-W Siberian (EWS)
39
Restricted
to region
395
Franz Joseph Land (ZF)
Number
of species
Restricted
to sector
Sector and region
Number
of species
Table 9.7. Numbers of moss species in sectors and regions of the
Russian Arctic after Afonina & Czernadjeva (1996).
113
Kanin-Pechora (KP)
255
3
Polar Ural (PU)
339
13
Novaya Zemlya (NZ)
203
4
Yamal-Gydan (JG)
252
3
309
8
E Siberian (ES)
385
27
Taimyr (TA)
Severnaya Zemlya (SZ)
99
Anabar-Olenik (AO)
192
4
Kharaulakh (KH)
282
6
Yana-Kolyma (YK)
248
6
174
2
Novosibirskiye Islands (NS)
Chukotka (C)
429
54
Continental Chukotka (CC)
171
1
Wrangel Island (WI)
241
5
S Chukotka (SC)
326
9
Beringian Chukotka (BC)
396
19
pends mainly on the knowledge of the regional floras,
the geographical position of the region and variation in
relief and diversity of habitats.
There are 395 species in the EWS sector, including 39
specific species (‘regional endemics’, so far only known
in the Russian Arctic from this sector). There are 385
recorded species from the ES sector including 27 specific
species, and 429 species from the C sector including 54
specific moss species. Species numbers are distinctly
different among regions. Species numbers of Severnaya
Zemlya (SZ) and Franz Joseph Land (ZF) are relatively
low (99 and 113, respectively), whereas Polar Ural (PU)
and Beringian Chukotka (BC) are rich in species (339
and 396, respectively). The Beringian Chukotka region
is the most diverse with 396 species, including 19 species only known in Russia from here. The high species
richness (429 species) of the Chukotka (C) sector (see
also Afonina 2004) and the presence of a high number of
species (54) not yet documented elsewhere in Russia may
be explained by geographical (variable mountain relief,
extensive areas of Paleozoic carbonate rocks) and historical (the existence in the past of the Bering Land Bridge)
factors. Some species are very rare and have not yet been
found elsewhere in Russia such as Funaria polaris, Seligeria
oelandica, Heterocladium procurrens, Orthotrichum pellucidum,
Racomitrium afoninae and Schistidium cryptocarpum. Other
species are rare with disjunct E Asian and North American distributions, e.g. Trachycistus ussuriensis, Bryoxiphium
norvegicum and Leptopterigynandrum austro-alpinum. Thus
the paramount position of Beringian Chukotka (and of
Wrangel Island, which belongs to Beringia as well), is
not only expressed in its vascular plant flora (Section
9.2), but also in its highly diverse and distinct moss flora
(see also Afonina 2004).
Mountain moss floras appear richer than Arctic plain
floras, as demonstrated by comparing the regions Polar
Ural (339 species) and Yamal-Gydan (252 species).
The richer flora of Polar Ural is certainly related to the
higher diversity of local climate and habitats.
Around 12% of all moss species of Arctic Russia are
common and widely distributed, and are often locally
dominant in the vegetation all over the Russian Arctic
(derived from Afonina & Czernyadjeva 1995). They
include many mainly circumboreal/Arctic-alpine species
(e.g. black rock-moss Andreaea rupestris, Aulacomnium turgidum, Brachythecium turgidum (turgid brachythecium moss),
Bryoerythrophyllum recurvirostrum (red beard-moss), Bryum
cyclophyllum (round-leaved bryum), Campylium stellatum
(bog star-moss), Cinclidium arcticum (Arctic cupola-moss),
Conostomum tetragonum (helmet-moss), Dicranum spadiceum,
Ditrichum lexicaule (slender-stemmed hair moss), Encalypta
rhaptocarpa (ribbed extinguisher-moss), Hylocomium splendens (glittering woodmoss), Limprichtia revolvens, Meesia
triquetra (three-ranked hump-moss), Oncophorus wahlenbergii (Wahlenberg’s spur-moss), Paludella squarrosa (tufted
fen-moss), Pohlia cruda (opal thread-moss), Polytrichastrum
alpinum (alpine haircap), Polytrichum piliferum (bristly
haircap), Racomitrium lanuginosum, Sanionia uncinata,
Sphagnum teres (rigid bogmoss), S. warnstorii (Warnstorf’s
bogmoss), Syntrichia ruralis (a screw-moss), Tetraplodon
mnioides (slender cruet-moss), Tomenthypnum nitens (woolly
feather-moss or golden silk moss) and Warnstoria exannulata (ringless hook-moss). Most of these species occur in
plant communities of the circumpolar classes Scheuchzerio-Caricetea (mires and rich fens), Carici-Kobresietea (graminoid and prostrate dwarf shrub vegetation on non-acidic
substrates) and snow beds (Salicetea herbaceae) (cf. Sieg et
al. 2006). Around 32% of the moss species have a sporadic distribution, occurring in a majority of the regions.
A large number of species, c. 40%, are considered rare,
known from only a few regions, although this may partly
be due to poor knowledge of some difficult taxonomic
groups. About 16% of the species are so far known only
from one locality in the Russian Arctic.
9.3.5.2. Latitudinal variation
High Arctic areas have fewer species than low Arctic
areas, as exemplified by comparing the numbers of moss
species of the non-Beringian E Siberian high Arctic
Novosibirsky Islands (NS) with that in the adjacent low
Arctic Yana-Kolyma (YK) region in Arctic Russia (174
and 248 species, respectively) (Afonina & Czernyadjeva
1996) (Tab. 9.7). The Polar Ural (PU) in the low Arctic
has 339 species, whereas Novaya Zemlya (NZ) in the
high Arctic, which belongs to the same Euro-Siberian
(EWS) sector, has only 203 species. In the Chukotka (C)
sector, Wrangel Island (WI) has 241 species, whereas
Beringian Chukotka (BC) to the south has 396. Species
richness of the Russian Arctic Islands is relatively low.
338
These islands are isolated from mainland Russia and situated at high latitudes, mostly in the high Arctic.
Other examples of latitudinal variation are found in
Greenland, where 134 moss species were documented
in Peary Land, N Greenland (Holmen 1960) versus
nearly 500 species for all of Greenland (Goldberg 2003).
Mogensen (1987) showed that c. 58% of the Greenlandic
moss flora consists of species with a southern distribution in Greenland, 18% are widely distributed and 24%
are northern species.
The northern local floras in high Arctic Ellesmere Island
have fewer species as compared with floras on the southern part of the island. The moss flora of the Lake Hazen
area, northern Ellesmere Island, comprises c. 45 species
(Powell 1967), whereas 84 moss species are reported
from the Alexandra Fiord lowlands in the middle of
Ellesmere Island (Maass et al. 1996).
The same trend is seen in liverwort floras, where diversity
decreases with increasing latitude. The liverwort flora of
S Greenland includes 139 species (Schuster 1988), that
of W Greenland (66° N-72° N) 136 species (Schuster &
Damsholt 1974), whereas northern floras seem to have
fewer species, e.g. Peary Land, N Greenland with 25 species (Arnell 1960), Svalbard with 85 (Frisvoll & Elvebakk
1996), the eastern Canadian Arctic with 72 (Polunin
1947) and northern Ellesmere Island with 43 (Schuster
1959). However all these figures should be considered
with caution, since the state of knowledge of the liverwort
flora from these areas varies greatly.
9.3.6. Origin of Arctic bryoloras and
distribution types
So far, assumptions on the origin of Arctic bryofloras
are generally based on the analysis of distribution patterns (e.g. Schuster & Damsholt 1974, Schuster 1988),
environmental and geographical conditions, glaciation
histories and refugia, degree of taxonomic isolation of
species (Brassard 1971a, 1974) and (sub)fossil records
(e.g. Mogensen 1984, Hedenäs 1994). However, ongoing
and future phylogenetic and phylogeographic studies certainly will add new insights and conclusions to this field.
Apart from several cosmopolitan mosses such as Bryum
argenteum (woolly silver moss) and Ceratodon purpureus
(redshank), and liverworts such as Aneura pinguis (greasewort) and Cephalozia bicuspidata (two-horned pincerwort)
together with some small groups of ecologically or geographically disjunct species, three general distribution
types are of special interest in a broader, circumpolar
Arctic perspective: circumboreal species, Arctic species
and amphi-Beringian species (cf. Schofield 1972, Steere
1976, 1979).
The circumboreal species are widely distributed in
temperate, boreal and Arctic climates of the Northern
Hemisphere. Most (c. 75-80%) of the bryoflora of
Arctic Alaska belongs to this group, and this percent-
Arctic Biodiversity Assessment
age might hold for the entire Arctic. Most of the species
considered sporadic (S), common (C) and widespread
(W) in Arctic Russia (Afonina & Czernyadjeva 1995,
1996, Afonina 2004) (see Section 9.3.5.1.) belong to this
distribution type. Most of these species are believed to
have expanded their distribution by colonizing the devegetated and deglaciated areas in the circumpolar North
after the Pleistocene glaciations (Crum 1966, Steere
1976, 1978, 1979).
Arctic (including Arctic-alpine) species comprise around
15% of the North American bryoflora (cf. Brassard
1971a, 1974, Steere 1979). Possible explanations for the
presence of these bryophyte species in the high Arctic
Canadian Queen Elizabeth Islands were thoroughly addressed by Brassard (1971a). In a later publication, the
possible evolution of these Arctic bryophyte taxa was
plausibly explained by taking into account their degree
of taxonomic isolation and a possible area change as a
Tertiary taxon, a newly evolved Quaternary taxon or a
recent taxon (Brassard 1974). Arnellia fennica, Bryobrittonia pellucida (Fig. 11 in Afonina 2004), Aplodon wormskjoldii, Philocrya aspera (Fig. 4 in Afonina 2004) might
represent old species already present in the Arctic before
the Quaternary. Regarding Arctoa anderssonii, Cyrtomnium
hymenophylloides, C. hymenophyllum, Psilopium cavifolium
(Fig. 2 in Schofield 1972), Seligeria polaris and Voitia
hyperborea (Fig. 9 in Afonina 2004), it is assumed that
these Arctic species might have evolved before or during
the Quaternary (cf. also Mogensen 1984). Aulacomnium
acuminatum, Cinclidium latifolium, C. arcticum, Timmia
comata, Tritomaria heterophylla, Bryum wrightii, Didymodon
leucostomus, D. johansenii, Fissidens arcticus, Funaria polaris
(Fig. 8 in Afonina 2004), Hygrohypnum polare, Hygrolejeunea polaris, Oligotrichum falcatum (Fig. 6 in Afonina 2004),
Scapania simmonsii and Trichostomum cuspidatissimum (Fig.1
in Schofield 1972) may be pre-Quaternary taxa or ones
that evolved during the Quaternary. Two other groups
totalling c. 24 species (Tab. 3 in Brassard 1974, see also
Schofield 1972) and including Barbula icmadophila, Campylium arcticum, Ceratodon heterophyllus, Distichium hagenii,
Mnium blyttii, Rhizomnium andrewsianum, Seligeria pusilla,
Tortella arctica, Bryum arcticum, B. calophyllum, B. cryophilum, Drepanocladus badius, D. brevifolius (Fig. 3 in Schofield
1972) and D. lycopodioides together with Lophozia species
such as L. hyperarctica, L. pellucida and L. quadriloba likely
represent young taxa that presumably evolved during
the early or late Pleistocene (Brassard 1974). Other taxa
mainly restricted to the Arctic proper with a circumpolar distribution include Cnestrum glaucescens (Fig. 27 in
Afonina 2004), Timmia sibirica (Fig. 34 in Afonina 2004),
Plagiothecium berggrenianum (Fig. 39 in Afonina 2004),
Schistidium cryptocarpum (Fig. 18 in Afonina 2004),
Sphagnum arcticum (Fig. 2 in Afonina 2004) and Encalypta
brevipes (Fig. 12 in Afonina 2004).
The smaller Arctic amphi-Beringian moss flora includes
Drepanocladus latinervis, Pohlia beringiensis (Fig. 30 in
Afonina 2004), Rhizomnium gracile (Fig. 31 in Afonina
2004), Schistidium andreaeopsis, Bryoxiphium norvegicum,
Grimmia pilifera, Herzogiella adscendens, Pseudotaxiphyllum
339
Chapter 9 • Plants
elegans, Didymodon subandreaeoides, Leptopterigynandrum
austro-alpinum (Afonina & Czernadjeva 1996, Fig. 37
in Afonina 2004) and Racomitrium afoninae (Fig. 15 in
Afonina 2004). Their distribution pattern is explained
by the existence of an ice-free land bridge between
northeastern Asia and northwestern North America during the last glaciations (Afonina 2004). These species are
likely fragments of a Tertiary bryoflora (Steere 1969).
Bryophytes have in general a wider distribution (reproduction by light spores and vegetative reproduction) than
vascular plants, and as a result endemism is much lower
in this group.
Endemism of Arctic bryophytes is apparently much higher
on the infra-species level (N.A. Konstantinova, unpubl.).
Endemism in liverworts was considered high in northern
Ellesmere Island and Greenland, maybe as the result of
refugia during the last Wisconsin Glaciation (cf. Schuster
1959, Holmen 1960, Brassard 1971a). However, most of
these species were later recorded in many Arctic regions
and must therefore now be considered pan-Arctic ‘endemics’.
Nevertheless, a large number of liverworts are restricted
to the Arctic and adjacent mountain ridges of the subArctic. Such species are defined in Russian geobotanical
literature as ‘meta-Arctic’ species.
According to Konstantinova (2000) 44 liverwort species
have a predominantly Arctic distribution. Most of these
are poorly known and were described mainly from Ellesmere Island and Greenland (Schuster 1969-1992, 1988,
Schuster & Damsholt 1974). During the last decades,
many species regarded as endemics of Ellesmere Island
or Greenland were found in several other regions of the
Arctic and sub-Arctic, particularly in Svalbard, Russia
and Alaska (e.g. Anastrophyllum sphenoloboides, Cephaloziella
polystratosa, Lophoziopsis (Lophozia) pellucida and Lophoziopsis
(Lophozia) polaris). Some recently described ‘Arctic’ liverworts were synonymized with earlier known species (e.g.
Leiocolea katenii was synonymized with Leiocolea badensis).
Several species referred by Konstantinova (2000) from
the Arctic were collected in the alpine belt of the Caucasus in southern Russia (e.g. Cephaloziella aspericaulis).
Moreover a majority of liverwort species considered previously as Arctic have been collected in the mountains of
Siberia and the Russian Far East. These areas are phytogeographical corridors (or ‘bridges’) between the Arctic
and the mountains of South and East Asia.
Nowadays only a few liverworts species can be considered
as true Arctic endemics. They were recently described
and only known from a single locality (e.g. Scapania matveyevae) or several localities (Gymnocolea fascinifera, Schistochilopsis hyperarctica). It is quite possible that most of such
taxa will turn out to be more widespread in the Arctic.
Most of the Arctic, sub-Arctic and alpine species (‘metaArctic’ species) have a circumpolar distribution. Several
species are really ‘old’ endemics, some are evidently
neoendemics (Schuster & Konstantinova 1996).
The majority of ‘old’ isolated species occur in the Beringian sector of the Arctic (e.g. Pseudolepicolea fryei, Radula
prolifera), and such species are absent in W Siberia and E
Canada. Thus as for vascular plants Beringia is an important refugium of a presumably relictual tertiary bryophyte flora (see also Schuster & Konstantinova 1996).
9.3.7. Trends
We refrain from speculations about changes in bryofloras, due to insufficient knowledge in many Arctic areas.
There are no known threatened species.
9.3.8. Conclusions and recommendations
The estimated species number of the bryophyte flora of
the Arctic is moderate (c. 900) compared with that of
lichens (c. 1,750) and vascular plants (c. 2,218). But it is
likely that this number will increase significantly in the
course of future studies. Arctic endemism is not strongly
pronounced, and is displayed mainly on an infra-species
level. The Arctic bryoflora is rather uniform. Almost
80% of the species have a broad circumboreal and
circumpolar distribution. In rather stable, wet-to-moist
sites they strongly contribute to vegetation biomass, and
they also contribute to species richness of many vegetation types in other habitats. Their ecosystem function is
poorly studied, and overall the bryofloras of most Arctic
regions are still incompletely known. Moreover, Arctic
material in the majority of taxonomic groups needs revision using modern molecular phylogenetic approaches (cf.
Konstantinova & Vilnet 2009, Söderström et al. 2010).
Records of localities of rare and recently described species need verification. There are no known threatened
species. The use of bryophytes by indigenous peoples
is very restricted. A circumpolar checklist according
to uniform taxonomic concepts and nomenclature is
urgently needed and will be highly beneficial for vegetation and ecosystem studies, especially for monitoring and
interpretation of change in the face of climate change.
9.4. ALGAE
This section surveys both freshwater and marine environments by inventorying the biodiversity of the algal
flora at a pan-Arctic scale in terms of species richness
and distribution. The current exercise should be taken as
a snapshot of the present situation regarding the accumulated knowledge of the biodiversity of these micro- and
macroalgal organisms.
Algae are oxygenic autotrophic eukaryotes characterized by chloroplasts containing chlorophyll and other
associated pigments, and reproducing by the formation
of spores and gametes. They differ from vascular plants
in physiological, cellular and morphological aspects.
However, the cyanobacteria are addressed here since
340
they were classified as blue-green algae with the Cyanophyceae for many years and are functionally oxygenic
autotrophic prokaryotes. All algae contain the photosynthetic chlorophyll a pigment. The eukaryotic algae
comprise heterogeneous and evolutionarily different
groups. The origin and development of the first eukaryotic algae is explained through an endosymbiotic event
where a heterotrophic eukaryote acquired or enslaved an
ancestral cyanobacterium (cf. Reyes-Prieto et al. 2007).
After genetic reduction and transformation, this event
gave rise to primary plastids (chloroplasts) present in the
Glaucophyta, Rhodophyta (red algae) and Chlorophyta
(green algae); the three lineages are classified as Plantae
with the higher plants (van den Hoek et al. 1993, Adl et
al. 2005, Raven et al. 2005, Cocquyt et al. 2010). The
Chlorophyta are ancestral to the algal Streptophyta and
hence to the bryophytes (Bryophyta) and vascular plants
(Tracheophyta). All are predominantly green with chlorophyll b as a secondary pigment.
Other algae are polyphyletic, lacking an identifiable
common ancestor, and for the most part, their chloroplasts originated as a secondary endosymbiotic event
where a single-celled pre-rhodophyte alga was acquired
or enslaved by another heterotrophic protist (see Lovejoy, Chapter 11). Over time, this ancestral red lineage is
thought to have given rise to other major algal phyla (e.g.
Friedl et al. 2003, Falkowski et al. 2004, Reyes-Prieto
et al. 2007, Armbrust 2009, Cocquyt 2009). Chlorophyll c is a secondary pigment common to most of these
other algae, and the Chromalveolata is a term used to
designate a supergroup of all the chlorophyll c containing
algae and their non-chloroplastic relatives. These algae
include the Dinophyta and the diverse heterokont algae.
Two algal phyla, the Cryptophyta and Haptophyta, with
chlorophyll c are now thought to have arisen through
separate endosymbiotic events with different protists
(Baurain et al. 2010), and since their phylogenetic positions are uncertain, the term Chromalveolata is used
descriptively in this text. There are two other algal phyla
that arose from endosymbiotic events where singlecelled green algae gave rise to chlorophyll b-containing
chloroplasts in the photosynthetic Euglenophyta and
Chlorarachniophyta. Several dinoflagellates from diverse
lineages have lost their original secondary, endosymbiotically acquired chloroplast and have acquired new
chloroplasts directly from green algae, cryptophytes and
even diatoms in what are termed tertiary endosymbiotic
events (Keeling 2010).
Algae are ecologically very important, contributing to
the biogenic carbon flux throughout aquatic systems,
being at the base of marine and freshwater food webs
(Forest et al. 2011). Algae have a worldwide distribution occurring in nearly all wet or aquatic habitats, on
land (terrestrial), in freshwater (limnic) and in seawater
(marine). They occur either free in the upper water
column (pelagic) and known as phytoplankton which
encompass autotrophic, single-celled eukaryotes ranging in size from 0.2 to 200 µm, and further segregated
into pico- (< 2 µm), nano- (2-20 µm) and micro-sized
Arctic Biodiversity Assessment
(20-200 µm) fractions of the scaling plankton classification (Sieburth et al. 1978). In a broader sense, plankton
includes microzooplankton, non-autotrophic eukaryotic
protists, bacteria, Archaea and viruses (Thomas et al.
2008, Poulin et al. 2011). Algae are also associated with
polar sea ice (sympagic, Różańska et al. 2009, Poulin et
al. 2011), or attached to soft and hard bottom substrates
(benthic, Totti et al. 2009). Single-celled algae as well as
large macroalgae (seaweeds) live in the intertidal zone.
Attached, benthic marine macroalgae are the main habitat structuring agents of several major marine ecosystems, such as kelp forests and sub-tidal red algal crusts.
Marine phytoplankton and sympagic algae are at the
base of the Arctic marine food web. Marine phytoplankton are responsible for more than 45% of the annual
net primary production of the Earth (Falkowski et al.
2004, Simon et al. 2009). Diatoms alone are responsible
for 20% of the Earth’s annual net primary production,
generating as much carbon as all terrestrial rainforests
together (Armbrust 2009). In Arctic seas, sympagic
algae contribute 57% of the total primary production in
the central Arctic Ocean (Gosselin et al. 1997) and up to
25% on Arctic shelves (Legendre et al. 1992).
Algae exhibit a tremendous variability in morphology from unicellular solitary and colonial microalgae
to multicellular macroalgae (e.g. seaweeds). Their size
range varies from 0.2 µm for pico-sized cells to more
than 100 m long Phaeophyta in giant kelps (Thomas et
al. 2008). Algae differ from vascular plants, bryophytes
and lichens by their diversified cell wall compounds,
which can be taxonomically specific with, for example,
the siliceous casing characterizing the Bacillariophyta
(diatoms) or the calcium carbonate distinctive of the
Haptophyta coccolithophorids. Other algae have cell
walls containing cellulose or chitin. In addition to some
differences in the main chlorophyll pigments, accessory
pigments also differ among algae, with different phyla
having specific profiles of xanthophylls (carotenoids
with molecules containing oxygen). Cryptophyta and
a few dinoflagellates with cryptophyte-origin plastids
contain phycobiliproteins that also absorb photons. Storage products also differ with starches, sugars and lipids
found among different phyla in various proportions. Life
cycles are haplontic, diplohaplontic or diplontic, without
embryonic stadia (e.g. van den Hoek et al. 1993, Raven et
al. 2005), and algae do not produce an early sporophytic
generation embedded in parental tissue (Friedl et al.
2003). The mode of locomotion among algae is highly
diverse; only the Rhodophyta and centric diatoms do
not usually have a flagellated stage. However, benthic
pennate diatoms are well known for their movement on
substrates and vertical migration through soft ediments
(Round et al. 1990). In addition, most algal groups lack
functional anatomical and morphological differentiation as shown in vascular plants (e.g. van den Hoek et al.
1993, Raven et al. 2005).
Cyanobacteria (blue-green algae) in the domain of
Bacteria (van den Hoek et al. 1993, Raven et al. 2005)
341
Chapter 9 • Plants
are fundamentally different from autotrophic eukaryotic
algae. Cyanobacteria lack a membrane-bound nucleus
and organelles. Most do not have accessory chlorophylls
(b or c). Cyanobacteria exhibit a very diverse morphological range from solitary coccoid cells to colonies with
simple thalli of a more restricted size range compared
with eukaryotic algae. Their blue-green color is caused
by high amounts of the accessory pigments phycocyanin
and allophycocyanin (e.g. van den Hoek et al. 1993). In
marine systems as well as in some deep freshwater environments, the accessory pigment phycoerythrin masks
the phycocyanin giving a pink coloration to the cyanobacteria. Cyanobacterial mats exposed to high ultraviolet
radiation in polar regions have high concentrations of
photoprotective sunscreen and other pigments and can
be black or orange (Vincent 2000). They are able to fix
atmospheric nitrogen and are thus key players in global
nitrogen budgets as well as in carbon flux.
Additionally, both green and blue-green algae can be
the photosynthetic partner for fungi to form lichens (see
Dahlberg & Bültmann, Chapter 10). They can also live
independently of lichens and with bryophytes and lichens
they cover bare soil as ‘black crusts’ and improve growth
conditions for vascular plants, which is particularly important in the Arctic (Elster et al. 2002).
Table 9.8. Worldwide recognized and estimated numbers of algal
species. Sources: Norton et al. (1996) and Poulin & Williams (2002).
Taxonomic
group
Recognized
taxa
Estimated
taxa
c. 2,000
?
Prasinophyceae
incl. Pedinophyceae
120-140
500
Ulvophyceae
1-1,000
3,000
2,500 -2,600
10,000-100,000
11,000-13,000
20,500
Prokaryotic algae
Cyanophyta
Eukaryotic algae
Archaeplastida
Chlorophyta (1)
Chlorophyceae
Streptophyta (2)
Charophyceae
incl. Zygnematales
Glaucophyta
13
50
Rhodophyta
4,000-6,000
5,500-20,000
200
1,200
300-500
2,000
2,000-4,000
3,500-11,000
Dictyochophyceae
incl. Pedinellophyceae
10
15
Eustigmatophyceae
12
1,000-10,000
Pelagophyceae
7
20
10,000-12,000
100,000-200,000
900-1,000
2,000
Chromalveolata
Cryptophyta
Haptophyta
(Prymnesiophyceae)
Dinophyta (7)
Stramenopiles (8)
9.4.1. Major algal groups
The diversity of marine phytoplankton is impressive.
About 25,000 species are known to date (Norton et al.
1996, Poulin & Williams 2002, Falkowski et al. 2004,
Poulin et al. 2011). Over the last 15 years with the
increasing use of molecular tools there has been some
upsurge in taxonomic descriptions, especially among
some of the smaller, less speciose groups such as Chlorarachniophyta (Ota et al. 2011). The major taxonomic
divisions discussed in the following sections are presented in Tab. 9.8, with phylogeny based on a consensus
of recent works (Adl et al. 2005, Raven et al. 2005,
Cocquyt 2009, Baurain et al. 2010, Cocquyt et al. 2010,
Keeling 2010, Marin & Melkonian 2010).
Among the Chlorophyta (chlorophytes, green algae sensu
stricto), the most frequently reported classes in the Arctic
are Chlorophyceae, Mamiellophyceae, Pedinophyceae,
Prasinophyceae and Ulvophyceae. The Chlorophyceae
are often found in snow and ice (Müller et al. 1998, Larose et al. 2010), whereas the Prasinophyceae are mainly
planktonic but also occur frequently in Arctic sea ice
(Poulin et al. 2011). The Pedinophyceae and Mamiellophyceae are truly planktonic, and all known species are
less than 5 µm in size. One phylotype of Micromonas in
the Mamiellophyceae is likely the most abundant singlecelled type in the Arctic Ocean (Lovejoy et al. 2007).
The Streptophyta (streptophytes) include all green land
plants and Charophyceae as well as the Zygnematales
(Leliaert et al. 2012). Charophytes (stoneworts) and Zygnematales occur predominantly in freshwater habitats,
and they have a worldwide distribution. The Zygne-
Bacillariophyceae
Phaeophyceae
Xanthophyceae
600-700
2,000
Chrysophyceae
1,000-2,000
2,400
15-27
100
900-1,000
2,000
60
?
<5
20
Rhaphidophyceae
Excavata
Euglenophyta (9)
Opisthokonta
Choanolagellidae (10)
Rhizaria
Chlorarachniophyta (11)
matales are highly diverse and often used as indicator
species of water quality; they have been reported from
Arctic tundra streams (Sheath et al. 1996).
The Rhodophyta are also highly diverse, and although
many have a tropical distribution, they occur in cold
waters and are present in freshwater, including in the
Arctic (Sheath et al. 1996).
The Chromalveolata constitute the group with the
highest diversity of autotrophic eukaryotic algae with
chlorophyll c pigment, including two major groups: the
stramenopiles and the alveolates. The alveolates are
defined by their cell wall characteristics, and include
ciliates, dinoflagellates and parasitic taxa (see Lovejoy,
342
Chapter 11). The autotrophic alveolates, with a few
exceptions, are in the Dinophyta (dinoflagellates). The
stramenopiles (heterokonts) have the highest diversity
within the Chromalveolata. There are six major groups
of algae within the stramenopiles that are frequently
reported from the Arctic. The Bacillariophyta (diatoms)
are planktonic, sympagic and benthic, occurring in
terrestrial, freshwater and marine habitats. The Eustigmatophyceae, Raphidophyceae and Dictyochophyceae
are mostly marine planktonic species and have been
rarely reported from Arctic seas (Comeau et al. 2011,
Poulin et al. 2011), except perhaps for Dictyocha speculum,
which occurs regularly all year round in Arctic seas. The
Chrysophyceae (golden-brown algae) are mostly limnic
and often dominate Arctic and sub-Arctic lakes (Charvet et al. 2011) and tundra streams (Sheath et al. 1996),
occasionally occurring as Arctic marine phytoplankton
(Lovejoy et al. 2002, 2006, Poulin et al. 2011), with
Dinobryon balticum dominating in late summer – early fall
around Svalbard. Most species are single cells but some
species are colonial including the spectacular arborescent colonies of Dinobryon. The Xanthophyceae (yellowgreen algae) are mostly freshwater species. Among the
macroalgal stramenopiles are the Phaeophyta (Phaeophyceae, brown algae), widespread along the Arctic coasts.
Also within the Chromalveolata, the Cryptophyta
(cryptophytes) and Haptophyta (haptophytes or prymnesiophytes) branch apart from other chlorophyll c containing algae and also from each other (Baurain et al. 2010).
The Cryptophyta are unicellular organisms and can have
a variety of colors derived from the phycoerythrin and
phycocyanin pigments. The cryptophytes are reported
from both cold marine and limnic environments (Lovejoy et al. 2002, 2006, Charvet et al. 2011, Poulin et al.
2011). The Haptophyta are unicellular algae and they
are mainly marine and coastal, although the calciumcarbonate-scale-bearing coccolithophorids are scarce in
Arctic regions. Small flagellated haptophytes that do not
have coccoliths are common in polar marine plankton.
The Euglenophyta (euglenids) belong to the lineage
Excavata and are mainly limnic, unicellular flagellates,
with some species present in Arctic seas (Poulin et al.
2011). The chlorarachniophytes (Chlorarachniophyta)
have been reported in the Arctic from molecular biological surveys (see Lovejoy, Chapter 11) and the group
belongs to the Rhizaria lineage. The Opisthokonta are
represented by small celled, non-autotrophic choanoflagellates considered in the broader sense as belonging
to marine phytoplankton. They are characterized by the
formation of a siliceous lorica protecting the cell and
have been recorded mainly in coastal Arctic seas (Thomsen et al. 1997).
9.4.2. Arctic algal taxonomic diversity and
regionality
The taxonomic diversity of algae on a worldwide scale
is estimated to be extremely high (Norton et al. 1996,
Poulin & Williams 2002). Many phycologists assume
Arctic Biodiversity Assessment
that the number of described species, varying between
30,000 and 40,000, is only a small fraction of the
total number of undescribed species estimated to vary
between 400,000 and more than 10 million (Norton et
al. 1996), this last figure having to be taken with great
caution. More recently, the number of diatom species
was estimated around 200,000 by both Poulin & Williams (2002) and Armbrust (2009). Recent environmental surveys using molecular techniques suggest that the
diversity at all levels has been underestimated, and much
work needs to be done to even make informed estimates
of how many species of algae exist on Earth.
The total species number of algae in the Arctic is unknown and assumed to be much lower than in warmer
regions of comparable size (e.g. van den Hoek 1984,
Lüning 1985, Norton et al. 1996, Kerswell 2006),
although recently Archambault et al. (2010) reported a
lower number of marine seaweeds and a higher number
of marine phytoplankton in the Canadian Arctic compared with eastern and western Canada. Seaweeds from
the Arctic were characterized by Kjellman (1883) as
having “monotony and luxuriance.” Species diversity of
the seaweed Chlorophyta, Phaeophyta and Rhodophyta
(Norton et al. 1996; Tab. 9.8) showed distinctly lower
species numbers for the Arctic regions: 80 and 252 taxa
for the Bering Sea and Arctic Ocean, respectively (Kjellman 1883), and 201 and 280 taxa for Arctic Canada and
Greenland, Iceland, Svalbard and northern Norway,
respectively (South & Tittley 1986). Species numbers of
algae from temperate and tropical regions were generally
much higher, with up to 1,058 species in the Caribbean
and adjacent waters (Wynne 1986) and 1,510 in Japan
(Yoshida et al. 1990).
The species diversity of microalgae in the Arctic, as
elsewhere, is generally poorly known. Okolodkov &
Dodge (1996) reported the occurrence of 250 species of
planktonic dinoflagellates from various localities in the
Arctic Ocean. All species appeared common in Arcticboreal marine waters, and species diversity was higher
in regions influenced by an influx of warmer water from
the south. In Arctic regions, marine diatoms are very
diverse and abundant in both annually formed sea ice and
pelagic waters (von Quillfeldt et al. 2003, Różańska et al.
2009, Poulin et al. 2011).
Recent molecular studies reported a high diversity in the
smallest sized-fraction of the phytoplankton in polar regions, frequently contributing to more than 50% of the
total phytoplankton biomass and production (Lovejoy et
al. 2006, Poulin et al. 2011). A more recent study using
high throughput sequencing technology to better capture
rare species indicates that in the Beaufort Sea, western
Canadian Arctic, alone there may be on the order of
10,000 single-celled eukaryotic species, at least half of
which are likely autotrophic (Comeau et al. 2011).
The knowledge of taxonomic biodiversity and geographical variation of marine, freshwater and terrestrial algal floras across the Arctic regions is obviously
343
Chapter 9 • Plants
less than that of the predominantly terrestrial vascular
plant and bryophyte floras. The knowledge of terrestrial and freshwater algae, including cyanobacteria, of
the comparatively well-explored Svalbard archipelago
was considered “still in its infancy” by Skulberg (1996),
whereas the bryophyte and vascular plant floras are
comparatively well known (Elvebakk & Prestrud 1996).
Nevertheless, knowledge of the diversity of marine microalgae distinctly increased in the last decades because
of increased and improved sampling techniques and
culture protocols, advanced microscopy and molecular
biology research methods, electronic archiving databases and gene libraries, and increased international
cooperation through climate change and biodiversity
research programs (e.g. ArcticNet, Conservation of
Arctic Flora and Fauna, Arctic Ocean Diversity-Census
of Marine Life).
The main difficulties in assessing biological diversity at
the subgeneric levels are the dissimilarities that exist in
the taxonomic species concept and classification among
the circumpolar countries. Moreover, current species
concepts from traditional morphological assessments are
challenged by the latest molecular phylogenetic analyses
(e.g. Pröschlod & Leliaert 2007), which may or may not
support traditional classifications. In particular for microalgae, this dichotomy between fundamental morphological and molecular phylogenetic interpretations makes
the assessment of their geographical distribution very
hard (cf. Rindi et al. 2009). The morphological variation
in picophytoplankton (< 2 µm) is poorly addressed with
current microscopic investigations, whereas its molecular biodiversity is enormous (Lovejoy et al. 2006). A
major challenge facing biodiversity assessments will be
matching morphology of a single-celled alga to a given
gene sequence, which obviously will mean developing
better sampling strategies and culture techniques for
these small-sized microalgae.
Since the first algal biodiversity census by Norton et
al. (1996), various floristic or biogeographic reports of
local or regional character have been reported (e.g. Lee
1980, Okolodkov & Dodge 1996, von Quillfeldt 1997,
Cremer 1998, Okolodkov 1998, Stenina et al. 2000, von
Quillfeldt et al. 2003, Kerswell 2006, Lindstrom 2006,
Matuła et al. 2007), including more recent pan-Arctic algal biodiversity assessments (e.g. Adey et al. 2008, Wulff
et al. 2009, Bluhm et al. 2011, Poulin et al. 2011).
The present contribution can be regarded as a snapshot
of the algal diversity obtained from a number of regional
studies as well as a few more recent pan-Arctic surveys.
It is far from an exhaustive account of the algal diversity of terrestrial, freshwater and marine habitats; more
inventory work is still needed to get a good picture of
the situation across the immense polar region. Here, the
Arctic pertains to the Arctic lands (CAVM Team 2003,
Walker et al. 2005), their bordering seas (Beaufort
Sea, Canadian Arctic Archipelago straits and channels,
Hudson Bay system, Melville Bay, Baffin Bay, Denmark
Strait, Greenland Sea, Barents Sea, Kara Sea, Laptev
Sea, East Siberian Sea, northernmost Bering Sea and
Chukchi Sea) and the Arctic Ocean (cf. Fig. 6.4 in Christiansen & Reist, Chapter 6).
9.4.2.1. Russia
Okolodkov (1992) studied the sympagic flora of 125
stations in the Laptev, East Siberian and Chukchi Seas.
Over 120 algal species were identified, predominantly
diatoms, primarily of the genera Navicula (24 species)
and Nitzschia (20). Allochtonous freshwater diatoms
and many marine planktonic Chaetoceros and Thalassiosira
species were recorded as well. Okolodkov (1998) also
presented a checklist of dinoflagellates recorded since
1878 in the Russian Arctic and the central Arctic Basin.
Apart from several incertae sedis (organisms of unknown
taxonomic placement), 189 species were recognized
belonging to 16 families and 34 genera. Peridinales species were most prominent, followed by Gymnodiniales
and Gonyaulacales. The genera Protoperidinium (c. 50
species), Peridinium (20), Dinophysis (20) and Gyrodinium
(18) were well represented. The diversity and distribution of marine benthic diatoms in the Laptev Sea were
studied by Cremer (1998), who recorded the occurrence
of 345 taxa in 56 genera, including 78 taxa mainly from
Arctic and sub-Arctic areas. Species-rich genera included
Navicula (72 species), Pinnularia (27), Nitzschia (21),
Cymbella (20), Eunotia (20), Fragilaria (20) and Achnanthes
(16). The taxonomic biodiversity of limnic phytoplankton
of the Pechora Delta and adjacent tundras was studied
by Stenina et al. (2000). They recorded the occurrence
of 440 species and 523 subspecific taxa (Tab. 6.2.7. in
Stenina et al. 2000). Diatoms showed the highest diversity with 360 taxa in 44 genera and 19 families, followed
by blue-green algae with 79 taxa in 26 genera and 18
families, and chlorophytes with 72 taxa in 34 genera and
21 families. Xanthophyta, Chrysophyta and Dinophyta
were less common. The diatom genera Navicula, Nitzschia,
Pinnularia, Fragilaria, Achnanthes and Cymbella appeared
very species-rich. A last report by Ratkova & Wassmann
(2005) listed the occurrence of 306 algal species in the
phytoplankton and sea-ice communities of the White Sea
and Russian Barents Sea, with 156 species common to
both environments. Most species observed in the sea ice
were similar to those recorded in other Arctic regions.
More recently, Poulin et al. (2011) listed the most
frequently recorded marine phytoplankton diatoms
(Chaetoceros contortus, Thalassiosira gravida, T. nordenskioeldii, Cylindrotheca closterium, Thalassionema nitzschioides)
and dinoflagellates (Protoperidinium brevipes, P. pellucidum) from the Russian Arctic seas, as well as the most
commonly recorded sea ice diatoms (Melosira arctica,
Cylindrotheca closterium, Entomoneis kjellmanii, Fragilariopsis
cylindrus, F. oceanica, Navicula directa, N. transitans var.
derasa, Nitzschia frigida, N. polaris, Pseudo-nitzschia delicatissima). Many of these sea ice diatoms are also well known
from, and commonly occurring during, the spring
blooms at the ice edge zone.
344
9.4.2.2. Svalbard
In Svalbard, Hansen & Jenneborg (1996) recorded the
occurrence of 163 species of benthic marine algae and
cyanobacteria, including 38 chlorophytes, 60 phaeophytes and 59 rhodophytes, whereas some 200 marine
microalgae were reported by Hasle & von Quillfeldt
(1996) and Okolodkov et al. (2000). Diatoms (108
species) and dinoflagellates (60) are the most diverse
groups, with Chaetoceros (21) and Thalassiosira (16) as
the most prominent diatom genera, and Protoperidium
(18) in the dinoflagellates. Werner et al. (2007) studied
sympagic algae in pack ice during winter and identified
54 taxa. Diatoms appeared prominent with at least 24
species. Conversely, Skulberg (1996) listed a total of 766
species of terrestrial and limnic algae and cyanobacteria, including a number of incertae sedis: more than half
the species (i.e. 409) are represented by diatoms with
the most speciose genera being Navicula (97 species),
Pinnularia (49), Cymbella (49) and Epithemia (25). Matuła
et al. (2007) identified 150 algal species from several terrestrial habitats in W Svalbard, including 100 blue-green
algae, with 55 species new to Svalbard.
In the European Arctic, macroalgal composition and
zonation patterns are most well known from Svalbard,
especially the Kongsfjorden area (Wulff et al. 2009).
A total of 80 seaweed species, mostly brown algae, are
known from various Svalbard fjords (Wulff et al. 2009).
The distribution of these polar seaweeds depends mainly
on exposure and depth. Kelp species, Laminaria digitata
and Saccharina latissima (syn. Laminaria saccharina), are
dominant at depths between 5 and 15 m. A similar macroalgal flora to the Svalbard archipelago extends eastwards along the Russian Arctic coast, but is less speciose
than in Svalbard.
9.4.2.3. Greenland
There are a fair number of old and detailed local and
regional floristic monographs on algae of Greenland,
such as those on marine macroalgae (Rosenvinge 1893,
1898, Lund 1951, 1959a, 1959b, Pedersen 1976), marine phytoplankton (Grøntved & Seidenfaden 1938), and
freshwater desmids (Grönblad 1952), diatoms (Foged
1953, 1955, 1973) and algae (Hansen 1967). A rather
poor knowledge of freshwater algal species diversity was
summarized by Kristiansen (2003). The most speciose
estimated algal groups were the diatoms (1,000 taxa),
the desmids with the Mesotaeniaceae and Desmidiaceae
(400) and the chlorophytes (200), followed by chrysophytes (83), dinoflagellates (21) and cyanobacteria (127),
with several confined to hot spring environments. The
total number of algal species was estimated at about
1,900. The species diversity of marine phytoplankton,
considered to be very fragmentary for Greenland, was
reviewed by Burmeister (2003) and Poulin et al. (2011).
Diatoms are the most speciose group of microalgae in
pelagic waters with some 250 species, mainly from the
following dominant genera: Chaetoceros, Nitzschia and
Thalassiosira, followed by prymnesiophytes (38 species),
Arctic Biodiversity Assessment
chrysophytes (15) and cryptophytes (10). An estimated
50 species of Pedinophyceae occur in the coastal waters
around Greenland. Benthic macroalgae are far better
known in Greenland with a long history of floristic
surveys. Eight classes of benthic algae occur along the
coasts of Greenland, with a total number of 215 species
consisting of 83 brown, 53 green and 52 red algae (Tab.
16 in Pedersen 2003).
Already in old times the zonation of macroalgae and
their main players along the Greenland rocky coast were
well known by the indigenous peoples, because these
informed the kayak hunters about the tidal conditions
(see Robbé 1994).
9.4.2.4. Canada
The Canadian Arctic has been surveyed for its freshwater
and marine algal flora, but unfortunately without entirely
covering this immense polar region. The first significant
algal records came when the eastern Canadian Arctic
was inventoried by Ross (1947), Seidenfaden (1947) and
Whelden (1947). Ross (1947) recorded a total of 245
freshwater diatom taxa, with the Naviculaceae being the
most speciose family (170 taxa) mainly represented by
the genera Navicula (47 taxa) and Pinnularia (43), while
the genus Nitzschia was represented by 16 taxa. Whelden
(1947) surveyed 423 algal taxa mainly from freshwater
habitats with only a few from the marine shoreline. The
main algal groups were the Chlorophyceae (284 taxa),
Cyanophyceae (112), Phaeophyceae (12) and Rhodophyceae (8). Among the green algae, the desmids (Desmidiaceae) showed the highest diversity with 220 taxa mainly
dominated by the genera Cosmarium (90 taxa) and Staurastrum (70), while Lyngbya, Gloeocapsa and Nostoc were the
dominant genera in the Cyanophyceae with 12, 10 and
nine species, respectively. Finally, marine phytoplankton
were inventoried by Seidenfaden (1947), who reported a
total of 128 taxa, including 84 diatoms, 39 dinoflagellates
and five small flagellates. The two most speciose phytoplankton genera were the diatom Chaetoceros (23 taxa) and
dinoflagellate Peridinium (19).
In the 1980s, Sheath & Steinman (1982) listed the freshwater algal flora of 279 bodies of waters from the Northwest Territories, which consisted of 1,577 taxa in 212
genera. The major algal classes included the Bacillariophyceae with 761 taxa, followed by Chlorophyceae (481)
and Cyanophyceae (173), which accounted for 90% of
the total freshwater flora diversity. The most speciose
genera were Navicula (119 taxa), Cymbella (78), Pinnularia
(73), Eunotia (55), Nitzschia (52), Achnanthes (45) and
Gomphonema (44) for the Bacillariophyceae; Cosmarium
(156 taxa), Staurastrum (56) and Closterium (35) for the
Chlorophyceae (Desmidiaceae); and Oscillatoria (18
taxa), Anabaena (13), Chroococcus (12), Lyngbya (12) and
Nostoc (12) for the Cyanophyceae. In 1980, Lee reported
the occurrence of 183 marine seaweed taxa from more
than 105 sites scattered throughout the Canadian Arctic,
including 37 new records. The most speciose genera
were the green alga Enteromorpha (8 taxa) and the brown
Chapter 9 • Plants
algae Laminaria (7) and Fucus (5). Later, Haber (1995)
reported a total of 171 marine benthic algae across the
Canadian Arctic, with Saccharina latissima (syn. Laminaria
saccharina) as the most common species.
Generally speaking, benthic algal diversity increases
with decreasing latitude and decreasing longitude from
the west to east along Parry Channel, the main marine
corridor linking the Beaufort Sea-Arctic Ocean to Baffin
Bay-North Atlantic Ocean. Hsiao (1983) recorded a total
of 685 marine phytoplankton and sea-ice microalgal
taxa across the Canadian Arctic, which mainly consisted
of 561 diatom taxa followed by 95 dinoflagellates, 22
chrysophytes, four chlorophytes and three euglenids.
For northern Baffin Bay in the eastern Canadian Arctic, Lovejoy et al. (2002) reported a total of 192 marine
phytoplankton taxa, mainly represented by 75 diatom
taxa, 58 dinoflagellates, 46 flagellates and 13 incertae
sedis. The most speciose genera were the diatom Chaetoceros (17 taxa) and the dinoflagellate Gymnodinium (12). In
the central Canadian Arctic near Resolute Bay, Riedel
et al. (2003) recorded 180 marine phytoplankton and
sea-ice algal taxa consisting of 99 diatoms, 45 dinoflagellates, 26 flagellates and 10 incertae sedis, with the most
speciose genera belonging to the diatoms Navicula (17
taxa), Nitzschia (14) and Chaetoceros (8), and the dinoflagellates Gymnodinium (8 taxa) and Gyrodinium (8). In the
northernmost part of the Canadian Arctic Archipelago,
Antoniades et al. (2008) described and illustrated 362
benthic diatom taxa from various freshwater habitats,
primarily consisting of these dominant genera: Pinnularia (37 taxa), Navicula (29), Nitzschia (25), Eunotia (15),
Gomphonema (14), Neidium (14), Caloneis (12), Cymbopleura
(12), Encyonema (12) and Stauroneis (12).
In a recent circumpolar biodiversity assessment of the
marine unicellular eukaryotes, Poulin et al. (2011) recorded a total of 1,350 phytoplankton and sea-ice algal
taxa for the entire Canadian Arctic, including the Hudson
Bay system (e.g. Hudson Bay, Hudson Strait, Foxe Basin).
They reported the most frequently occurring phytoplankton species, which mainly consisted of the diatoms
Attheya septentrionalis, Chaetoceros decipiens, C. furcillatus,
C. wighamii, Eucampia groenlandica, Thalassiosira gravida,
T. nordenskioeldii, Cylindrotheca closterium, Fragilariopsis
cylindrus, F. oceanica, Nitzschia frigida and Pseudo-nitzschia
seriata. They also reported the most commonly occurring
sea-ice diatom taxa, consisting of Attheya septentrionalis,
Melosira arctica, Cylindrotheca closterium, Entomoneis kjellmanii, Fragilariopsis cylindrus, Navicula directa, N. transitans,
Nitzschia frigida, N. longissima and Pauliella taeniata.
9.4.2.5. Alaska
Historical seaweed collections from Alaska were
reviewed by Lindstrom (2006). The total number of
recognized seaweed species for Alaska increased from
376 in 1977 to about 550 today. Marine unicellular
eukaryotes from the Alaskan Arctic were addressed by
Poulin et al. (2011) who reported a total of 443 phyto-
345
plankton and sea-ice microalgal taxa, mainly represented
by diatoms (331 taxa) and dinoflagellates (74). The low
number of freshwater and marine algae reported for the
Alaskan Arctic can be explained by the low number of
investigations conducted in these regions.
9.4.3. Pan-Arctic surveys
As a constant feature of the Arctic regions, snow offers
a suitable habitat for the development of freshwater
microbial communities (e.g. Gradinger & Nürnberg
1996, Müller et al. 1998, Takeuchi et al. 2001, Vincent et al. 2004, Larose et al. 2010). Such microbial
mats were recorded from ice floes in the Arctic Ocean
(Gradinger & Nürnberg 1996), snow fields in Svalbard
(Müller et al. 1998, Larose et al. 2010), Devon and
Penny ice cap glaciers in the Canadian Arctic (Takeuchi
et al. 2001) and on the ice shelf in the Canadian high
Arctic (Vincent et al. 2004). Algae on snow fields and
on the surface of ice floes are mainly represented by
the chlorophytes Chlamydomonas nivalis (green and red
forms) and Chloromonas nivalis (Gradinger & Nürnberg
1996, Müller et al. 1998), while Larose et al. (2010)
recorded 19 different bacterial classes from 16S rRNA
gene sequencing, mainly dominated by Betaproteobacteria and Sphingobacteria from snow fields in Svalbard.
Exposed microbial mats on Arctic ice shelves were
dominated by the chlorophyte genera Chlorosarcinopsis,
Pleurastrum, Palmellopsis and Bracteococcus, and the cyanobacteria Nostoc, Phormidium, Leptolyngbya and Gloeocapsa
(Vincent et al. 2004). Glaciers farther inland such as
the Devon and Penny ice caps on Devon and Baffin
Islands, respectively, exhibit a microbial community
characterized by seven chlorophyte and cyanophyte
taxa (Takeuchi et al. 2001).
Biodiversity assessments of algae across the Arctic are
extremely scarce with only a handful of reports on marine seaweeds and unicellular eukaryotes (e.g. Kjellman
1883, van den Hoek 1984, Lüning 1985, Wiencke et al.
2007, Wulff et al. 2009, Poulin et al. 2011). This type
of information is simply lacking for the freshwater and
terrestrial polar environments. Arctic species counts
are likely to be underestimated due to few collections,
extremely remote areas and sampling logistics.
According to Lüning (1985), there are approximately
150 seaweed species across the Arctic; the most recent
estimates total 210-215 (Pedersen 2003, Archambault
et al. 2010). The Arctic seaweed flora is of Atlantic and
by and large Pacific origin (Adey et al. 2008), with many
species having a circumpolar distribution and a few cosmopolitan or endemic species (Kjellman 1883, Lüning
1985, Wiencke et al. 2007, Wulff et al. 2009). About a
dozen seaweed species are restricted to the Arctic, including the brown algae Punctaria glacialis and Platysiphon
verticillatus, and the red alga Petrocelis polygyna (Wiencke
et al. 2007). Most species have a southern range extension into the temperate region, such as the kelp Laminaria solidungula and the red algae Devaleraea ramentacea,
Turnerella pennyi, Neodilsea integra (now Dilsea socialis)
346
Arctic Biodiversity Assessment
Total
Russia
Canada
Alaska
Algal group
Scandinavia
Table 9.9. Survey of total numbers of marine unicellular eukaryote
taxa in Arctic regions (Poulin et al. 2011).
Archaeplastida/Plantae
Chlorophyta
12
34
3
17
55
Prasinophyta
1
42
25
18
60
Chromalveolata
Centric diatoms
99
199
132
202
297
Pennate diatoms
232
604
251
563
930
Bicosoecida
0
7
5
1
10
Chrysophyceae
9
22
18
8
38
Dictyochophyceae
3
14
9
9
19
Pelagophyceae
1
0
0
0
1
Rhaphidophyceae
0
2
1
2
3
Synurales
0
3
3
0
6
Xanthophyceae
1
3
0
0
3
Cryptophyceae
0
23
10
9
30
Prymnesiophyceae
2
33
45
10
70
74
266
183
257
441
Euglenida
3
14
4
10
20
Kinetoplastea
1
8
1
0
9
0
30
39
9
46
0
4
0
9
12
Dinolagellates
Excavata
Ophistokonta
Choanolagellates
Cyanophyta
Incertae sedis
Total species number
5
42
25
4
56
443
1,350
754
1,128
2,106
and Pantoneura baerii (Lüning 1985). At a pan-Arctic
scale, macroalgal species richness tends to decrease with
increasing latitude and from the Atlantic sector to the
Pacific sector (Wiencke et al. 2007). Arctic seaweeds are
almost entirely subtidal with, however, some specialized
species exclusively in the supralittoral or spray zone such
as the green alga Prasiola crispa and the red alga Bangia
atropurpurea (Wiencke et al. 2007).
The pan-Arctic diversity of marine pelagic and sea-ice
unicellular eukaryotes was thoroughly reviewed by
Poulin et al. (2011) based on current inventories and literature. A total of 2,106 marine single-celled eukaryote
taxa were reported from the four main Arctic regions,
namely Alaska, Canada, Scandinavia and Greenland, and
the Russian Federation (Tab. 9.9): 1,027 sympagic taxa
associated with sea-ice and 1,874 phytoplankton taxa.
More than three-quarters of the total microalgal flora
belongs to diatoms (1,227 taxa; 58%) and dinoflagellates
(441; 21%), with pennate and centric diatoms accounting for 44% (930 taxa) and 14% (297), respectively.
Landfast and pack ice are predominantly colonized by
pennate diatoms. Some colonial diatoms Entomoneis
kjellmannii, Fragilariopsis cylindrus, F. oceanica, Nitzschia
frigida, Pauliella taeniata, and solitary pennate diatoms
Cylindrotheca closterium, Navicula directa, together with the
colonial centric Melosira arctica and solitary, epiphytic Attheya septentrionalis are considered highly associated with
or strictly confined to the Arctic sea ice, whereas Navicula frigida can be regarded as the sentinel endemic species
of sympagic communities (Różańska et al. 2009, Poulin
et al. 2011). Some colonial centric diatoms such as Chaetoceros furcillatus, Thalassiosira gravida, T. nordenskioeldii and
pennate Fragilariopsis oceanica together with the solitary
pennate diatom Cylindrotheca closterium are marine cold
water plankton and widespread across the Arctic seas.
The biodiversity of the smaller cell-sized phytoplankton
(< 20 µm) is not very well known and is estimated at
20% of the diversity of the known pan-Arctic unicellular marine pelagic and sea-ice eukaryotes (Poulin et
al. 2011). Nanoalgae mainly consist of cyanobacteria,
prasinophytes, dinoflagellates, diatoms and prymnesiophytes (Tab. 6.1 in Thomas et al. 2008). In addition to
this first biodiversity assessment of the marine unicellular eukaryotes, there are 37 potentially toxic species
recorded including 25 dinoflagellates, nine diatoms, two
prymnesiophytes and one raphidophyte (Tab. 4 in Poulin
et al. 2011).
9.4.4. Trends
It is extremely hazardous to provide estimates of trends
in Arctic phytoplankton, sea ice and benthic seaweed assemblages due to insufficient and fragmented knowledge
of this algal biodiversity. However, recent studies have
provided further information about the various marine
algal groups present across the Arctic (Wiencke et al.
2007, Li et al. 2009, Różańska et al. 2009). For instance,
it has been reported from the Arctic Ocean that the size
class of marine phytoplanton is changing from large to
small cells (Li et al. 2009). It has now to be seen if this
trend will effectively apply to the entire Arctic region.
Therefore, indeed more inventories and monitoring
of micro- and macroalgae are needed to better define
populations and assess trends across the Arctic.
9.4.5. Conclusions and recommendations
The total number of recognized algal species for the
Arctic is at present likely around 4,000, which represents 10% of the world’s recognized species. There are
between 30,000 and 40,000 described species of algae
worldwide, which correspond to only a small fraction of
the estimated number of about 200,000 species (Poulin
& Williams 2002). The total species number of algae
and cyanobacteria in the Arctic is still largely unknown,
especially in terrestrial and freshwater environments.
Regarding their huge ecological importance for all life
on Earth, both in the sea and on land, better inventories
and monitoring of algae are strongly needed, particularly considering that the Arctic regions are and will be
severely impacted by global warming.
Chapter 9 • Plants
“The urgent need for more studies of all aspects of
the biodiversity and ecology of polar algae has never
been more apparent than at present” (Wulff et al.
2009).
“Such an initiative will require that the field of taxonomy be better financially supported by the panArctic countries … it would also be imperative to
develop some training of the next generation of expert scientists in the field of phytoplankton taxonomy
and systematics, which has been entirely neglected at
present” (Poulin et al. 2011).
A major effort should be undertaken to establish a
complete baseline of the biodiversity of marine and
freshwater phytoplankton and macroalgae and polar sea
ice microalgae, especially since these algae will become
part of the CAFF Circumpolar Biodiversity Monitoring
Program (CBMP). Reaching that goal requires more
taxonomic studies in order to elucidate the species
concept and harmonize it across the Arctic. The fields
of taxonomy and systematics should be considered more
than a descriptive exercise and rather as fundamental
tools of discovery, conservation and management. Future efforts should focus particularly on the biodiversity
of small-celled (< 20 µm) microalgae. Finally, all this
research effort should be undertaken through international networks leveraging the costs associated with such
pan-Arctic programs.
ACKNOWLEDGEMENTS
We thank Heidi Solstadt and Reidar Elven (Oslo) for
providing the then unpublished PAF report in 2009.
Thanks are due to Robbert Gradstein (Paris) and Heinjo
During (Utrecht) for their advice regarding the bryophytes. Paul Sokoloff (Ottawa) provided literature.
We thank Irina Goldberg (Copenhagen) for providing
information about the mosses of Greenland and Kell
Damsholt (Copenhagen) for permission to publish his
checklist of the liverworts of Greenland (Appendix 9.7).
Sandra Lindstrom, Vancouver, gave advice and support.
We thank Kenneth Høegh for information about Lupinus
nootkatensis in SW Greenland. Finally thanks are due to
anonymous reviewers for their useful comments on the
manuscript.
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Appendix 9: www.abds.is/aba-2013-appendix-9
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Arctic Biodiversity Assessment
The golden colored blackening waxcap Hygrocybe conica var. aurantiolutea is a colorful member of waxcaps that grows in grasslands
in the low and sub-Arctic zones. At appropriate climatic conditions, the cryptically growing long-lived mycelia produce sporocarps in
August-September. Waxcaps are sensitive to nitrogen, and their occurrence is strongly reduced in temperate and boreal zones due to
anthropogenic deposition of nitrogen and fertilization. Tasiusaq at Qassiarsuk in South Greenland, 1987. Photo: Flemming Rune.