LONG-TOED Ambystoma macrodactylum Baird, 1849 (Ambystomatidae)

Global rank G5 (14Dec2001) State rank S2? (25Feb1992 reviewed 1Nov2004) State rank reasons Restricted distribution in Southeast Alaska. Possible island endemism, but genetics not available to validate. Relatively common throughout its range, but overall abundance and trends unknown. Major threats include destruction of wetland habitats, predation by introduced trout, and exposure to UV-B.

Taxonomy For phylogenetic analyses of North American Ambystoma see Kraus (1988), Shaffer et al. (1991), and Jones et al. (1993). insects and garter snakes (Thamnophis spp.);

garter snakes and bullfrogs (Rana catesbeiana) Five subspecies are currently recognized, one eat adults (Nussbaum et al. 1983). occurs in Alaska. It has been suggested that the mainland and island population in the vicinity of Migration Migrates between breeding ponds and the Stikine River of coastal Alaska are non-breeding habitat; usually migrates at night in phenotypically and taxonomically distinct conjunction with precipitation. Males reach ponds (MacDonald 2003). before females and stay longer.

General description Food A delicate, smooth-skinned dusky or black Larvae feed on zooplankton, immature insects, salamander with an irregular yellow, tan, or light snails, and occasionally other salamander larvae, green dorsal stripe. White flecking on sides. including conspecifics. Adults eat terrestrial and Ventral surface is gray. Long toes and faint costal aquatic invertebrates including: insects, insect grooves (Hodge 1976). Adults measure up to 17 larvae, spiders, slugs, earthworms, amphipods, cm. etc.

This species is easily distinguished from Alaska’s Phenology May be active almost all winter in other by its prominent dorsal stripe Pacific Northwest coastal ponds (Stebbins 1985). and long fourth toe.

Length (cm) 17 Habitat

Found in a wide variety of habitats, from semiarid Reproduction sagebrush deserts to sub-alpine meadows, Breeding season is longer and earlier (fall-early including dry woodlands, humid forests, and rocky spring) in coastal lowlands, shorter and later shores of mountain lakes. Adults are (summer) in interior mountains. Clutch size is subterranean except during the breeding season. larger at lower elevations (Howard and Wallace A terrestrial habitat use survey near Hinton, 1985). Eggs laid in water, attached to vegetation Alberta determined that individuals were found or loose on bottom. Larvae metamorphose in first primarily in well-drained areas with thick litter on summer or overwinter (high elevations). In the forest floor and close to relatively permanent Alberta, sexually mature in 2+ years; maximum water bodies (Graham 1997). were life span 10 years, usually 6 years or less (Russell also found in seral stages ranging from three- et al. 1996). year-old clear-cuts to 180-year-old forests and

occurred in active logging areas (Graham 1997). Ecology Breeds in temporary or permanent ponds, or in Predators of larvae probably include fish, aquatic quiet water at the edge of lakes and streams. During the breeding season adults may be found

Long-toed salamander 1 under logs, rocks, and other debris near water. native amphibians without a valid permit. Occurs Eggs are attached to vegetation or loose on in Stikine-Leconte Wilderness Area of the bottom. Tongass National Forest.

Global range Global threats Southeastern Alaska southward to Tuolumne In the Cascades of northern Washington, larval County, California, east to Rocky Mountains (east abundance was related to both lake productivity to east-central British Columbia, west-central and the presence of introduced trout (reduced Alberta, western Montana, and central Idaho). larval abundance when trout present) (Tyler et al. Isolated populations in Santa Cruz and Monterey 1998). In Montana, introduced trout populations counties, California (Bury et al. 1980). Sea level clearly excluded salamanders from lakes (Funk to about 10,000 ft (Stebbins 1985). and Dunlap 1999).

State range In developed areas, the destruction of wetland Alaska distribution restricted to southeastern habitats may be the greatest threat. Human coastal forests adjacent to the Stikine and Taku disturbance such as road and trail construction, River watersheds (Hodge 1976). Reported near timber harvest, grazing, and fire management can the mouth of the Stikine River at Figure Eight result in fragmentation of terrestrial habitat and Lake (Twin Lakes), Mallard Slough, Cheliped Bay, breeding ponds (Fukumoto 1995 in Graham and Andrew Slough, Farm Island and Sokolof Island. Powell 1999, Maxell 2000, Paton 2002). Larvae Also collected on the Alaska side of the Coast are sensitive to a combination of low pH and Range in the Taku River Valley (MacDonald aluminum. In the Pacific Northwest, this species 2003). appears to be particularly sensitive to UV-B exposure (Belden et al. 2000). Possible effects of Global abundance exposure to UV-B include increased mortality and Total adult population size is unknown but surely incidence of deformities, slowed growth and skin exceeds 10,000. darkening (Belden and Blaustein 2002).

State abundance State threats Relatively common throughout its range; the In developed areas, the destruction of wetland overall Alaska population size is unknown but habitats may be the greatest concern. Human considered relatively small. Waters (1992) disturbance such as road and trail construction, surveyed the Stikine River Basin during summer timber harvest, grazing, and fire management can 1991 and failed to observe this species. result in fragmentation of terrestrial habitat and breeding ponds (Fukumoto 1995 in Graham and State trend Powell 1999, Maxell 2000, Grialou et al. 2000, Unknown. Data currently not available. Paton 2002). Larvae are sensitive to a combination of low pH and aluminum. In the Global protection Pacific Northwest, this species appears to be Would benefit from protection of habitat near particularly sensitive to UV-B exposure (Belden et breeding ponds (Bury et al. 1980). Prohibit al. 2000). Possible effects of exposure to UV-B introductions of non-native fishes in salamander include increased mortality and incidence of habitat. deformities, slowed growth and skin darkening (Belden and Blaustein 2002). Trout introduction is In British Columbia, protection occurs through the of concern as they are known predators to larvae. Wildlife Act, which includes all native amphibians (Graham and Powell 1999). In Alberta, long-toed State research needs salamanders are designated a sensitive species Establish programs to monitor population trends; (Pearson 2003). identify threats and limiting factors. Genetic research needed to determine if island endemicity State protection exists. Research is needed on the effects of roads In Alaska, amphibians are managed by Alaska and logging on population persistence. Department of Fish and Game under statute 16.05.030, which legally includes amphibians in Global inventory needs the definition of “fish”. This statute makes it illegal Since there can be significant year to year for anyone to “hold, transport or release” any variation in breeding population size,

Long-toed salamander 2 long-term monitoring is necessary to determine UV-B radiation. Journal of Zoology 251:473- population trends and size (Graham and Powell 479. 1999, Paton 2002). Bury, R.B., C.K. Dodd, Jr., and G.M. Fellers. State inventory needs 1980. Conservation of the amphibia of the Additional inventory to precisely determine the United States: a review. U.S. Fish and Wildlife species' range and population estimates for each Service, Washington, DC. Resource Publ. area of occurrence are needed. 134. 34 pp.

Global conservation and management needs Funk, W.C., and W.W. Dunlap. 1999. Fisheries management could improve the status Colonization of high-elevation lakes by long- of salamander populations by preventing toed salmanders (Ambystoma introduction of fishes into salamander habitats macrodactylum) after the extinction of where fishes are not native. Removal of non- introduced trout populations. Canadian native fishes from otherwise favorable Journal of Zoology. 77:1759-1767. salamander habitat is appropriate in many locations. Information on genetic variation on a Graham, K.L. 1997. Habitat use by long-toed large scale (subspecies) and population scale salamanders (Ambystoma macrodactylum) at would be useful for management decision-making three different scales. M.S. Thesis. University (Graham and Powell 1999). Montana researchers of Guelph, Ottawa, Ontario. 71 pp. recommend using only herbicide and pesticide brands that rapidly decompose and not spraying Graham, K.L. and G.L. Powell. 1999. Status of within 300m of water bodies or wetlands (Joslin the long-toed salamander (Ambystoma and Youmans 1999 in Paton 2002). Logging macrodactylum) in Alberta. Edmonton, AB. activities in areas with long-toed salamanders Alberta Sustainable Resource Development, should be scheduled to occur during the winter to Fish and Wildlife Division, Alberta Species at minimize soil compaction and litter layer Risk Report No. 22. disturbance (Graham 1997, Paton 2002). Grialou, J.A., S.D. West, and R.N. Wilkins. 2000. State conservation and management needs The effects of forest clearcut harvesting and Fisheries management could improve the status thinning on terrestrial salamanders. Journal of of salamander populations by not introducing Wildlife Management. 64:105-113. fishes into salamander habitats where fishes are not native. Removal of non-native fishes from Hodge, R.P. 1976. Amphibians and reptiles in otherwise favorable salamander habitat is Alaska, the Yukon, and Northwest Territories. appropriate in many locations. Information on Alaska Northwest Publishing Company, genetic variation and island endemism would Anchorage, AK. 89 pp. provide useful data to aid in management decisions. Whenever possible, logging activities in Howard, J.H., and R.L. Wallace. 1985. Life history areas with long-toed salamanders should be characteristics of populations of the long-toed scheduled to occur during the winter to minimize salamander (Ambystoma macrodactylum) soil compaction and litter layer disturbance from different altitudes. Amer. Midl. Naturalist (Graham 1997, Paton 2002). 113:361-373.

Jones, T.R., A.G. Kluge, and A.J. Wolf. 1993. LITERATURE CITED When theories and methodologies clash: a Belden, L.K. and A.R. Blaustein. 2002. Population phylogenetic reanalysis of the North American differences in sensitivity to UV-B radiation for ambystomatid salamanders (: larval long-toed salamanders. Ecology 83: Ambystomatidae). Systematic Biology 42:92- 1586-1590. 102.

Belden, L.K., E.L. Wildy and A.R. Blaustein. 2000. Kraus, F. 1988. An empirical evaluation of the use Growth, survival and behavior of larval long- of the ontogeny polarization criterion in toed salamanders (Ambystoma phylogenetic inference. Systematic Zoology macrodactylum) exposed to ambient levels of 37:106-141.

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MacDonald, S.O. 2003.The amphibians and macrodactylum) in high-elevation lakes. reptiles of Alaska. A Field Handbook. Conservation Biology 12:94-105. Unpublished report to U.S. Fish and Wildlife Service, Juneau, AK. Waters, D.L. 1992. Habitat associations, phenology, and biogeography of amphibians Maxell, B.A. 2000. Management of Montana’s in the Stikine River basin and southeast amphibians: A review of factors that present a Alaska. Unpubl. rep. of the 1991 pilot project. risk to population viability and accounts on the U.S. Dept. Interior, Fish and Wildlife Service, identification, distribution, , habitat California Cooperative Fishery Research Unit, use, natural history and the status and Humboldt State University, Arcata, CA. 61 pp. conservation of individual species. Missoula, MT. Region 1, USDA Forest Service.

Nussbaum, R.A., E.D. Brodie, Jr., and R.M. Acknowledgements Storm. 1983. Amphibians and reptiles of the Pacific Northwest. Univ. Press of Idaho. 332 State Conservation Status, pp. Element Ecology & Life History Author(s): Gotthardt, T.A. Paton, D. 2002. Columbia mountain amphibian State Conservation Status, Element Ecology & surveys, 2001. Edmonton, AB. Alberta Life History Edition Date: 22Mar2005 Sustainable Resource Development, Fish and Wildlife Division, Alberta Species at Risk Reviewer(s): Stephen MacDonald, University of Report No. 39. New Mexico; Blain Anderson, National Park Service, Anchorage, AK. Pearson, K. 2003. Distribution and habitat associations of the long-toed salamander Life history and Global level information were (Ambystoma macrodactylum) in Oldman obtained from the on-line database, NatureServe River drainage. Edmonton, AB. Alberta Explorer (www.natureserve.org/explorer). In many Sustainable Resource Development, Fish and cases, life history and Global information were Wildlife Division, Alberta Species at Risk updated for this species account by Alaska Report No. 75. Natural Heritage Program zoologist, Tracey Gotthardt. All Global level modifications will be Russell, A.P., G.L. Powell, and D.R. Hall. 1996. sent to NatureServe to update the on-line version. Growth and age of Alberta long-toed salamanders (Ambystoma macrodactylum NatureServe Conservation Status Factors krausei): a comparison of two methods of Edition Date: 28Dec2001 estimation. Canadian Journal of Zoology NatureServe Conservation Status Factors 74:397-412. Author: Hammerson, G. Global Element Ecology & Life History Edition Shaffer, H.B., J.M. Clark, and F. Kraus. 1991. Date: 19Sep1994 When molecules and morphology clash: a Global Element Ecology & Life History phylogenetic analysis of the North American Author(s): Hammerson, G. ambystomatid salamanders (Caudata: Ambystomatidae). Systematic Zoology 40:284-303.

Stebbins, R.C. 1985. A field guide to western reptiles and amphibians. Second Edition. Houghton Mifflin Company, Boston, MA. xiv + 336 pp.

Tyler, T., W.J. Liss, L.M. Ganio, G.L. Larson, R. Hoffman, E. Deimling, and G. Lomnicky. 1998. Interaction between introduced trout and larval salamanders (Ambystoma

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