Arctic Biodiversity Assessment

Home , Common frog, Eastern Hemisphere, Manchuria, Moor Frog, Rana (genus), Rana amurensis

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. 183

Chapter 5 Amphibians and Reptiles

Authors Sergius L. Kuzmin and David F. Tessler

Contents Summary ��184 We have lizards. We have seen them this year even at 5.1. Introduction ��184 » the upper reaches of ‘Afanas’ki’. 5.2. Status of knowledge ��184 Leader of the Saami indigenous obschina ‘Piras’ in the Kola 5.2.1. Historical overview ��184 Peninsula, Andrey Yulin on northwards expansion of lizards; 5.3. Status and trends ��185 Zavalko & Mustonen (2004). 5.3.1. Species richness and distribution ��185 5.3.1.1. Eastern hemisphere species ��185 5.3.1.2. Western hemisphere species ��186 5.3.2. Status ��187 5.3.3. Trends ��187 5.3.4. Prospects ��187 5.4. Conclusions and recommendations ��189 5.4.1. Sensitive areas and hotspots ��189 5.4.2. Key knowledge gaps ��189 5.4.3. Recommended conservation actions ��189 5.4.3.1. Research recommendations ��189 5.4.3.2. Conservation action recommendations ��189 Acknowledgements ��189 References ��189 184 Arctic Biodiversity Assessment

SUMMARY 5.2. STATUS OF KNOWLEDGE The herpetofauna of the Arctic is depauperate relative The first scientific records of Arctic amphibians are from to temperate and tropical regions. Only five amphibians expeditions in the 19th and early 20th centuries. Since and a single reptile range into the Arctic, and none are then, despite intensive study of Arctic regions, little circumpolar. All Arctic amphibian and reptile taxa are attention has been paid to understanding the distribu- currently categorized as ‘Least Concern’ according to tion and ecology of Arctic amphibians and reptiles. IUCN criteria. However, basic survey and inventory data Few works on the distribution, morphology, genetics, for these species are lacking across most of the Arctic, phenology, development, hibernation and diet of Arctic and there are few quantitative data on abundance, status amphibians have been published. or trends for Arctic herpetofauna. At the same time, isolated populations of amphibians and reptiles in the Amphibians and reptiles reach Arctic regions only on the Arctic exist at or near their current physiological limits periphery of their ranges, where their overall abundance and likely face a number of escalating challenges stem- is low. Due to their limited dispersal capabilities and ming primarily from habitat alteration. because their distributions are defined by fine-scale microhabitat associations, their actual distributions may be quite patchy (Olson 2009). Because focused research 5.1. INTRODUCTION on amphibian and reptile biology in the Arctic is scarce, much of the data available were obtained as a by-product Although amphibians and reptiles account for nearly of other efforts. 15,000 species worldwide, only five amphibians and a single reptile are found in the Arctic. The majority 5.2.1. Historical overview of Arctic herpetofauna are found in the eastern hemisphere; there are no circumpolar taxa (Tab. 5.1). Am- Historical information on the Arctic distributions of phibian species richness (number of species) in the Arctic amphibians and reptiles is very limited. To date, there is as low as in desert regions. Amphibians and reptiles have been only two published studies on the historical are phylogenetically the oldest of terrestrial vertebrates, phylogeography of Arctic herpetofauna using molecular and their limited representation in the Arctic is due in techniques. large measure to their poikilothermic physiology (body temperature determined by ambient conditions). Poyarkov & Kuzmin (2008) investigated molecular genetics in the Siberian newt throughout its range and A number of recent publications have suggested ma- found that patterns of genetic differentiation of Siberian jor changes to herpetological systematics (Frost et al. populations are likely the result of repeated processes of 2006, Roelants et al. 2007), but because these proposed colonization of new territories during inter-glacial ep- changes are not yet universally accepted and many names ochs and subsequent retreats into more temperate belts remain in a state of flux, we follow stable herpetological during glacial peaks. Following the peak of the most taxonomy as described in Collins & Taggart (2002) and recent glacial maximum, the Siberian newt likely first Kuzmin & Semenov (2006). colonized territories in E Siberia, followed by colonization events west towards the Urals and east towards Eastern hemisphere taxa: Beringia and Kamchatka. Dispersal to the north and Siberian newt Salamandrella keyserlingii west appears to have taken place very quickly. Common frog Rana temporaria Moor frog Rana arvalis In North America, an investigation of the historic phy- Siberian wood frog Rana amurensis logeography of the wood frog using mitochondrial genes Common lizard Lacerta vivipara suggests a post glacial range expansion that differs from Western hemisphere taxa: most other herpetofaunal taxa on the continent (Lee- Wood frog Rana sylvatica 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 Subspecies 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

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. from a number of high-latitude refugia in what is now species richness of amphibians and reptiles appears to be the northeastern United States, while most amphibians zero or one. currently found in western and northwestern North America appear to have radiated from lower-latitude ref- 5.3.1.1. Eastern hemisphere species ugia to the west. Colonization following the last glacial maximum appears to have been rapid, with expansion to The Siberian newt is the most widespread amphibian the north and northwest (Arctic Alaska and most of subspecies in the Arctic and sub-Arctic (Fig. 5.1), and has Arctic Canada) having occurred from a single putative the widest geographical range of any recent amphibian refugium near the edge of the Laurentide ice-sheet in species, c. 12 million km2. The northernmost habitats present-day Wisconsin, while radiation into sub-Arctic of the Siberian newt consist of grass-undershrub-lichen- Quebec, Newfoundland and Labrador took place from moss bogs and low shrub-moss and grass-moss tundras. another refugium near the Appalachian Mountains in the The newt penetrates the Arctic in the area of the polar vicinity of Pennsylvania. 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 5.3. STATUS AND TRENDS some areas of the Republic of Sakha-Yakutia, in particular, near Chukochya Guba in Nizhnekolymskii. The 5.3.1. Species richness and distribution Amguema River in the Shmidtovskii District of Chukot- ka marks the northeastern extent of its distribution. Maximum Arctic amphibian and reptile richness, three species, occurs south of the Yamal Peninsula, where Frogs Rana spp. are more narrowly distributed in the the European common frog and moor frog coexist with Arctic than the Siberian newt. The common frog and the Siberian newt. The moor frog and common frog the moor frog are mainly inhabitants of Europe but both are sympatric on the Kanin Peninsula and in the Vor- range eastwards to the Urals, with the moor frog found kuta region, while the Siberian newt and Siberian wood into E Siberia. The common frog crosses into the low frog both inhabit the Khaiyr Settlement in Republic of Arctic only in the northernmost peninsulas of Norway Sakha-Yakutia. Throughout the remainder of the Arctic, (Gasc et al.1997) and along the eastern slope of the polar 186 Arctic Biodiversity Assessment

Urals at the southern border of the Yamal Peninsula, in The common lizard is the only reptile species found in the Priuralskii District of Yamal-Nenetskii Autonomous the Arctic. It ranges from northwestern Spain through Okrug, Tyumenskaya Province, Russia (Toporkova & Europe and Siberia to Mongolia and eastwards to Sakha- Shvarts 1960, Toporkova & Zubareva 1965, Toporkova lin Island in the Pacific. It reaches the Arctic in Europe 1973, Ishchenko 1978). The common frog has been in the vicinity of the Kanin Peninsula and around Vor- known to occur on the Kanin Peninsula since at least kuta City in the Komi Republic in Russia (Anufriev & 1902 (Zhitkov 1905). Bobretsov 1996). The common lizard is found in the Po- lar Urals, southwards from the Yamal Peninsula, where The moor frog has a broader Arctic distribution than it occurs primarily in river valleys (Vershinin 2007). In the common frog. It occurs in southern Yamal and the the area of Taimyr Peninsula the common lizard occurs Polar Urals, and as far north as the Khadyta-Yakha River in the sub-Arctic near the southern boundary of the low (Shvarts 1959). A few records indicate it is found on the Arctic (Bannikov et al. 1977). sub-Arctic/low Arctic margin to the east as far as the Malaya Khadyta River in Yamal (Anufriev 1984). 5.3.1.2. Western hemisphere species The Siberian wood frog is a widespread brown frog, The wood frog is the only herpetofaunal species found whose distribution covers a large part of Siberia, far in the Arctic of the Western hemisphere. This widely eastern Russia, and northern Mongolia and Manchuria. distributed North American frog is found throughout the Nevertheless, this species is known from only one Arctic sub-Arctic (in Labrador, Quebec, Ontario, Manitoba, locality: Khaiyr Settlement in Ust-Yanskii District of the Saskatchewan, Alberta). It extends into the low Arctic in Republic of Sakha-Yakutia (Borkin et al. 1984). 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 indicative 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 & The common frog is considered common and stable Rudis 1983, Cook 1984, Russell & Bauer 1993, Weller throughout its range, though local declines have been & Green 1997, Chubbs & Phillips 1998, Blackburn et noted in Switzerland and Spain (Kuzmin et al. 2008c), al. 2001, Trust & Tangermann 2002, Carstensen et al. and since the 1970s in Norway along the northern limits 2003, MacDonald 2003, Anderson 2004, Desroches & of its distribution (Gasc et al. 1997). Rodrigue 2004, Slough and Mennell 2006, MacDonald & Cook 2007, Lee-Yaw et al. 2008, NatureServe 2010). Globally, the Siberian wood frog is considered com- This unique frog ranges at least as far north as the south mon, stable and a species of least concern (Kuzmin et side of the Brooks Range in Alaska, the Old Crow Flats in al. 2008d), though there is no information on Arctic the Yukon, and the Mackenzie River Delta in Northwest abundance or trends. Territories (Fig. 5.2). Despite its wide distribution and apparent ubiquity, there are only eight confirmed records Population size, trend and status data for the common of wood frogs extending into the Arctic as defined by lizard are lacking throughout its large distribution, in- Geographic Information System data layers developed for cluding the Arctic, but it is classified as ‘Least Concern’ the Arctic Biodiversity Assessment (MacDonald 2003, by the IUCN (2010). Gotthardt & Pyare 2009, Environment and Natural Resources, GNWT 2010). The species is probably more The wood frog is the most widespread amphibian in widespread in the Arctic, and the limited number of North America (Martof 1970), the most common am- records is likely due in part to very limited survey effort. phibian in Alaska (MacDonald 2003), and is considered relatively common throughout most of its range, includ- 5.3.2. Status ing the sub-Arctic. There are no data on abundance, status or trends for this species in the Arctic. The global All of the amphibians and reptiles found in the Arctic are population trend is unknown but is suspected to be considered taxa of ‘Least Concern’ by IUCN, suggest- stable, and the wood frog is considered a species of least ing their populations are currently stable. However, as a conservation concern (Hammerson 2004). class, amphibians are among the most globally threatened groups, with roughly 32% (1,856 species) considered 5.3.4. Prospects threatened and 43% (2,469 species) declining. At least 168 species worldwide have become extinct over the last In general, the outlook for amphibian persistence in the 20 years, and the numbers of threatened and extinct taxa Arctic is probably good. The species all appear secure are expected to continue to climb (Stuart et al. 2004). globally, with Arctic populations representing the northern fringe of much larger distributions to the south. While there are no indications of declines of amphibian However, conservation of Arctic reptiles and amphibians and reptile populations in the Arctic, their apparent sta- will face a number of challenges in the immediate future bility may be a result of the almost complete lack of his- and over the next century, including contaminants from toric and contemporary abundance data. In a few cases local and global sources, anthropogenic habitat altera- there are some qualitative estimates of local abundance, tion, emerging infectious diseases, climate change-in- but these are insufficient to determine the dynamics of duced habitat alteration and loss, and the introduction of even local populations. Overall numbers of amphibians novel pathogens and predators. and reptiles in the Arctic appear to be low and populations appear scattered and isolated. Environmental contaminants originating either from local point sources or from diffuse regional and/or global 5.3.3. Trends sources may impact many otherwise undisturbed Arctic wetlands. Ackerman et al. (2008) and Landers et al. Global population estimates and trend data are largely (2008) found concentrations of atmospherically depos- lacking for all herpetofauna found in the Arctic, and ited organic and other contaminants in fish from remote few if any data exist on local Arctic population sizes or lakes in Arctic and sub-Arctic Alaska that exceeded trends for any of the taxa. thresholds of health concern for humans and wildlife. Because the study sites were all located in remote areas The Siberian newt has been known as an Arctic spe- with no local contaminant sources, their presence was cies since 1909 (Nikolsky 1918), and is considered to be attributed to long-range trans-Pacific transport and to common and stable throughout its range, with the ex- global sources. The risk to amphibians is unclear and ception of some small, declining and isolated populations undocumented, but due to their aquatic developmental in Mongolia, which are considered threatened (Kuzmin phase, moist and highly permeable integuments, larval et al. 2008a). diets of zooplankton, phytoplankton and periphyton and adult diets of higher trophic level invertebrates, they may The moor frog, though extinct in Switzerland, is con- be predisposed to bio-accumulate a variety of contami- sidered common, stable and a species of ‘Least con- nants when present in the environment. cern’ by the IUCN throughout its distribution (Kuzmin et al. 2008b). 188 Arctic Biodiversity Assessment

As development inevitably proceeds in the Arctic, of ranaviruses (genus Iridoviridae) are also associated with many wetland habitats will be transformed, and even recent amphibian declines (Stuart et al. 2004). There has those that remain intact may be exposed to additional been little surveillance for Bd in the Arctic to date, but threats as roads and other infrastructure elements are the fungus has already been detected in sub-Arctic Alas- constructed. Research on wood frog breeding ponds ka, and both Bd and ranaviruses have been detected in at two National Wildlife Refuges in sub-Arctic Alaska Canada’s Northwest Territories (Reeves & Green 2006, documented some of the highest rates of skeletal abnor- Schock et al. 2009, T. Chestnut pers. com.). Infrastruc- malities found in amphibians anywhere; as high as 20% ture development and increased human presence in the of individuals from some breeding sites had pronounced Arctic may contribute to the spread of these pathogens: skeletal abnormalities (Reeves et al. 2008). The normal Both ranaviruses and Bd may be translocated between background rate for such abnormalities is estimated as water bodies via transport of contaminated water or either 0-2% (Ouellet 2000) or 0-5% (Johnson et al. sediments by humans (on footwear and equipment), birds 2001). The prevalence of structural abnormalities in (e.g. on feathers), or other animals (Johnson & Speare Alaska was found to increase with proximity to roads, 2005, Harp & Petranka 2006, Phillot et al. 2010). and more recently, the presence of contaminants (both organic and inorganic) and odonate larvae were identi- Warmer temperatures and expanded human industry fied as predictors of the frequency of skeletal abnormali- and infrastructure will increase the risks of introducing ties at these sites (Reeves et al. 2010). exotic or novel species into Arctic environments. Arctic amphibians breed primarily in fish-free water bodies, in- As climatic conditions in the Arctic continue to amelio- cluding ephemeral pools and small lakes and ponds, and rate, we might expect the amphibians already present to it is widely believed that this preference is an adaptation expand their current distributions and colonize previ- to avoid predation. Fish are successful predators on the ously unoccupied areas. However, a warming climate eggs and aquatic larvae of amphibians (Semlitch 1988), will also catalyze a suite of interconnected changes in and many amphibians have few defenses against fish the physical and biological environments of the Arctic, (Grubb 1972). Introductions of non-native fishes have the ramifications of which are poorly understood for been implicated as one of the major causes of amphib- Arctic amphibians. Warmer winter and summer tem- ian population declines (Kats & Ferrer 2003), and even peratures, alteration in both the rates and the timing of native fish introduced into previously fish free habitats snow deposition and melt-off, and permafrost thaw will have the potential to extirpate local amphibian popula- have profound effects on hydrology and hydroperiod tions. Increased human presence may serve as a vector across the Arctic. Both the total volume of water and for the spread of native and non-native fish across naïve the timing of its availability to plants and animals may be Arctic wetlands. Amphibian larvae are also preyed on by significantly altered (McDonald et al. 2004, Borner et al. a variety of birds, other amphibians (Sours & Petranka 2008; see Wrona & Reist, Chapter 13) resulting in shifts 2007) and a number of aquatic invertebrates, especially in the composition of the biotic community. Permafrost odonates, whose abundance appears related to water thaw, which is forecast to continue at an accelerated temperature (Reeves et al. 2010). Trophic webs and rate, is already causing the draining of Arctic wetlands competitive interactions among amphibians in Arctic in Alaska (Yoshikawa & Hinzman 2003). In regions of wetlands aren’t well understood, but changes in spe- thawing discontinuous permafrost in Siberia, the num- cies composition or density can strongly influence the ber of large lakes declined by 11% and overall wetland outcomes of competition and predation in these aquatic surface area declined by 6% since the 1970s (Smith et communities (Relyea 2000, Sours & Petranka 2007). al. 2005). These changes in hydroperiod and hydrology, The consequences for Arctic amphibian populations of and their attendant impacts on the biotic community, any new or heightened predatory or competitive interac- have the potential to disrupt many facets of amphibian tions are completely unknown. ecology, including: (1) breeding phenology relative to the timing and availability of algae and the emergence of While most sub-Arctic amphibians are unlikely to important aquatic invertebrate prey (and predator) spe- colonize the Arctic in the next century (due to their cies, (2) habitat connectivity, (3) dispersal movements, southerly distributions, inherently slow rate of dispersal (4) juvenile and adult survival, and ultimately (5) the and less freeze-tolerant physiologies) the boreal cho- persistence of local populations. rus frog Pseudacris maculata in the Western hemisphere may already have reached it (Fig. 5.2). In 2009 a single Emerging infectious diseases are also likely to play a un-vouchered observation was recorded in the Arctic in greater role in the ecology of Arctic amphibians. Warm- Canada’s Northwest Territories near the border of Nuna- ing climatic conditions may aid the spread and survival of vut in the vicinity of Big Lake (Environment and Natural deadly new pathogens, and infrastructure development Resources, GNWT 2010). Very low mitochondrial may provide new vectors for transmission across the land- genetic diversity found in populations occupying regions scape. The amphibian chytrid fungus, Batrachochytrium north of the last glacial maximum suggests the boreal dendrobatidis (Bd) is a recently recognized pathogen widely chorus frog colonized its northern frontiers recently and believed to be expanding due to climate change and im- rather rapidly (Lemmon et al. 2007). If it is not already, plicated in a number of amphibian declines and extirpa- the boreal chorus frog will soon be the second Nearctic tions (Berger et al. 1998, Pounds et al. 2006). A number amphibian species. Chapter 5 • Amphibians and Reptiles 189

5.4. CONCLUSIONS AND monitoring, and to improve our understanding of the RECOMMENDATIONS dynamics of Arctic ecosystems. • Conduct research into the impacts of climate-induced changes to hydrology/hydroperiod on reproduction, 5.4.1. Sensitive areas and hotspots persistence and habitat connectivity for Arctic amphibians. Hotspots are difficult to identify because the distribu- • Determine the geographic prevalence of contaminant tions of Arctic amphibians and reptiles are so poorly burdens and chief pathogens for amphibians across the characterized. Furthermore, these species are found Arctic. in small, isolated and patchily distributed populations for a variety of reasons: (1) their Arctic range limits These efforts may involve citizen science projects. likely represent each species’ physiological limitations, (2) amphibians require a number of different micro- 5.4.3.2. Conservation action recommendations habitat features to intersect in close proximity due to their limited movement potential, and (3) they may not • Develop guidelines for human development projects occupy all suitable habitats within their apparent range that require land managers and developers to consider due to the susceptibility of small populations to stochas- amphibian and reptile habitats and populations in their tic events and the residual effects of past disturbances development plans. (Olson 2009). Nonetheless, a few areas in the Eastern • Determine which areas are of special importance for hemisphere appear to be of particular importance: the amphibian and reptile species richness and for the corridors and deltas of the Khadyta-Yakha River on the long-term persistence of individual taxa. Use data Yamal Peninsula, the Chaunskaya Tundra in the lowlands from survey and inventory efforts to identify hotspots of the Chaunsky Administrative District of the Chu- and areas of likely significance by modeling species’ kotsky Autonomous Okrug, and the Khalerchinskaya habitat and micro-habitat associations across the Arc- Tundra in the Kolyma lowlands. tic landscape. • Establish or strengthen protections for areas of key 5.4.2. Key knowledge gaps importance to reptiles and amphibians. Arctic amphibians have complex life cycles, and require a range The principal knowledge gap is the near complete lack of habitats throughout their annual cycles and life of survey and inventory data for status and population histories. Conservation of these species will require trends of Arctic amphibians and reptiles. Distributions a landscape-level approach, conserving various vital are poorly and incompletely characterized, and are habitats at appropriate spatial scales and maintaining known only in broad general terms. connectivity between conservation units, while ac- counting for expected wetland loss and alteration. There are no reliable abundance estimates for local or regional populations for any Arctic herpetofauna, and there are no statistically meaningful monitoring efforts ACKNOWLEDGEMENTS currently in place. General lack of understanding of the factors which limit amphibian and reptile populations in We wish to thank M. Reeves, B. Slough and D. Payer the Arctic is also a principal knowledge gap. for their thoughtful reviews of this manuscript, O.L. Leontyeva and S. Aikens for their unpublished data on 5.4.3. Recommended conservation actions Arctic frogs, and T. Chestnut for unpublished data on the prevalence of Batrachochytrium dendrobatidis in Alaska.

5.4.3.1. Research recommendations REFERENCES • Establish effective survey and inventory efforts to bet- ter define the actual distributions and ecology of these Ackerman, L.K., Schwindt, A.R., Simonich, S.L., Koch, D.C., species. Blett, T.F., Schreck, C.B. et al. 2008. Atmospherically de- • Construct statistically defensible baselines of abun- posited PBDEs, pesticides, PCBs, and PAHs in western U.S. National Park fish: concentrations and consumption guidelines. dance data in specific locations against which changes Environ Sci. Technol. 42: 2334-2341. in abundance can be monitored. Alaska Natural Heritage Program, University of Alaska Anchorage • Establish monitoring programs with replicate schema (NatureServe) 2007. Wood frog, Alaska Status Report. aknhp. representative of the range of habitats and microhabi- uaa.alaska.edu/zoology/Zoology_Amphibs_track07.htm. [ac- tats inhabited by each species. Monitoring locations cessed 1 March 2010] Amphibians of the former USSR, Databank ® 0229803415, State should also be chosen in such a way so as to minimize Register of Databanks of Russian Federation. the effort and expense to reach them in order to in- Anderson, B.C. 2004. An opportunistic amphibian inventory in crease the likelihood that monitoring will be contin- Alaska’s national parks 2001-2003. National Park Service, ued into the future. If practicable, monitoring efforts Inventory and Monitoring Program, Anchorage. should be collocated with monitoring efforts for other Anufriev, V.M. 1984. Number and distribution of Rana arvalis in taxa in order to develop economies of scale for all 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: Harp, E.M. & Petranka, J.W. 2006. Ranavirus in wood frogs Voprosy Gerpetologii. Sverdlovsk: Inst. Plant and Anim. Ecol. (Rana sylvatica): Potential Sources of Transmission Within and Ural Branch of USSR Acad. Sci. Between Ponds. Journal of Wildlife Diseases 42: 307-318. Anufriev, V.M. & Bobretsov, A.B. 1996. Fauna Evropeiskogo Ishchenko, V.G. 1978. Dinamicheskii Polimophizm Burykh Severo-Vostoka Rossii 4. Amfibii i Reptilii [Fauna of the Euro- Lyagushek Fauny SSSR [Dynamic Polymorphism of the Brown pean North-East of Russia 4. Amphibians and Reptiles]. Nauka, Frogs of USSR Fauna]. Nauka, Moscow. St. Petersburg. IUCN, Conservation International, and NatureServe 2004. Global Bannikov, A.G., Darevsky, I.S., Ishchenko, V.G., Rustamov, A.K. & Amphibian Assessment. IUCN, Conservation International, Szczerbak, N.N. 1977. Opredelitel Zemnovodnykh i Presmyk- and NatureServe, Washington, DC and Arlington. ayushchikhsya Fauny SSSR [Guide to Amphibians and Reptiles Johnson, M.L. & Speare, R. 2005. Possible modes of dissemina- of the USSR Fauna]. Prosveshchenie, Moscow. tion of the amphibian chytrid Batrachochytrium dendrobatidis in Berger, L., Speare, R., Daszak, P., Green, D.E. et al. 1998. the environment. Dis. Aquat. Org. 65: 181-186. Chytridiomycosis causes amphibian mortality associated with Johnson, P.T.J., Lunde, K.B., Ritchie, E.G., Reaser, J.K. & population declines in the rain forest of Australia and Central Launer, A.E. 2001. Morphological abnormality patterns in a America. Proc. Natl. Acad. Sci. USA 95: 9031-9036. California amphibian community. Herpetologica 57: 336-352. Blackburn, L., Nanjappa, P. & Lannoo, M.J. 2001. An Atlas of the Kats, L.B. & Ferrer, R.P. 2003. Alien predators and amphibian Distribution of U.S. Amphibians. Ball State University, Muncie. declines: Review of two decades of science and the transition Borkin, L.J., Belimov, G.T. & Sedalishchev, V.T. 1984. New to conservation. Diversity and Distributions 9: 99-110. data on distribution of amphibians and reptiles in Yakutia. In: Kuzmin, S.L. 1994. Chapter 2. Areal. In: E.I. Vorobyeva (ed.). The L.J. Borkin (ed.). Ecology and Faunistics of Amphibians and Siberian newt (Salamandrella keyserlingii Dybowski, 1870), pp Reptiles of the USSR and Adjacent Countries, pp 89-101. 15-52. Zoogeography, Systematics, Morphology, Moscow. Leningrad. Kuzmin, S.L. & Maslova, I.V. 2003. The amphibians of the Russian Borner, A.P., Kielland, K. & Walker, M.D. 2008. Effects of Far East. Advances in Amphibian Res. in the f. Soviet Union, simulated climate change on plant phenology and nitrogen vol. 8. Pensoft Publ., Sofia-Moscow. mineralization in Alaskan arctic tundra. Arct. Antarct. Alp. Res. Kuzmin, S.L. & Semenov, D.V. 2006. Conspect of the fauna of 40: 27-38. amphibians and reptiles of Russia. KMK, Moscow. Carstensen, R., Willson, M. & Armstrong, R. 2003. Habitat use of Kuzmin, S., Ishchenko, V. ,Matsui, M., Wenge, Z. & Kaneko, Y. amphibians in northern southeast Alaska. Unpublished report 2008a. Salamandrella keyserlingii. In: IUCN 2010. IUCN Red to Alaska Department of Fish and Game. Juneau. Discovery List of Threatened Species. Version 2010.3. www.iucnredlist. Southeast. org. [accessed 22 October 2010] Chubbs, T.E. & Phillips, F.R. 1998. Distribution of the wood frog, Kuzmin, S., Tarkhnishvili, D., Ishchenko, V., Tuniyev, B., Beebee, Rana sylvatica, in Labrador: an update. Canadian Field-Natural- T., Anthony, B. et al. 2008b. Rana arvalis. In: IUCN 2010. ist 112: 329-331. IUCN Red List of Threatened Species. Version 2010.3. www. Collins, J.T. & Taggart, T.W. 2002. Standard common and current iucnredlist.org. [accessed 22 October 2010] scientific names for North American amphibians, turtles, rep- Kuzmin, S., Ishchenko, V., Tuniyev, B., Beebee, T., Andreone, F., tiles, and crocodilians. Fifth Edition. Publication of the Center Nyström, P. et al. 2008c. Rana temporaria. In: IUCN 2010. for North American Herpitology, Lawrence. IUCN Red List of Threatened Species. Version 2010.3. www. Conant, R. & Collins, J.T. 1998. A field guide to reptiles and iucnredlist.org. [accessed 22 October 2010] amphibians: eastern and central North America. Third edition. Kuzmin, S., Maslova, I., Matsui, M. & Wenge, Z. 2008d. Rana Houghton Mifflin Co., Boston. amurensis. In: IUCN 2010. IUCN Red List of Threatened Spe- Cook, F.R. 1984. Introduction to Canadian amphibians and rep- cies. Version 2010.3. www.iucnredlist.org. [accessed 22 Octo- tiles, National Museum of Natural Sciences. National Museums ber 2010] of Canada, Ottowa. Landers, D.H., Simonich, S.L., Jaffe, D.A., Geiser, L.H., Camp- DeGraaf, R.M. & Rudis, D.D. 1983. Amphibians and reptiles of bell, D.H., Schwindt, A.R. et al. 2008. The Fate, Transport, New England. Habitats and natural history. Univ. Massachu- and Ecological Impacts of Airborne Contaminants in Western setts Press. National Parks (USA). EPA/600/R-07/138. U.S. Environ- Desroches, J.F. & Rodrigue, D. 2004. Amphibiens et reptiles du mental Protection Agency, Office of Research and Devel- Québec et des Maritimes. Éditions Michel Quintin. opment, NHEERL, Western Ecology Division, Corvallis, Environment and Natural Resources, GNWT 2010. NWT Oregon. Amphibian and Reptile Observations 1849 to present. NWT Lee-Yaw, J.E., Irwin, J.T. & Green, D.M. 2008. Postglacial range Wildlife Management Information System Project 140. Gov- expansion from northern refugia by the wood frog, Rana syl- ernment of the Northwest Territories Yellowknife, NT, Canada. vatica. Molecular Ecology 17: 867-884. [accessed 18 October 2010] Lemmon, E.M., Lemmon, A.R., Collins, J.T., Lee-Yaw, J.A. Frost, D.R., Grant, T., Faivovich, J., Bain, R.H., Haas, A., & Cannatella, D.C. 2007. Phylogeny-based delimitation of Haddad, C.F.B. et al. 2006. The amphibian tree of life. Bulletin species boundaries and contact zones in the trilling chorus of the American Museum of Natural History 297: 1-370. frogs (Pseudacris). Molecular Phylogenetics and Evolution 44: Gasc, J.P., Cabela, A., Crnobrnja-Isailovic, J., Dolmen, D., Gros- 1068-1082. senbacher, K., Haffner, P. et al. (eds) 1997. Atlas of amphibians MacDonald, S.O. 2003. The amphibians and reptiles of Alaska. A and reptiles in Europe. Collection Patrimoines Naturels, 29, Field Handbook. Unpublished report to U.S. Fish and Wildlife Societas Europaea Herpetologica, Muséum National d’Histoire Service, Juneau, AK. aknhp.uaa.alaska.edu/herps/index.htm. Naturelle & Service du Patrimoine Naturel, Paris. [accessed 22 October 2010] Gotthardt, T.A. & Pyare, S. 2009. Alaska Amphibian Occurrence MacDonald, S.O. & Cook, J.A. 2007. Mammals and amphibians Database. Alaska Natural Heritage Program, University of of Southeast Alaska. The Museum of Southwestern Biology, Alaska Anchorage. Unpublished database available by request. Special Publication 8: 1-191. [accessed 18 October 2010] Martof, B.S. 1970. Rana sylvatica. Catalogue of American Am- Grubb, J.C. 1972. Differential Predation by Gambusia affinis on phibians and Reptiles: 1-4. the Eggs of Seven Species of Anuran Amphibians. American NatureServe 2010. NatureServe Explorer: An online encyclopedia Midland Naturalist 88: 102-108. of life [web application]. Version 7.1. NatureServe, Arlington. Hammerson, G. 2004. Lithobates sylvaticus. In: IUCN 2010. IUCN www.natureserve.org/explorer. [accessed 1 March 2010] Red List of Threatened Species. Version 2010.3. www.iucn- Nikolsky, A.M. 1918. Faune de la Russie et des Pays Limitrophes: redlist.org. [accessed 22 October 2010] 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- Vershinin, V.L. 2007. Amfibii i Reptilii Urala [Amphibians and western North America. Northwestern Naturalist 90: 61-96. Reptiles of the Urals]. UrO RAN, Ekaterinburg. Ouellet, M. 2000. Amphibian Deformities: Current State of Weller, W.F. & Green, D.M. 1997. Checklist and current status Knowledge. In: D.W. Sparling, G. Linder & C.A. Bishop of Canadian amphibians. In: D.M. Green (ed.). Amphibians in (eds). Ecotoxicology of Amphibians and Reptiles, pp 617-646. decline: Canadian studies of a global problem, pp 309-328. So- Society of Environmental Toxicology and Chemistry (SETAC), ciety for the Study of Amphibians and Reptiles, Herpetological Pensacola. Conservation 1. Phillott, A.D., Speare, R., Hines, H.B., Skerratt, L.F., Meyer, E., Yoshikawa, K. & Hinzman, LD. 2003. Shrinking thermokarst McDonald, K.R. et al. 2010. Minimizing exposure of amphib- ponds and groundwater dynamics in discontinuous permafrost ians to pathogens during field studies. Diseases of Aquatic near Council, Alaska. Permafrost Periglac. Process 14: 151- Organisms, Special Issue 4. 160. Pounds, J.A., Bustamonte, M.R., Coloma, L.A., Consuegra, J.A., Zavalko, S. & Mustonen, T. 2004. Tusovka of Piras. In: T. Mun- Fogden, M.P.L., Foster, P.N. et al. 2006. Widespread amphib- stonen & E. Helander (eds). Snowscapes, Dreamscapes – A ian extinctions from epidemic disease driven by global warm- Snowchange Community Book on Community Voices of ing. Nature 439: 161-167. Change, pp 336-353. Tampere University of Applied Sciences, Poyarkov, N.A. & Kuzmin, S.L. 2008. Phylogeography of the Tampere. Siberian newt Salamandrella keyserlingii by mitochondrial DNA Zhitkov, B.M. 1905. Travel in Kaninskaya tundra. Report of Ex- sequence analysis. Russian J. Genetics 44: 948-958. pedition of Russian Geographical Society on Kanin Peninsula Reeves, M.K. & Green, D.E. 2006. Rana sylvatica Chytridiomycosis. in 1902. Zapiski Russkogo Geograficheskogo Obshchestva po Herpetological Review. 37: 450. Obshchei Geografii 41: 1-170. 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 amphibians 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.