ArcticBiodiversity

AssessmentStatus and trends in Arctic biodiversity

ARCTIC COUNCIL

i

ARCTIC COUNCIL

Lead authors:

Tom BarryDominique Berteaux

Helga BültmannJørgen S. Christiansen

Joseph A. CookAnders Dahlberg

Fred J.A. DaniëlsDorothee Ehrich

Jon FjeldsåFinnur FriðrikssonBarbara GanterAnthony J. GastonLynn J. GillespieLenore GrenobleEric P. HobergIan D. HodkinsonHenry P. HuntingtonRolf A. ImsAlf B. JosefsonSusan J. KutzSergius L. KuzminKristin L. LaidreDennis R. Lassuy

Patrick N. LewisConnie Lovejoy

Hans MeltofteChristine Michel

Vadim MokievskyTero Mustonen

David C. PayerMichel Poulin

Donald G. ReidJames D. Reist

David F. TesslerFrederick J. Wrona

Chief scientist and executive editor:Hans Meltofte

Assistant editors on species chapters:Alf B. Josefson and David Payer

Arctic Biodiversity AssessmentStatus and trends in Arctic biodiversity

ARCTIC COUNCIL

ii

Arctic Biodiversity AssessmentStatus 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.orMeltofte, 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 reflect 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 financial 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: caff@caff.is

Note: This report may contain images for which permission for use will need to be obtained from original copyright holders.

Print: Narayana Press, DenmarkImpression: 1,300ISBN: 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.

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Preface by CAFF and Steering Committee Chairmen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v

Foreword by the Chief Scientist . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

Indigenous peoples and biodiversity in the Arctic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

Synthesis: Implications for Conservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

Species Diversity in the Arctic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66

Mammals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78

Birds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142

Amphibians and Reptiles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182

Fishes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 192

Terrestrial and Freshwater Invertebrates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 246

Marine Invertebrates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 276

Plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 310

Fungi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 354

Microorganisms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 372

Terrestrial Ecosystems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 384

Freshwater Ecosystems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 442

Marine Ecosystems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 486

Parasites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 528

Invasive Species: Human-Induced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 558

Genetics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 566

Provisioning and Cultural Services . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 592

Disturbance, Feedbacks and Conservations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 628

Linguistic Diversity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 652

Lead author biographies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 664

Author affiliations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 670

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Contents

182 Arctic Biodiversity Assessment

A few amphibians and reptiles extend their distribution into the Arctic, such as this moor frog Rana arvalis of Eurasia. Photo: Konstantin Mikhailov.

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Chapter 5

Amphibians and ReptilesAuthorsSergius L. Kuzmin and David F. Tessler

ContentsSummary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184

5.1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184

5.2. Status of knowledge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1845.2.1. Historical overview . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184

5.3. Status and trends . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1855.3.1. Species richness and distribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . 185

5.3.1.1. Eastern hemisphere species . . . . . . . . . . . . . . . . . . . . . . . . . 1855.3.1.2. Western hemisphere species . . . . . . . . . . . . . . . . . . . . . . . . . 186

5.3.2. Status . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1875.3.3. Trends . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1875.3.4. Prospects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187

5.4. Conclusions and recom mendations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1895.4.1. Sensitive areas and hotspots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1895.4.2. Key knowledge gaps . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1895.4.3. Recommended conservation actions . . . . . . . . . . . . . . . . . . . . . . . . 189

5.4.3.1. Research recommendations . . . . . . . . . . . . . . . . . . . . . . . . . 1895.4.3.2. Conservation action recommendations . . . . . . . . . . . . . . 189

Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189

References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189

» We have lizards. We have seen them this year even at the upper reaches of ‘Afanas’ki’.

Leader of the Saami indigenous obschina ‘Piras’ in the Kola Peninsula, Andrey Yulin on northwards expansion of lizards; Zavalko & Mustonen (2004).

184 Arctic Biodiversity Assessment

SUMMARY

The herpetofauna of the Arctic is depauperate relative to temperate and tropical regions. Only five amphibians and a single reptile range into the Arctic, and none are circumpolar. All Arctic amphibian and reptile taxa are currently categorized as ‘Least Concern’ according to IUCN criteria. However, basic survey and inventory data for these species are lacking across most of the Arctic, and there are few quantitative data on abundance, status or trends for Arctic herpetofauna. At the same time, isolated populations of amphibians and reptiles in the Arctic exist at or near their current physiological limits and likely face a number of escalating challenges stem-ming primarily from habitat alteration.

5.1. INTRODUCTIONAlthough amphibians and reptiles account for nearly 15,000 species worldwide, only five amphibians and a single reptile are found in the Arctic. The majority of Arctic herpetofauna are found in the eastern hemi-sphere; there are no circumpolar taxa (Tab. 5.1). Am-phibian species richness (number of species) in the Arctic is as low as in desert regions. Amphibians and reptiles are phylogenetically the oldest of terrestrial vertebrates, and their limited representation in the Arctic is due in large measure to their poikilothermic physiology (body temperature determined by ambient conditions).

A number of recent publications have suggested ma-jor changes to herpetological systematics (Frost et al. 2006, Roelants et al. 2007), but because these proposed changes are not yet universally accepted and many names remain in a state of flux, we follow stable herpetological taxonomy as described in Collins & Taggart (2002) and Kuzmin & Semenov (2006).

Eastern hemisphere taxa: Siberian newt Salamandrella keyserlingii Common frog Rana temporaria Moor frog Rana arvalis Siberian wood frog Rana amurensis Common lizard Lacerta viviparaWestern hemisphere taxa: Wood frog Rana sylvatica

5.2. STATUS OF KNOWLEDGE

The first scientific records of Arctic amphibians are from expeditions in the 19th and early 20th centuries. Since then, despite intensive study of Arctic regions, little attention has been paid to understanding the distribu-tion and ecology of Arctic amphibians and reptiles. Few works on the distribution, morphology, genetics, phenology, development, hibernation and diet of Arctic amphibians have been published.

Amphibians and reptiles reach Arctic regions only on the periphery of their ranges, where their overall abundance is low. Due to their limited dispersal capabilities and because their distributions are defined by fine-scale microhabitat associations, their actual distributions may be quite patchy (Olson 2009). Because focused research on amphibian and reptile biology in the Arctic is scarce, much of the data available were obtained as a by-product of other efforts.

5.2.1. Historical overviewHistorical information on the Arctic distributions of amphibians and reptiles is very limited. To date, there have been only two published studies on the historical phylogeography of Arctic herpetofauna using molecular techniques.

Poyarkov & Kuzmin (2008) investigated molecular genetics in the Siberian newt throughout its range and found that patterns of genetic differentiation of Siberian populations are likely the result of repeated processes of colonization of new territories during inter-glacial ep-ochs and subsequent retreats into more temperate belts during glacial peaks. Following the peak of the most recent glacial maximum, the Siberian newt likely first colonized territories in E Siberia, followed by coloniza-tion events west towards the Urals and east towards Beringia and Kamchatka. Dispersal to the north and west appears to have taken place very quickly.

In North America, an investigation of the historic phy-logeography of the wood frog using mitochondrial genes suggests a post glacial range expansion that differs from most other herpetofaunal taxa on the continent (Lee-Yaw et al. 2008). The wood frog appears to have radiated

Table 5.1. Amphibian and reptile taxa of the Arctic. ‘Redlisted’ denotes taxa on the IUCN Red List of Threatened Species. * denotes known trends over the past 10-20 years.

Region Orders Families Genera Species Sub species Stable* Increasing* Decreasing* No Info.* Redlisted

Arctic 2 3 3 5 – ? – – 5 0

WH Arctic 1 1 1 1 – ? – – 1 0

WH high Arctic – – – – – ? – – – 0

WH low Arctic 1 1 1 1 – ? – – 1 0

EH Arctic 2 3 3 4 – ? – – 4 0

EH high Arctic – – – – – ? – – – 0

EH low Arctic 2 3 3 4 – ? – – 4 0

Chapter 5 • Amphibians and Reptiles 185

from a number of high-latitude refugia in what is now the northeastern United States, while most amphibians currently found in western and northwestern North America appear to have radiated from lower-latitude ref-ugia to the west. Colonization following the last glacial maximum appears to have been rapid, with expansion to the north and northwest (Arctic Alaska and most of sub-Arctic Canada) having occurred from a single putative refugium near the edge of the Laurentide ice-sheet in present-day Wisconsin, while radiation into sub-Arctic Quebec, Newfoundland and Labrador took place from another refugium near the Appalachian Mountains in the vicinity of Pennsylvania.

5.3. STATUS AND TRENDS

5.3.1. Species richness and distributionMaximum Arctic amphibian and reptile richness, three species, occurs south of the Yamal Peninsula, where the European common frog and moor frog coexist with the Siberian newt. The moor frog and common frog are sympatric on the Kanin Peninsula and in the Vor-kuta region, while the Siberian newt and Siberian wood frog both inhabit the Khaiyr Settlement in Republic of Sakha-Yakutia. Throughout the remainder of the Arctic,

species richness of amphibians and reptiles appears to be zero or one.

5.3.1.1. Eastern hemisphere species

The Siberian newt is the most widespread amphibian species in the Arctic and sub-Arctic (Fig. 5.1), and has the widest geographical range of any recent amphibian species, c. 12 million km2. The northernmost habitats of the Siberian newt consist of grass-undershrub-lichen-moss bogs and low shrub-moss and grass-moss tundras. The newt penetrates the Arctic in the area of the polar Urals and eastwards, and is found along the Khatanga River on the Taimyr Peninsula just south of the low Arctic (Kuzmin 1994). It reaches the Arctic Ocean in some areas of the Republic of Sakha-Yakutia, in par-ticular, near Chukochya Guba in Nizhnekolymskii. The Amguema River in the Shmidtovskii District of Chukot-ka marks the northeastern extent of its distribution.

Frogs Rana spp. are more narrowly distributed in the Arctic than the Siberian newt. The common frog and the moor frog are mainly inhabitants of Europe but both range eastwards to the Urals, with the moor frog found into E Siberia. The common frog crosses into the low Arctic only in the northernmost peninsulas of Norway (Gasc et al.1997) and along the eastern slope of the polar

Figure 5.1. Known locations of Eastern Hemisphere Arctic amphibian and reptile species. Colored dots indicate known locations: Red dots represent Siberian Newt; Blue connotes the common frog; yellow signifies Siberian wood frog; Pink indicates moor frog; and brown signifies common lizard. The graphic only includes data from the vicinity of the Arctic and sub-Arctic and is not indicative of the global distribution of these species. Data from Amphibians of the former USSR, Databank ® 0229803415 and Gasc et al. 1997.

186 Arctic Biodiversity Assessment

Urals at the southern border of the Yamal Peninsula, in the Priuralskii District of Yamal-Nenetskii Autonomous Okrug, Tyumenskaya Province, Russia (Toporkova & Shvarts 1960, Toporkova & Zubareva 1965, Toporkova 1973, Ishchenko 1978). The common frog has been known to occur on the Kanin Peninsula since at least 1902 (Zhitkov 1905).

The moor frog has a broader Arctic distribution than the common frog. It occurs in southern Yamal and the Polar Urals, and as far north as the Khadyta-Yakha River (Shvarts 1959). A few records indicate it is found on the sub-Arctic/low Arctic margin to the east as far as the Malaya Khadyta River in Yamal (Anufriev 1984).

The Siberian wood frog is a widespread brown frog, whose distribution covers a large part of Siberia, far eastern Russia, and northern Mongolia and Manchuria. Nevertheless, this species is known from only one Arctic locality: Khaiyr Settlement in Ust-Yanskii District of the Republic of Sakha-Yakutia (Borkin et al. 1984).

The common lizard is the only reptile species found in the Arctic. It ranges from northwestern Spain through Europe and Siberia to Mongolia and eastwards to Sakha-lin Island in the Pacific. It reaches the Arctic in Europe in the vicinity of the Kanin Peninsula and around Vor-kuta City in the Komi Republic in Russia (Anufriev & Bobretsov 1996). The common lizard is found in the Po-lar Urals, southwards from the Yamal Peninsula, where it occurs primarily in river valleys (Vershinin 2007). In the area of Taimyr Peninsula the common lizard occurs in the sub-Arctic near the southern boundary of the low Arctic (Bannikov et al. 1977).

5.3.1.2. Western hemisphere species

The wood frog is the only herpetofaunal species found in the Arctic of the Western hemisphere. This widely distributed North American frog is found throughout the sub-Arctic (in Labrador, Quebec, Ontario, Manitoba, Saskatchewan, Alberta). It extends into the low Arctic in Alaska, and is suspected to do so in portions of Yukon,

Figure 5.2. Known locations of Western Hemisphere Arctic amphibians. Wood frog locations are represented by brown dots. Boreal chorus frog locations are represented by yellow dots. The graphic only includes data from the vicinity of the Arctic and sub-Arctic and is not indica-tive of the global distribution of these species. Data from Martof 1970, IUCN, Conservation International and NatureServe, 2004, Alaska Natu-ral Heritage Program, 2007, Gotthardt & Pyare 2009, Environment and Natural Resources Office, Government of the Northwest Territories 2010, Frogwatch Database, National Wildlife Research Centre, Environment Canada 2010.

Chapter 5 • Amphibians and Reptiles 187

Northwestern Territories, and Nunavut (DeGraaf & Rudis 1983, Cook 1984, Russell & Bauer 1993, Weller & Green 1997, Chubbs & Phillips 1998, Blackburn et al. 2001, Trust & Tangermann 2002, Carstensen et al. 2003, MacDonald 2003, Anderson 2004, Desroches & Rodrigue 2004, Slough and Mennell 2006, MacDonald & Cook 2007, Lee-Yaw et al. 2008, NatureServe 2010). This unique frog ranges at least as far north as the south side of the Brooks Range in Alaska, the Old Crow Flats in the Yukon, and the Mackenzie River Delta in Northwest Territories (Fig. 5.2). Despite its wide distribution and apparent ubiquity, there are only eight confirmed records of wood frogs extending into the Arctic as defined by Geographic Information System data layers developed for the Arctic Biodiversity Assessment (MacDonald 2003, Gotthardt & Pyare 2009, Environment and Natural Resources, GNWT 2010). The species is probably more widespread in the Arctic, and the limited number of records is likely due in part to very limited survey effort.

5.3.2. StatusAll of the amphibians and reptiles found in the Arctic are considered taxa of ‘Least Concern’ by IUCN, suggest-ing their populations are currently stable. However, as a class, amphibians are among the most globally threatened groups, with roughly 32% (1,856 species) considered threatened and 43% (2,469 species) declining. At least 168 species worldwide have become extinct over the last 20 years, and the numbers of threatened and extinct taxa are expected to continue to climb (Stuart et al. 2004).

While there are no indications of declines of amphibian and reptile populations in the Arctic, their apparent sta-bility may be a result of the almost complete lack of his-toric and contemporary abundance data. In a few cases there are some qualitative estimates of local abundance, but these are insufficient to determine the dynamics of even local populations. Overall numbers of amphibians and reptiles in the Arctic appear to be low and popula-tions appear scattered and isolated.

5.3.3. TrendsGlobal population estimates and trend data are largely lacking for all herpetofauna found in the Arctic, and few if any data exist on local Arctic population sizes or trends for any of the taxa.

The Siberian newt has been known as an Arctic spe-cies since 1909 (Nikolsky 1918), and is considered to be common and stable throughout its range, with the ex-ception of some small, declining and isolated populations in Mongolia, which are considered threatened (Kuzmin et al. 2008a).

The moor frog, though extinct in Switzerland, is con-sidered common, stable and a species of ‘Least con-cern’ by the IUCN throughout its distribution (Kuzmin et al. 2008b).

The common frog is considered common and stable throughout its range, though local declines have been noted in Switzerland and Spain (Kuzmin et al. 2008c), and since the 1970s in Norway along the northern limits of its distribution (Gasc et al. 1997).

Globally, the Siberian wood frog is considered com-mon, stable and a species of least concern (Kuzmin et al. 2008d), though there is no information on Arctic abundance or trends.

Population size, trend and status data for the common lizard are lacking throughout its large distribution, in-cluding the Arctic, but it is classified as ‘Least Concern’ by the IUCN (2010).

The wood frog is the most widespread amphibian in North America (Martof 1970), the most common am-phibian in Alaska (MacDonald 2003), and is considered relatively common throughout most of its range, includ-ing the sub-Arctic. There are no data on abundance, status or trends for this species in the Arctic. The global population trend is unknown but is suspected to be stable, and the wood frog is considered a species of least conservation concern (Hammerson 2004).

5.3.4. ProspectsIn general, the outlook for amphibian persistence in the Arctic is probably good. The species all appear secure globally, with Arctic populations representing the north-ern fringe of much larger distributions to the south. However, conservation of Arctic reptiles and amphibians will face a number of challenges in the immediate future and over the next century, including contaminants from local and global sources, anthropogenic habitat altera-tion, emerging infectious diseases, climate change-in-duced habitat alteration and loss, and the introduction of novel pathogens and predators.

Environmental contaminants originating either from lo-cal point sources or from diffuse regional and/or global sources may impact many otherwise undisturbed Arctic wetlands. Ackerman et al. (2008) and Landers et al. (2008) found concentrations of atmospherically depos-ited organic and other contaminants in fish from remote lakes in Arctic and sub-Arctic Alaska that exceeded thresholds of health concern for humans and wildlife. Because the study sites were all located in remote areas with no local contaminant sources, their presence was attributed to long-range trans-Pacific transport and to global sources. The risk to amphibians is unclear and undocumented, but due to their aquatic developmental phase, moist and highly permeable integuments, larval diets of zooplankton, phytoplankton and periphyton and adult diets of higher trophic level invertebrates, they may be predisposed to bio-accumulate a variety of contami-nants when present in the environment.

188 Arctic Biodiversity Assessment

As development inevitably proceeds in the Arctic, many wetland habitats will be transformed, and even those that remain intact may be exposed to additional threats as roads and other infrastructure elements are constructed. Research on wood frog breeding ponds at two National Wildlife Refuges in sub-Arctic Alaska documented some of the highest rates of skeletal abnor-malities found in amphibians anywhere; as high as 20% of individuals from some breeding sites had pronounced skeletal abnormalities (Reeves et al. 2008). The normal background rate for such abnormalities is estimated as either 0-2% (Ouellet 2000) or 0-5% (Johnson et al. 2001). The prevalence of structural abnormalities in Alaska was found to increase with proximity to roads, and more recently, the presence of contaminants (both organic and inorganic) and odonate larvae were identi-fied as predictors of the frequency of skeletal abnormali-ties at these sites (Reeves et al. 2010).

As climatic conditions in the Arctic continue to amelio-rate, we might expect the amphibians already present to expand their current distributions and colonize previ-ously unoccupied areas. However, a warming climate will also catalyze a suite of interconnected changes in the physical and biological environments of the Arctic, the ramifications of which are poorly understood for Arctic amphibians. Warmer winter and summer tem-peratures, alteration in both the rates and the timing of snow deposition and melt-off, and permafrost thaw will have profound effects on hydrology and hydroperiod across the Arctic. Both the total volume of water and the timing of its availability to plants and animals may be significantly altered (McDonald et al. 2004, Borner et al. 2008; see Wrona & Reist, Chapter 13) resulting in shifts in the composition of the biotic community. Permafrost thaw, which is forecast to continue at an accelerated rate, is already causing the draining of Arctic wetlands in Alaska (Yoshikawa & Hinzman 2003). In regions of thawing discontinuous permafrost in Siberia, the num-ber of large lakes declined by 11% and overall wetland surface area declined by 6% since the 1970s (Smith et al. 2005). These changes in hydroperiod and hydrology, and their attendant impacts on the biotic community, have the potential to disrupt many facets of amphibian ecology, including: (1) breeding phenology relative to the timing and availability of algae and the emergence of important aquatic invertebrate prey (and predator) spe-cies, (2) habitat connectivity, (3) dispersal movements, (4) juvenile and adult survival, and ultimately (5) the persistence of local populations.

Emerging infectious diseases are also likely to play a greater role in the ecology of Arctic amphibians. Warm-ing climatic conditions may aid the spread and survival of deadly new pathogens, and infrastructure development may provide new vectors for transmission across the land-scape. The amphibian chytrid fungus, Batrachochytrium dendrobatidis (Bd) is a recently recognized pathogen widely believed to be expanding due to climate change and im-plicated in a number of amphibian declines and extirpa-tions (Berger et al. 1998, Pounds et al. 2006). A number

of ranaviruses (genus Iridoviridae) are also associated with recent amphibian declines (Stuart et al. 2004). There has been little surveillance for Bd in the Arctic to date, but the fungus has already been detected in sub-Arctic Alas-ka, and both Bd and ranaviruses have been detected in Canada’s Northwest Territories (Reeves & Green 2006, Schock et al. 2009, T. Chestnut pers. com.). Infrastruc-ture development and increased human presence in the Arctic may contribute to the spread of these pathogens: Both ranaviruses and Bd may be translocated between water bodies via transport of contaminated water or sediments by humans (on footwear and equipment), birds (e.g. on feathers), or other animals (Johnson & Speare 2005, Harp & Petranka 2006, Phillot et al. 2010).

Warmer temperatures and expanded human industry and infrastructure will increase the risks of introducing exotic or novel species into Arctic environments. Arctic amphibians breed primarily in fish-free water bodies, in-cluding ephemeral pools and small lakes and ponds, and it is widely believed that this preference is an adaptation to avoid predation. Fish are successful predators on the eggs and aquatic larvae of amphibians (Semlitch 1988), and many amphibians have few defenses against fish (Grubb 1972). Introductions of non-native fishes have been implicated as one of the major causes of amphib-ian population declines (Kats & Ferrer 2003), and even native fish introduced into previously fish free habitats have the potential to extirpate local amphibian popula-tions. Increased human presence may serve as a vector for the spread of native and non-native fish across naïve Arctic wetlands. Amphibian larvae are also preyed on by a variety of birds, other amphibians (Sours & Petranka 2007) and a number of aquatic invertebrates, especially odonates, whose abundance appears related to water temperature (Reeves et al. 2010). Trophic webs and competitive interactions among amphibians in Arctic wetlands aren’t well understood, but changes in spe-cies composition or density can strongly influence the outcomes of competition and predation in these aquatic communities (Relyea 2000, Sours & Petranka 2007). The consequences for Arctic amphibian populations of any new or heightened predatory or competitive interac-tions are completely unknown.

While most sub-Arctic amphibians are unlikely to colonize the Arctic in the next century (due to their southerly distributions, inherently slow rate of dispersal and less freeze-tolerant physiologies) the boreal cho-rus frog Pseudacris maculata in the Western hemisphere may already have reached it (Fig. 5.2). In 2009 a single un-vouchered observation was recorded in the Arctic in Canada’s Northwest Territories near the border of Nuna-vut in the vicinity of Big Lake (Environment and Natural Resources, GNWT 2010). Very low mitochondrial genetic diversity found in populations occupying regions north of the last glacial maximum suggests the boreal chorus frog colonized its northern frontiers recently and rather rapidly (Lemmon et al. 2007). If it is not already, the boreal chorus frog will soon be the second Nearctic amphibian species.

Chapter 5 • Amphibians and Reptiles 189

5.4. CONCLUSIONS AND RECOM MENDATIONS

5.4.1. Sensitive areas and hotspots

Hotspots are difficult to identify because the distribu-tions of Arctic amphibians and reptiles are so poorly characterized. Furthermore, these species are found in small, isolated and patchily distributed populations for a variety of reasons: (1) their Arctic range limits likely represent each species’ physiological limitations, (2) amphibians require a number of different micro-habitat features to intersect in close proximity due to their limited movement potential, and (3) they may not occupy all suitable habitats within their apparent range due to the susceptibility of small populations to stochas-tic events and the residual effects of past disturbances (Olson 2009). Nonetheless, a few areas in the Eastern hemisphere appear to be of particular importance: the corridors and deltas of the Khadyta-Yakha River on the Yamal Peninsula, the Chaunskaya Tundra in the lowlands of the Chaunsky Administrative District of the Chu-kotsky Autonomous Okrug, and the Khalerchinskaya Tundra in the Kolyma lowlands.

5.4.2. Key knowledge gapsThe principal knowledge gap is the near complete lack of survey and inventory data for status and population trends of Arctic amphibians and reptiles. Distributions are poorly and incompletely characterized, and are known only in broad general terms.

There are no reliable abundance estimates for local or regional populations for any Arctic herpetofauna, and there are no statistically meaningful monitoring efforts currently in place. General lack of understanding of the factors which limit amphibian and reptile populations in the Arctic is also a principal knowledge gap.

5.4.3. Recommended conservation actions

5.4.3.1. Research recommendations

• Establish effective survey and inventory efforts to bet-ter define the actual distributions and ecology of these species.

• Construct statistically defensible baselines of abun-dance data in specific locations against which changes in abundance can be monitored.

• Establish monitoring programs with replicate schema representative of the range of habitats and microhabi-tats inhabited by each species. Monitoring locations should also be chosen in such a way so as to minimize the effort and expense to reach them in order to in-crease the likelihood that monitoring will be contin-ued into the future. If practicable, monitoring efforts should be collocated with monitoring efforts for other taxa in order to develop economies of scale for all

monitoring, and to improve our understanding of the dynamics of Arctic ecosystems.

• Conduct research into the impacts of climate-induced changes to hydrology/hydroperiod on reproduction, persistence and habitat connectivity for Arctic am-phibians.

• Determine the geographic prevalence of contaminant burdens and chief pathogens for amphibians across the Arctic.

These efforts may involve citizen science projects.

5.4.3.2. Conservation action recommendations

• Develop guidelines for human development projects that require land managers and developers to consider amphibian and reptile habitats and populations in their development plans.

• Determine which areas are of special importance for amphibian and reptile species richness and for the long-term persistence of individual taxa. Use data from survey and inventory efforts to identify hotspots and areas of likely significance by modeling species’ habitat and micro-habitat associations across the Arc-tic landscape.

• Establish or strengthen protections for areas of key importance to reptiles and amphibians. Arctic am-phibians have complex life cycles, and require a range of habitats throughout their annual cycles and life histories. Conservation of these species will require a landscape-level approach, conserving various vital habitats at appropriate spatial scales and maintaining connectivity between conservation units, while ac-counting for expected wetland loss and alteration.

ACKNOWLEDGEMENTSWe wish to thank M. Reeves, B. Slough and D. Payer for their thoughtful reviews of this manuscript, O.L. Leontyeva and S. Aikens for their unpublished data on Arctic frogs, and T. Chestnut for unpublished data on the prevalence of Batrachochytrium dendrobatidis in Alaska.

REFERENCESAckerman, L.K., Schwindt, A.R., Simonich, S.L., Koch, D.C.,

Blett, T.F., Schreck, C.B. et al. 2008. Atmospherically de-posited PBDEs, pesticides, PCBs, and PAHs in western U.S. National Park fish: concentrations and consumption guidelines. Environ Sci. Technol. 42: 2334-2341.

Alaska Natural Heritage Program, University of Alaska Anchorage (NatureServe) 2007. Wood frog, Alaska Status Report. aknhp.uaa.alaska.edu/zoology/Zoology_Amphibs_track07.htm. [ac-cessed 1 March 2010]

Amphibians of the former USSR, Databank ® 0229803415, State Register of Databanks of Russian Federation.

Anderson, B.C. 2004. An opportunistic amphibian inventory in Alaska’s national parks 2001-2003. National Park Service, Inventory and Monitoring Program, Anchorage.

Anufriev, V.M. 1984. Number and distribution of Rana arvalis in the valley of the Malaya Khadyta River, Southern Yamal. – In:

190 Arctic Biodiversity Assessment

V.G. Ishchenko (ed.). Vid i Ego Produktivnost v Areale, pt. 5: Voprosy Gerpetologii. Sverdlovsk: Inst. Plant and Anim. Ecol. Ural Branch of USSR Acad. Sci.

Anufriev, V.M. & Bobretsov, A.B. 1996. Fauna Evropeiskogo Severo-Vostoka Rossii 4. Amfibii i Reptilii [Fauna of the Euro-pean North-East of Russia 4. Amphibians and Reptiles]. Nauka, St. Petersburg.

Bannikov, A.G., Darevsky, I.S., Ishchenko, V.G., Rustamov, A.K. & Szczerbak, N.N. 1977. Opredelitel Zemnovodnykh i Presmyk-ayushchikhsya Fauny SSSR [Guide to Amphibians and Reptiles of the USSR Fauna]. Prosveshchenie, Moscow.

Berger, L., Speare, R., Daszak, P., Green, D.E. et al. 1998. Chytridiomycosis causes amphibian mortality associated with population declines in the rain forest of Australia and Central America. Proc. Natl. Acad. Sci. USA 95: 9031-9036.

Blackburn, L., Nanjappa, P. & Lannoo, M.J. 2001. An Atlas of the Distribution of U.S. Amphibians. Ball State University, Muncie.

Borkin, L.J., Belimov, G.T. & Sedalishchev, V.T. 1984. New data on distribution of amphibians and reptiles in Yakutia. In: L.J. Borkin (ed.). Ecology and Faunistics of Amphibians and Reptiles of the USSR and Adjacent Countries, pp 89-101. Leningrad.

Borner, A.P., Kielland, K. & Walker, M.D. 2008. Effects of simulated climate change on plant phenology and nitrogen mineralization in Alaskan arctic tundra. Arct. Antarct. Alp. Res. 40: 27-38.

Carstensen, R., Willson, M. & Armstrong, R. 2003. Habitat use of amphibians in northern southeast Alaska. Unpublished report to Alaska Department of Fish and Game. Juneau. Discovery Southeast.

Chubbs, T.E. & Phillips, F.R. 1998. Distribution of the wood frog, Rana sylvatica, in Labrador: an update. Canadian Field-Natural-ist 112: 329-331.

Collins, J.T. & Taggart, T.W. 2002. Standard common and current scientific names for North American amphibians, turtles, rep-tiles, and crocodilians. Fifth Edition. Publication of the Center for North American Herpitology, Lawrence.

Conant, R. & Collins, J.T. 1998. A field guide to reptiles and amphibians: eastern and central North America. Third edition. Houghton Mifflin Co., Boston.

Cook, F.R. 1984. Introduction to Canadian amphibians and rep-tiles, National Museum of Natural Sciences. National Museums of Canada, Ottowa.

DeGraaf, R.M. & Rudis, D.D. 1983. Amphibians and reptiles of New England. Habitats and natural history. Univ. Massachu-setts Press.

Desroches, J.F. & Rodrigue, D. 2004. Amphibiens et reptiles du Québec et des Maritimes. Éditions Michel Quintin.

Environment and Natural Resources, GNWT 2010. NWT Amphibian and Reptile Observations 1849 to present. NWT Wildlife Management Information System Project 140. Gov-ernment of the Northwest Territories Yellowknife, NT, Canada. [accessed 18 October 2010]

Frost, D.R., Grant, T., Faivovich, J., Bain, R.H., Haas, A., Haddad, C.F.B. et al. 2006. The amphibian tree of life. Bulletin of the American Museum of Natural History 297: 1-370.

Gasc, J.P., Cabela, A., Crnobrnja-Isailovic, J., Dolmen, D., Gros-senbacher, K., Haffner, P. et al. (eds) 1997. Atlas of amphibians and reptiles in Europe. Collection Patrimoines Naturels, 29, Societas Europaea Herpetologica, Muséum National d’Histoire Naturelle & Service du Patrimoine Naturel, Paris.

Gotthardt, T.A. & Pyare, S. 2009. Alaska Amphibian Occurrence Database. Alaska Natural Heritage Program, University of Alaska Anchorage. Unpublished database available by request. [accessed 18 October 2010]

Grubb, J.C. 1972. Differential Predation by Gambusia affinis on the Eggs of Seven Species of Anuran Amphibians. American Midland Naturalist 88: 102-108.

Hammerson, G. 2004. Lithobates sylvaticus. In: IUCN 2010. IUCN Red List of Threatened Species. Version 2010.3. www.iucn-redlist.org. [accessed 22 October 2010]

Harp, E.M. & Petranka, J.W. 2006. Ranavirus in wood frogs (Rana sylvatica): Potential Sources of Transmission Within and Between Ponds. Journal of Wildlife Diseases 42: 307-318.

Ishchenko, V.G. 1978. Dinamicheskii Polimophizm Burykh Lyagushek Fauny SSSR [Dynamic Polymorphism of the Brown Frogs of USSR Fauna]. Nauka, Moscow.

IUCN, Conservation International, and NatureServe 2004. Global Amphibian Assessment. IUCN, Conservation International, and NatureServe, Washington, DC and Arlington.

Johnson, M.L. & Speare, R. 2005. Possible modes of dissemina-tion of the amphibian chytrid Batrachochytrium dendrobatidis in the environment. Dis. Aquat. Org. 65: 181-186.

Johnson, P.T.J., Lunde, K.B., Ritchie, E.G., Reaser, J.K. & Launer, A.E. 2001. Morphological abnormality patterns in a California amphibian community. Herpetologica 57: 336-352.

Kats, L.B. & Ferrer, R.P. 2003. Alien predators and amphibian declines: Review of two decades of science and the transition to conservation. Diversity and Distributions 9: 99-110.

Kuzmin, S.L. 1994. Chapter 2. Areal. In: E.I. Vorobyeva (ed.). The Siberian newt (Salamandrella keyserlingii Dybowski, 1870), pp 15-52. Zoogeography, Systematics, Morphology, Moscow.

Kuzmin, S.L. & Maslova, I.V. 2003. The amphibians of the Russian Far East. Advances in Amphibian Res. in the f. Soviet Union, vol. 8. Pensoft Publ., Sofia-Moscow.

Kuzmin, S.L. & Semenov, D.V. 2006. Conspect of the fauna of amphibians and reptiles of Russia. KMK, Moscow.

Kuzmin, S., Ishchenko, V. ,Matsui, M., Wenge, Z. & Kaneko, Y. 2008a. Salamandrella keyserlingii. In: IUCN 2010. IUCN Red List of Threatened Species. Version 2010.3. www.iucnredlist.org. [accessed 22 October 2010]

Kuzmin, S., Tarkhnishvili, D., Ishchenko, V., Tuniyev, B., Beebee, T., Anthony, B. et al. 2008b. Rana arvalis. In: IUCN 2010. IUCN Red List of Threatened Species. Version 2010.3. www.iucnredlist.org. [accessed 22 October 2010]

Kuzmin, S., Ishchenko, V., Tuniyev, B., Beebee, T., Andreone, F., Nyström, P. et al. 2008c. Rana temporaria. In: IUCN 2010. IUCN Red List of Threatened Species. Version 2010.3. www.iucnredlist.org. [accessed 22 October 2010]

Kuzmin, S., Maslova, I., Matsui, M. & Wenge, Z. 2008d. Rana amurensis. In: IUCN 2010. IUCN Red List of Threatened Spe-cies. Version 2010.3. www.iucnredlist.org. [accessed 22 Octo-ber 2010]

Landers, D.H., Simonich, S.L., Jaffe, D.A., Geiser, L.H., Camp-bell, D.H., Schwindt, A.R. et al. 2008. The Fate, Transport, and Ecological Impacts of Airborne Contaminants in Western National Parks (USA). EPA/600/R-07/138. U.S. Environ-mental Protection Agency, Office of Research and Devel-opment, NHEERL, Western Ecology Division, Corvallis, Oregon.

Lee-Yaw, J.E., Irwin, J.T. & Green, D.M. 2008. Postglacial range expansion from northern refugia by the wood frog, Rana syl-vatica. Molecular Ecology 17: 867-884.

Lemmon, E.M., Lemmon, A.R., Collins, J.T., Lee-Yaw, J.A. & Cannatella, D.C. 2007. Phylogeny-based delimitation of species boundaries and contact zones in the trilling chorus frogs (Pseudacris). Molecular Phylogenetics and Evolution 44: 1068-1082.

MacDonald, S.O. 2003. The amphibians and reptiles of Alaska. A Field Handbook. Unpublished report to U.S. Fish and Wildlife Service, Juneau, AK. aknhp.uaa.alaska.edu/herps/index.htm. [accessed 22 October 2010]

MacDonald, S.O. & Cook, J.A. 2007. Mammals and amphibians of Southeast Alaska. The Museum of Southwestern Biology, Special Publication 8: 1-191.

Martof, B.S. 1970. Rana sylvatica. Catalogue of American Am-phibians and Reptiles: 1-4.

NatureServe 2010. NatureServe Explorer: An online encyclopedia of life [web application]. Version 7.1. NatureServe, Arlington. www.natureserve.org/explorer. [accessed 1 March 2010]

Nikolsky, A.M. 1918. Faune de la Russie et des Pays Limitrophes: Amphibiens (Amphibia). Petrograd: Russ. Acad. Sci. (English transl. 1963): Fauna of Russia and Adjacent Countries: Am-phibians. Israel Progr. for Sci., Jerusalem.

Chapter 5 • Amphibians and Reptiles 191

Olson, D.H. (ed.) 2009. Herpetological Conservation in North-western North America. Northwestern Naturalist 90: 61-96.

Ouellet, M. 2000. Amphibian Deformities: Current State of Knowledge. In: D.W. Sparling, G. Linder & C.A. Bishop (eds). Ecotoxicology of Amphibians and Reptiles, pp 617-646. Society of Environmental Toxicology and Chemistry (SETAC), Pensacola.

Phillott, A.D., Speare, R., Hines, H.B., Skerratt, L.F., Meyer, E., McDonald, K.R. et al. 2010. Minimizing exposure of amphib-ians to pathogens during field studies. Diseases of Aquatic Organisms, Special Issue 4.

Pounds, J.A., Bustamonte, M.R., Coloma, L.A., Consuegra, J.A., Fogden, M.P.L., Foster, P.N. et al. 2006. Widespread amphib-ian extinctions from epidemic disease driven by global warm-ing. Nature 439: 161-167.

Poyarkov, N.A. & Kuzmin, S.L. 2008. Phylogeography of the Siberian newt Salamandrella keyserlingii by mitochondrial DNA sequence analysis. Russian J. Genetics 44: 948-958.

Reeves, M.K. & Green, D.E. 2006. Rana sylvatica Chytridiomycosis. Herpetological Review. 37: 450.

Reeves, M.K., Dolph, C.L., Zimmer, H., Tjeerdema, R.S. & Trust, K.A. 2008. Road proximity increases risk of skeletal abnormalities in wood frogs from National Wildlife Refuges in Alaska. Environmental Health Perspectives 116: 1009-1015.

Reeves, M.K., Jensen, P., Dolph, C.L. & Trust, K.A. 2010. Multiple stressors and the cause of amphibian abnormalities. Ecological Monographs 80: 423-440.

Relyea, R.A. 2000. Trait-mediated effects in larval anurans: re-versing competition with the threat of predation. Ecology 81: 2278-2289.

Roelants, K., Gower, D.J., Wilkinson, M., Loader, S.P., Biju, S.D., Guillaume, K. et al. 2007. Global patterns of diversifica-tion in the history of modern amphibians. Proceedings of the National Academy of Sciences of the United States of America 104: 887-892.

Russell, A.P. & Bauer, A.M. 1993. The amphibians and reptiles of Alberta. University of Calgary Press, Calgary, and University of Alberta Press, Edmonton.

Schock, D.M., Ruthig, G.R., Collins, J.P. & Kutz, S.J. 2009. Amphibian chytrid fungus and ranaviruses in the Northwest Territories, Canada. Dis. Aquat. Org. doi:10.3354/dao02134.

Shvarts, S.S. 1959. On the biology of amphibians above the Polar Circle (Rana terrestris and Hynobius kayserlingi). Trudy Sale-khardskogo Statsionara Uralskogo Filiala Akademii Nauk SSSR (1): 393-396.

Slough, B.G. & Mennell, R.L. 2006. Diversity and range of am-phibians of the Yukon Territory. Canadian Field-Naturalist 120: 87-92.

Smith, L.C., Sheng, Y., MacDonald, G.M. & Hinzman, L.D. 2005. Disappearing Arctic Lakes. Science 308: 1429.

Sours, G.N. & Petranka, J.W. 2007. Intraguild Predation and Competition Mediated Stage-Structured Interactions between Wood Frog (Rana sylvatica) and Upland Chorus Frog (Pseu-dacris feriarum) Larvae. Copeia 2007: 131-139.

Storey, K.B. & Storey, J.M. 1987. Persistence of Freeze Tolerance in Terrestrially Hibernating Frogs after Spring Emergence. Copeia 1987: 720-726.

Stuart, S.N., Chanson, J.S., Cox, N.A., Young, B.E., Rodrigues, A.S.L., Fischman, D.L. & Waller R.W. 2004. Status and trends of amphibian declines and extinctions worldwide. Science 306: 1783-1786.

Toporkova, L.Y. 1973. [Amphibians and reptiles of Urals]. In: Fauna Evropeiskogo Severa, Urala i Zapadnoi Sibiri, pp 84-116. Sverdlovsk.

Toporkova, L.Y. & Shvarts, S.S. 1960. Amphibians above the Polar Circle. Priroda (Moscow) (10): 85-86.

Toporkova, L.Y. & Zubareva, E.L. 1965. Materials on the ecology of Rana temporaria on the Polar Urals. Trudy Instituta Biologii Uralskogo Filiala Akademii Nauk SSSR. (38): 189-194.

Trust, K.A. & Tangermann, H. 2002. National malformed amphibian study. FY 2000: Kenai National Wildlife Refuge. Annual Progress Report. U.S. Fish and Wildlife Service. Anchorage.

Vershinin, V.L. 2007. Amfibii i Reptilii Urala [Amphibians and Reptiles of the Urals]. UrO RAN, Ekaterinburg.

Weller, W.F. & Green, D.M. 1997. Checklist and current status of Canadian amphibians. In: D.M. Green (ed.). Amphibians in decline: Canadian studies of a global problem, pp 309-328. So-ciety for the Study of Amphibians and Reptiles, Herpetological Conservation 1.

Yoshikawa, K. & Hinzman, LD. 2003. Shrinking thermokarst ponds and groundwater dynamics in discontinuous permafrost near Council, Alaska. Permafrost Periglac. Process 14: 151-160.

Zavalko, S. & Mustonen, T. 2004. Tusovka of Piras. In: T. Mun-stonen & E. Helander (eds). Snowscapes, Dreamscapes – A Snowchange Community Book on Community Voices of Change, pp 336-353. Tampere University of Applied Sciences, Tampere.

Zhitkov, B.M. 1905. Travel in Kaninskaya tundra. Report of Ex-pedition of Russian Geographical Society on Kanin Peninsula in 1902. Zapiski Russkogo Geograficheskogo Obshchestva po Obshchei Geografii 41: 1-170.