Long-toed Salamander - Ambystoma macrodactylum
Adults are dark gray to black with an irregular (sometimes broken or rarely absent) green to yellow stripe down the middle of the back. The longest toe on the hind foot is longer than the sole; this species lacks a groove running vertically from the nostril to the mouth. Adult body length is 2 to 3.25 inches. Eggs and Larvae: Eggs are typically laid in ponds in small clusters of 5 to 100, but eggs may be laid singly. Larvae are usually brown, have three external gills, and are relatively small (less than 1.75-inch body) and slender.
Western Hemisphere Range
Summary of Observations Submitted for Montana
Number of Observations:
(Click on the following maps and charts to see full sized version)
Map Help and Descriptions
(Observations spanning multiple months or years are excluded from time charts)
Long-toed Salamanders are found in a variety of habitats from sagebrush to alpine. They typically breed in ponds or lakes, usually those without fish present. Adults go to the breeding ponds immediately after snowmelt and in western Montana are usually the first amphibians to breed. Like all salamanders, they have internal fertilization. Following breeding, they move to adjacent uplands. Eggs hatch in 3 to 6 weeks and metamorphosis takes 2 to 14 months. Migration routes to breeding pools showed no preference to habitat, relative soil moisture or vegetation, although individuals tended to move through same habitat (Beneski et al. 1986).
Ecological Systems Associated with this Species
- Details on Creation and Suggested Uses and Limitations
How Associations Were Made
We associated the use and habitat quality (high, medium, or low) of each of the 82 ecological systems mapped in Montana for
vertebrate animal species that regularly breed, overwinter, or migrate through the state by:
- Using personal observations and reviewing literature that summarize the breeding, overwintering, or migratory habitat requirements of each species (Dobkin 1992, Hart et al. 1998, Hutto and Young 1999, Maxell 2000, Foresman 2001, Adams 2003, and Werner et al. 2004);
- Evaluating structural characteristics and distribution of each ecological system relative to the species’ range and habitat requirements;
- Examining the observation records for each species in the state-wide point database associated with each ecological system;
- Calculating the percentage of observations associated with each ecological system relative to the percent of Montana covered by each ecological system to get a measure of “observations versus availability of habitat”.
Species that breed in Montana were only evaluated for breeding habitat use, species that only overwinter in Montana were only evaluated for overwintering habitat use, and species that only migrate through Montana were only evaluated for migratory habitat use.
In general, species were associated as using an ecological system if structural characteristics of used habitat documented in the literature were present in the ecological system or large numbers of point observations were associated with the ecological system.
However, species were not associated with an ecological system if there was no support in the literature for use of structural characteristics in an ecological system, even if
point observations were associated with that system.
High, medium, and low habitat quality was assigned based on the degree to which the structural characteristics of an ecological system matched the preferred structural habitat characteristics for each species in the literature.
The percentage of observations associated with each ecological system relative to the percent of Montana covered by each ecological system was also used to guide assignments of habitat quality.
If you have any questions or comments on species associations with ecological systems, please contact Bryce Maxell at email@example.com
or (406) 444-3655.
Suggested Uses and Limitations
Species associations with ecological systems should be used to generate potential lists of species that may occupy broader landscapes for the purposes of landscape-level planning.
These potential lists of species should not be used in place of documented occurrences of species (this information can be requested at: http://mtnhp.org/requests/default.asp
) or systematic surveys for species and evaluations of habitat at a local site level by trained biologists.
Users of this information should be aware that the land cover data used to generate species associations is based on imagery from the late 1990s and early 2000s and was only intended to be used at broader landscape scales.
Land cover mapping accuracy is particularly problematic when the systems occur as small patches or where the land cover types have been altered over the past decade.
Thus, particular caution should be used when using the associations in assessments of smaller areas (e.g., evaluations of public land survey sections).
Finally, although a species may be associated with a particular ecological system within its known geographic range, portions of that ecological system may occur outside of the species’ known geographic range.
- Adams, R.A. 2003. Bats of the Rocky Mountain West; natural history, ecology, and conservation. Boulder, CO: University Press of Colorado. 289 p.
- Dobkin, D. S. 1992. Neotropical migrant land birds in the Northern Rockies and Great Plains. USDA Forest Service, Northern Region. Publication No. R1-93-34. Missoula, MT.
- Foresman, K.R. 2001. The wild mammals of Montana. Special Publication No. 12. Lawrence, KS: The American Society of Mammalogists. 278 p.
- Hart, M.M., W.A. Williams, P.C. Thornton, K.P. McLaughlin, C.M. Tobalske, B.A. Maxell, D.P. Hendricks, C.R. Peterson, and R.L. Redmond. 1998. Montana atlas of terrestrial vertebrates. Montana Cooperative Wildlife Research Unit, University of Montana, Missoula, MT. 1302 p.
- Hutto, R.L. and J.S. Young. 1999. Habitat relationships of landbirds in the Northern Region, USDA Forest Service, Rocky Mountain Research Station RMRS-GTR-32. 72 p.
- Maxell, B.A. 2000. Management of Montana’s amphibians: a review of factors that may present a risk to population viability and accounts on the identification, distribution, taxonomy, habitat use, natural history, and the status and conservation of individual species. Report to U.S. Forest Service Region 1. Missoula, MT: Wildlife Biology Program, University of Montana. 161 p.
- Werner, J.K., B.A. Maxell, P. Hendricks, and D. Flath. 2004. Amphibians and reptiles of Montana. Missoula, MT: Mountain Press Publishing Company. 262 p.
- Commonly Associated with these Ecological Systems
Forest and Woodland Systems
Shrubland, Steppe and Savanna Systems
Wetland and Riparian Systems
- Occasionally Associated with these Ecological Systems
Forest and Woodland Systems
Shrubland, Steppe and Savanna Systems
Larvae; ostracods/cyclops; also red water mites, insect egg masses, algae (Franz 1971). Adult: terrestrial arthropods (mostly formicidae coleoptera, diptera) 74%; aquatic insect larvae (mostly trichoptera) 37% (Farner 1947).
Paedogenesis is unknown in this species. Earliest amphibian to breed in Pacific Northwest (Nussbaum et al. 1983).
In ID: breed February to May below 2100m; June to July above 2100m. Clutch size = 167 low elevation, 90 high elevation. Metamorphose: below 2100m, at SVL 35 to 40mm, year 1; above 2100m, at SVL 47mm, year 2 to 4 (Howard and Wallace 1985). Metamorphs: August to September (Brunson and Demaree 1951, Franz 1971). Sexually mature at SVL 50mm.
- Literature Cited AboveLegend: View Online Publication
- Beneski, J.T. Jr., E.J. Zalisko and J.H. Larsen Jr. 1986. Demography and migratory patterns of the eastern long-toed salamander, Ambystoma macrodactylum columbianum. Copeia 1986(2): 398-408.
- Brunson, R.B. and H.A. Demaree, Jr. 1951. The herpetology of the Mission Mountains, Montana. Copeia (4):306-308.
- Farner, D.S. 1947. Notes on the food habits of the salamanders of Crater Lake, Oregon. Copeia 1947(4): 259-261.
- Franz, R. 1971. Notes on the distribution and ecology of the herpetofauna of northwestern Montana. Bulletin of the Maryland Herpetological Society 7: 1-10.
- Howard, J.H. and R.L. Wallace. 1985. Life history characteristics of populations of the long-toed salamander (Ambystoma macrodactylum) from different altitudes. American Midland Naturalist 113(2): 361-373.
- Nussbaum, R.A., E.D. Brodie, Jr. and R.M. Storm. 1983. Amphibians and reptiles of the Pacific Northwest. University of Idaho Press. Moscow, ID. 332 pp.
- Additional ReferencesLegend: View Online Publication
Do you know of a citation we're missing?
- [NDTI] Northrop, Devine, and Tarbell Incorporated. 1994. Cabinet Gorge and Noxon Rapids hydroelectric developments 1993 wildlife study. Northrop, Devine, and Tarbell Incorporated, Portland, ME. 197 p.
- [WWPC] Washington Water Power Company. 1995. 1994 wildlife report Noxon Rapids and Cabinet Gorge Reservoirs. Washington Water Power Company. Spokane, WA.
- Adams, M.J., B.R. Hossack, and R.A. Knapp. 2002. Distribution patterns of lentic-breeding amphibians in relation to ultraviolet radiation in western North America. Northwestern Naturalist 83:62.
- Adams, M.J., B.R. Hossack, and R.A. Knapp. 2005. Distribution patterns of lentic-breeding amphibians in relation to ultraviolet radiation in western North America. Ecosystems 8(5):488-500.
- Anderson, A.R. and J.W. Petranka. 2003. Odonate predator does not affect hatching time or morphology of embryos of two amphibians. Journal of Herpetology 37(1):65-71.
- Anderson, J.D. 1961. The courtship behavior of Ambystoma macrodactylum croceum. Copeia 1961(2): 132-139.
- Anderson, J.D. 1963. Reactions of the western mole to skin secretions of Ambystoma macrodactylum croceum. Herpetologica 19(4): 282-284.
- Anderson, J.D. 1967. A comparison of life histories of coastal and montane populations of Ambystoma macrodactylum in California. American Midland Naturalist 77(2): 323-355.
- Anderson, J.D. 1972. Behavior of three subspecies of Ambystoma macrodactylum in a soil moisture gradient. Journal of Herpetology 6(3-4): 191-194.
- Anderson, J.D. 1972. Phototactic behavior of larvae and adults of two subspecies of Ambystoma macrodactylum. Herpetologica 28(3):222-226.
- Anderson, J.D. 1968a. Thermal histories of two populations of Ambystoma macrodactylum. Herpetologica 24(1): 29-35.
- Anderson, J.D. 1968b. A comparison of the food habits of Ambystoma macrodactylum sigillatum, Ambystoma macrodactylum croceum and Ambystoma tigrinum californiense. Herpetologica 24(4): 273-284.
- Anderson, J.D. and R.E. Graham. 1967. Vertical migration and stratification of larval Ambystoma. Copeia 2:371-374.
- Anderson, M.E. 1977. Aspects of the ecology of two sympatric species of Thamnophis and heavy metal accumulation with the species. M.S. thesis, University of Montana, Missoula. 147 pp.
- Bailey, J.L. 1948. Supplementary observations on the geographic variation of Ambystoma macrodactylum. Herpetologica 4: 171-174.
- Belden, L.K., E.L. Wildy, and A.R. Balustein. 2000a. Growth, survival and behaviour of larval long-toed salamanders (Ambystoma macrodactylum) exposed to ambient levels of UV-B radiation. Journal of Zoology : Proceedings of the Zoological 251(4): 473.
- Beneski, J.T., Jr., J.H. Larsen Jr., and B.T. Miller. 1995. Variation in the feeding kinematics of mole salamanders (Ambystomatidae: Ambystoma). Canadian Journal of Zoology 73(2): 353-366.
- Berner, N.J. and R.L. Ingermann. 1988. Interaction of exogenous proteins with the jelly coat of eggs from the salamander, Ambystoma macrodactylum. Comparative Biochemistry and Physiology 90A(2): 265-268.
- Berner, N.J. and R.L. Ingermann. 1990. Role of sialic acid in exogenous protein accumulation and water retention by the egg jelly of the salamander Ambystoma macrodactylum. Journal of Experimental Zoology 256(1): 38-43.
- Blackwell, E.A., R.A. Angus, G.R. Cline, and K.R. Marion. 2003. Natural growth rates of Ambystoma maculatum in Alabama. Journal of Herpetology 37(3):608-612.
- Blaustein, A.R., E.L. Wildy, L.K. Beldan, and A. Hatch. 2001. Influence of abiotic and biotic factors on amphibians in ephemeral ponds with special reference to long-toed salamanders (Ambystoma macrodactylum). Israel Journal of Zoology 47(4):333-345.
- Blaustein, A.R., J.J. Beatty, H. Deanna, and R.M. Storm. 1995. The biology of amphibians and reptiles in old-growth forests in the Pacific Northwest. General Technical Report PNW-GTR-337. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 98 p.
- Blaustein, A.R., J.M. Keisecker, D.P. Chivers, and R.G. Anthony. 1997. Ambient UV-B radiation causes deformities in amphibian embryos. Proceedings of the National Academy of Science USA 94: 13735-13737.
- Boone, M.D., E.E. Little, and R.D. Semlitsch. 2004. Overwintered bullfrog tadpoles negatively affect salamanders and anurans in native amphibian communities. Copeia 3:683-690.
- Boundy, J. 2001. Herpetofaunal surveys in the Clark Fork Valley region, Montana. Herpetological Natural History 8: 15-26.
- Bradford, D.F., and M.S. Gordon. 1992. Aquatic amphibians in the Sierra Nevada: current status and potential effects of acidic deposition on populations. Final Report, Contract No. A932-139. California Air Resources Board. Sacramento, CA.
- Bradford, D.F., C. Swanson, and M.S. Gordon. 1994. Effects of low pH and aluminum on amphibians at high elevation in Sierra Nevada, California. Canadian Journal of Zoology 72: 1272-1279.
- Branch, L.C. and R. Altig. 1981. Nocturnal stratification of three species of Ambystoma larvae. Copeia 1981(4): 870-873.
- Brodie, E.D., Jr. 1977. Salamander antipredator postures. Copeia 1977(3): 523-535.
- Brodman, R. 1993. The efffect of acidity on interactions of Ambystoma salamander larvae. Journal of Freshwater Ecology 8(3): 209-214.
- Brodman, R. 2004. Intraguild predation on congeners affects size, aggression, and survival among Ambystoma salamander larvae. Journal of Herpetology 38(1):21-26.
- Brodman, R., J. Ogger, M. Kolaczyk, R.A. Pulver, A.J. Long, T. Bogard. 2003. Mosquito control by pond-breeding salamander larvae. Herpetologocal Review 34(2):116-119.
- Brunson, R.B. 1955. Check list of the amphibians and reptiles of Montana. Proceedings of the Montana Academy of Sciences 15: 27-29.
- Bull, E.L., J.W. Deal, and J.E. Hohmann. 2001. Avian and amphibian use of fenced and unfenced stock ponds in northeastern Oregon forests. US Forest Service Research Paper PNW 539: 1-9.
- Calfee, R.D., C.M. Bridges, and E.E. Little. 2006. Sensitivity of two salamaners (Ambystoma) species to ultraviolet radiation. Journal of Herpetology 40(1):35-42.
- Carl, G.C. 1942. The long-toed salamander on Vancouver Island. Copeia 1942(1): 56.
- Carl, G.C. and I.M. Cowan. 1945. Notes on the salamanders of British Columbia. Copeia 1945(1): 43-44.
- Chivers, D.P., E.L. Wildy, and A.R. Blaustein. 1997. Eastern long-toed salamander (Ambystoma macrodactylum columbianum) larvae recognize cannabilistic conspecifics. Ethology 103: 187-197.
- Chivers, D.P., J.M. Kiesecker, M.T. Anderson, E.L. Wildy, and A.R. Blaustein. 1996. Avoidance response of a terrestrial salamander (Ambystoma macrodactylum) to chemical alarm cues. Journal of Chemical Ecology 22(9): 1709-1716.
- Clark, K.L. 1985. Responses of spotted salamander (Ambystoma maculatum) populations in central Ontario (Candada) to habitat acidity. Canadian Field Naturalist 100(4): 463-469.
- Cook, R. and K. Boland. 2005. A comparison of approaches to counting spotted salamander (Ambystoma maculatum) egg masses in vernal ponds. Herpetological Review 36(3):272-274.
- Cope, E.D. 1867. A review of the species of the Amblystomidae. Proceedings of the Academy of Natural Sciences of Philadelphia 19: 166-211.
- Cope, E.D. 1875. Check-list of North American Batrachia and Reptilia; with a systematic list of the higher groups, and an essay on geographical distribution. Based on the specimens contained in the U.S. National Museum. U.S. Natioanl Museum Bulletin 1: 1-104.
- Cope, E.D. 1889. The Batrachia of North America. Bulletin of the U.S. National Museum 34: 1-525, figs. 1-119, pls. 1-86.
- Corn, P.S., B.R. Hossack, E. Muths, D.A. Palta, C.R. Peterson, and A.L. Gallant. 2005. Status of amphibians on the Continental Divide: aurveys on a transect from Montana to Colorado, USA. Alytes 22(3-4):85-94.
- Crother, B.I. (ed.) 2008. Scientific and standard English names of amphibians and reptiles of North America north of Mexico. SSAR Herpetological Circular No. 37:1-84.
- Cunnington, D.C. and R.J. Brooks. 2000. Optimal egg size theory: Does predation by fish affect egg size in Ambytroma maculatum? Journal of Herpetology 34(1):46-53.
- Dempster, W.T. 1930. The growth of larvae of Ambystoma maculatum under natural conditions. Biology Bulletin 58: 182-192.
- Dougherty, C.K., D.A. Vaala, and G.R. Smith. 2005. Within-pond oviposition site selection in two spring-breeding amphibians (Ambystoma maculatum and Rana sylvatica). Journal of Freshwater Ecology 20(4):781-782.
- Ducey, P.K. 1989. Agonistic behavior and biting during intraspecific encounters in Ambystoma salamanders. Herpetologica 45(2): 155-160.
- Dunlap, W.W. and F.W. Allendorf. No Date. Population structure of the long-toed salamander. Unpublished report. University of Montana. Missoula, MT. 24 p. + 8 figs.
- Efford, I.E. and J.A. Mathias. 1969. A comparison of two salamander populations in Marion Lake, British Columbia. Copeia 1969: 723-735.
- Faccio, S.D. 2003. Postbreeding emigration and habitat use bu Jefferson and Spotted Salamanders in Vermont. Journal of Herpetology 37(3):479-489.
- Farmer, P. and S.B. Heath. 1987. Wildlife baseline inventory, Rock Creek study area, Sanders County, Montana. Western Technology and Engineering, Inc. Helena, MT.
- Farmer, Patrick J., and Thomas W. Butts, Western Technology & Eng., Inc., Helena, MT., 1994, McDonald Project Terrestrial Wildlife Study, November 1989 - November 1993. April 1994. In McDonald Gold Project: Wildlife & Fisheries. [#18]. Seven-up Pete Joint Venture, Lincoln, MT. Unpub. No date.
- Ferguson, D.E. 1961. The geographic variation of Ambystoma macrodactylum Baird, with the description of two new subspecies. American Midland Naturalist 65: 311-338.
- Ferguson, D.E. 1963. Ambystoma macrodactylum. Catalogue of American Amphibians and Reptiles 4.1-4.2.
- Figiel, C.R. Jr. and R.D. Semlitsch. 1990. Population variation in survival and metamorphosis of larval salamanders (Ambystoma maculatum) in the presence and absence of fish predation. Copeia 1990(3): 818-826.
- Fukumoto, J.M. 1995. Long-toed salamander (Ambystoma macrodactylum) ecology and mangement in Waterton Lakes National Park. M.S. Thesis. Faculty of Environmental Design, University of Calgary, Calgary, Alberta.
- Fukumoto, J.M. and S. Herrero. 1998. Observations of the long-toed salamander, Ambystoma macrodactylum, in Waterton Lakes National Park, Alberta. TCanadian Field Naturalist 112(4): 579-585.
- Funk, W.C. and W.W. Dunlap. 1999. Colonization of high-elevation lakes by long-toed salamanders (Ambystoma macrodactylum) after the extinction of introduced trout populations. Canadian Journal of Zoology 77: 1759-1767.
- Funk, W.C., D.A. Tallmon, and F.W. Allendorf. 1999. Small effective population size in the long-toed salamander. Molecular Ecology 8: 1633-1640.
- Gibson, J.D. and D.A. Merkle. 2005. Ambystoma maculatum (Spotted Salamander). Reproduction. Herpetological Review 36:294.
- Graham, K.L. 1997. Habitat use of long-toed salamanders (Ambystoma macrodactylum) at three different scales. M.S. Thesis, University of Guelph, Guelph, ON. 71pp.
- Graham, K.L. and G.L. Powell. 1999. Status of the Long-toed salamander (Ambystoma macrodactylum) in Alberta. Alberta Wildlife Status Report No. 22., Edmonton, AB. 19p.
- Grant, E.H.C. and P. Nanjappa. 2006. Adressing errors in identification of Ambystoma maculatum (spotted salamanders) using spot patterns. Herpetological Review 37(1):57-60.
- Greer, K.R. 1955. Yearly food habits of the river otter in the Thompson Lakes region, northwest Montana. Am. Midl. Nat. 54(2):299-313.
- Hamilton, I.M., K.L. Graham, L. Powell, and A. Russell. 1996. The range of long-toed salamanders in northwestern Alberta. Alberta Environmental Protection, Fish and Wildlife Service, Edmonton, AB. 186 pp.
- Hendricks, P. 1997. Lee Metcalf National Wildlife Refuge preliminary amphibian and reptile investigations: 1996. Montana Natural Heritage Program, Helena, MT. 21 p.
- Hendricks, P. 2000. Amphibian and reptile survey of the Thompson Chain of Lakes. A report to the Montana Department of Fish, Wildlife, and Parks. Montana Natural Heritage Program, Helena, MT. 15 p.
- Hendricks, P. and J.D. Reichel. 1996. Amphibian and reptile survey of the Bitterroot National Forest: 1995. Montana Natural Heritage Program. Helena, MT. 95 p.
- Hilliard, J., H. Minkus, and M. Weber. 1997. Amphibian survey of the Birch Creek drainage, Beaverhead County. Wildland Studies Project, San Francisco State University, San Francisco, CA. 12 p.
- Hoffman, R.L., G.L. Larson, and B.J. Brokes. 2003. Habitat segregation of Ambystoma gracile and Ambystoma macrodactylum in mountain ponds and lakes, Mt. Ranier National Park, Washington, USA. Journal of Herpetology 37(1): 24-34.
- Homan, R.N., J.M. Reed, and B.S. Windmiller. 2005. Analysis of spotted salamander (Ambystroma maculatum) growth rates based on long-bone growth rings. Journal of Herpetology 37(3):617-621.
- Hossack, B. 2002. Ambystoma macrodactylum krausei (northern long-toed salamander) vocalization. Herpetological Review 33(2): 121.
- Hossack, B., D. Pilliod, and P.S. Corn. 2001b. Preliminary amphibian surveys of the National Bison Range, Lost Trail National Wildife Refuge, and Swan River National Wildlife Refuge: 2001. USGS Northern Rocky Mountain Science Center, Aldo Leopold Wilderness Research Institute, Missoula, MT. 15 p.
- Hossack, B.R. 2006. Amphibians and wildfire in the U.S. Northwest. International Journal of Wilderness 12(1):26.
- Hossack, B.R. and P.S. Corn. 2004. Responses of pond-breeding amphibians to wildfire in Glacier National Park. Abstract. Northwestern Naturalist 85:78.
- Howard, J.H. 1979. Ecological, genetic and physiological variation in populations of the long-toed salamander (Ambystoma macrodactylum columbianum) from different altitudes. Ph.D. dissertation, University of Idaho. Moscow, Idaho. 82pp.
- Howard, J.H. and J.R. Stauffer. 1983. Critical thermal maxima in populations of Ambystoma macrodactylum from different elevations. Journal of Herpetology 17(4): 402-404.
- Howard, J.H. and R.L. Wallace. 1980. Effects of altitude on selected blood parameters in populations of the salamander Ambystoma macrodactylum. Comparative Biochemistry and Physiology 65A(2): 243-245.
- Howard, J.H. and R.L. Wallace. 1981. Microgeographic variation of electrophoretic loci in populations of Ambystoma macrodactylum columbianum (Caudata: Ambystomatidae). Copeia 1981(2): 466-471.
- Husting, E.L. 1965. Survival and breeding structure in a population of Ambystoma maculatum. Copeia 1965: 352-362.
- Ingermann, R.L., D.C. Bencic, and P. Verrell. 2002. Methoxychlor alters the predator-prey relationship between dragonfly naiads and salamander larvae. Bulletin of Environmental Contamination and Toxicology 68(6):771-778.
- Johnson, P.T.J., K.B. Lunde, E.M. Thurman, E.G. Ritchie, S.N. Wray, D.R. Sutherland, J.M. Kapfer, T.J. Frest, J. Bowerman, and A.R. Blaustein. 2002. Parasite (Ribeiroia ondatrae) infection linked to amphibian malformations in the western United States. Ecological Monographs 72(2):151-168.
- Kezer, J. and D.S. Farner. 1955. Life history patterns of the salamander Ambystoma macrodactylum in the high Cascade Mountains of southern Oregon. Copeia 1955(2): 127-131.
- Knudsen, J.W. 1960. The courtship and egg mass of Ambystoma gracile and Ambystoma macrodactylum. Copeia 1960(1): 44-46.
- Kraus, F. 1988. An empirical evaluation of the use of the ontogeny polarization criterion in phylogenetic inference. Systematic Zoology 37(2): 106-141.
- Kutka, F.J. 1994. Low pH effects of swimming activity of Ambystoma salamander larvae. Environment, Toxicology and Chemistry 13(11): 1821-1824.
- L. L. C. Jones, W. P. Leonard and D. H. Olson, eds. 2005. Amphibians of the Pacific Northwest. Seattle Audubon Society: Seattle, WA, 227 pp.
- Laselle, B.T. 2000. Association of wetland area with breeding activity for multiple amphibian species. Undergraduate Honors Thesis. Carroll College, Helena, MT. 18 p.
- Leonard, W.P., and K.O. Richter. 1994. Western long-toed salamander demographics and oviposition in a small vernal wetland of the Puget Sound lowlands. Northwest Science 68(2): 135.
- Manville, R.H. 1957. Amphibians and reptiles of Glacier National Park, Montana. Copeia 1957: 308-309.
- Marnell, L. E. 1997. Herpetofauna of Glacier National Park. Northwestern Naturalist 78:17-33.
- Marnell, L.F. 1996. Amphibian survey of Glacier National Park, Montana. Abstract. Intermountain Journal of Sciences 2(2): 52.
- Maxell, B. A. 2000. Management of Montana's amphibians: a review of factors that may present a risk to population viability and accounts on the identification, distribution, taxonomy, habitat use, natural history, and the status and conservation of individual species. Report to USFS Region 1, Order Number 43-0343-0-0224. University of Montana, Wildlife Biology Program. Missoula, MT. 161 p.
- Maxell, B.A. 2002a. Amphibian and aquatic reptile inventories in watersheds in the South and Middle Forks of the Flathead River drainage that contain lakes being considered for application of piscicides and subsequent stocking of west slope cutthroat trout. Report to the Region 1 Office of the U.S. Forest Service and the Montana Department of Fish, Wildlife, and Parks. Montana Cooperative Wildlife Research Unit and Wildlife Biology Program, University of Montana, Missoula, MT. 62 pp.
- Maxell, B.A., J.K. Werner, P. Hendricks, and D.L. Flath. 2003. Herpetology in Montana: a history, status summary, checklists, dichotomous keys, accounts for native, potentially native, and exotic species, and indexed bibliography. Society for Northwestern Vertebrate Biology, Northwest Fauna Number 5. Olympia, WA. 135 p.
- McAvoy, S.A. 2003. Wetland characteristics associated with amphibian presence in the Rocky Mountain Region. Undergraduate Honors Thesis. Carroll College, Helena, MT. 23 p.
- McCoy, K.A., R.N. Harris. 2003. Integrating development stability analysis and current amphibian monitoring techniques: an experimental evaluation with the salamander Ambystoma maculatum. Herpetologica 59(1): 22-36.
- McGraw, R.L. 1997. Timber harvest effects on long-toed salamanders (Ambystoma macrodactylum). Abstract from SSAR meeting in Seattle, Washington.
- McGraw, R.L., II. 1997. Timber harvest effects on metamorphosed and larval long-toed salamanders (Ambystoma macrodactylum). M.S. Thesis. University of Montana, Missoula, MT. 74 p.
- McLaughlin, G.S. 1999. Draft revised recovery plan for the Santa Cruz long-toed salamander. U.S. Fish and Wildlife Service Region 1. Portland, Oregon.
- Menninger, K.L. 2004. The effects of hydroperiod on the head morphology of long-toed salamanders (Ambystoma macrodactylum). Undergraduate Honors Thesis. Carroll College, Helena, MT. 20 p.
- Miller, J. D. 1975. Interspecific food relationships of anurans in northwestern Montana and fluoride accumulation in amphibians and reptiles in northwestern Montana. M.S. thesis. University of Montana, Missoula, MT. 105 p.
- Miller, M. 1995. Amphibian survey Birch Creek July 1995. Wildland Studies Project, San Francisco State University, San Francisco, CA. 9 p.
- Miller, M. 1995. Amphibians survey, Birch Creek, July 1995. Unpublished report. University of California, Berkley. 3 pp.
- Mittleman, M.B. 1948, American Caudata II. Geographic variation in Ambystoma macrodactylum. Herpetologica 4: 81-95.
- Monello, R.J. and R.G. Wright. 2001. Predation by goldfish (Carassius auratus) on eggs and larvae of the eastern long-toed salamander (Ambystoma macrodactylum columbianum). Journal of Herpetology 35(2): 350-353.
- Naughton, G.P., C.B. Henderson, K.R. Foresman, and R.L. McGraw, II. 2000. Long-toed salamanders in harvested and intact Douglas-fir forests of western Montana. Ecological Applications 10(6): 1681-1689.
- Nebeker, A.V., G.S. Schuytema, W.L. Griffis, and A. Cataldo. 1998. Impact of guthion on survival and growth of the frog Psuedacris regilla and the salamanders Ambystoma gracile and Ambystoma maculatum. Archives of Environmental Contamination and Toxicology 35(1):48-51.
- Nelson, S.J., G.L. Powell, and A.P. Russell. 1995. Population survey of the long-toed salamander (Ambystoma macrodactylum) in southwestern Alberta. Alberta Environmental Protection, Fish and Wildlife Service, Edmonton, AB. 161pp.
- Oseen, K.L., G.L. Powell, and A.P. Russell. 1995b. The distribution of the long-toed salamander (Ambystoma macrodactylum) in northwestern Alberta. Unpublished report submitted to Alberta Fish and Wildlife, Environmental Protection, Edmonton, Alberta. p
- Palen, W.J., C.E. Williamson, A.A. Clauser, and D.E. Schindler. 2005. Impact of UV-B exposure on amphibian embryos: linking species physiology and oviposition behaviour. Proceedings of the Royal Society Biological Sciences Series B 272(1569):1227-1234.
- Pearman, P.B. 2002. Interactions between Ambystoma salamander larvae: evidence for competitive asymmetry. Herpetologica 58(2):156-165.
- Pearson, K.J. 2004. Long-toed salamander (Ambystoma macrodactylum) Alberta Species at Risk Report 90:136-147.
- Peters, W. 1882. Legte drei neue Batrachier (Amblystoma krausei, Nyctibatrachus sinensis, Bufo buchneri) vor. Sitzungs - bericht der Gesellschaft Naturforschender Freunde zu Berlin 10: 145-148.
- Powell, G.L., A.P. Russell, J.D. James, S.J. Nelson, and S.M. Watson. 1997. Population biology of the long-toed salamander (Ambystoma macrodactylum) in the front range of Alberta. In: D.M. Green (Ed.). Amphibians in decline: Canadian studies of a glob
- Powell, G.L., A.P. Russell, S.J. Nelson, I.M. Hamilton, and K.L. Graham 1997. The status of the Long-toed salamander (Ambystoma macrodactylum) in Alberta. Alberta Environmental Protection, Fish and Wildlife Division, Edmonton, AB.
- Powell, G.L., S.J. Nelson, and A.P. Russell. 1993. The Bow Valley long-toed salamander population study: a preliminary report on the 1992 field season. Alberta Environmental Protection, Fish and Wildlife Service, Edmonton, AB. 124 pp.
- Reed, R.J. 1978. Population study of the Santa-Cruz long-toed salamander (Ambystoma marcrodactylum croceum) at Valencia Lagoon 1977-1978 with notes on habitat and occurrence in Santa Cruz and Monterey countites. California Department of Fish and Game R
- Reed, R.J. 1980. The population dynamics of Ambystoma macrodactylum croceum as a response to climatological variables. American Zoologist 20(4): 879.
- Reichel, J. and D. Flath. 1995. Identification of Montana's amphibians and reptiles. Montana Outdoors 26(3):15-34.
- Reichel, J.D. 1996. Preliminary amphibian and reptile survey of the Helena National Forest: 1995. Montana Natural Heritage Program. Helena, MT. 87 pp.
- Reichel, J.D. 1997. Amphibian, reptile and northern bog lemming survey on the Rocky Mountain Front: 1996. Montana Natural Heritage Program, Helena, MT. 81 p.
- Reichel, J.D. 1995. Preliminary amphibian and reptile survey of the Lewis and Clark National Forest: 1994. Montana Natural Heritage Program. Helena, MT. 92 p.
- Reichel, J.D. and S.G. Beckstrom. 1993. Northern bog lemming survey: 1992. Unpublished report. Montana Natural Heritage Program, Helena, MT. 64 pp.
- Rodgers, T. L. and W. L. Jellison. 1942. A collection of amphibians and reptiles from western Montana. Copeia (1):10-13.
- Roedel, M.D. and P. Hendricks. 1998. Amphibian and reptile survey on the Bureau of Land Management Lewistown District: 1995-1998. Montana Natural Heritage Program, Helena, MT. 53 p.
- Roedel, M.D. and P. Hendricks. 1998b. Amphibian and reptile inventory on the Headwaters and Dillon Resource Areas in conjunction with Red Rocks Lakes National Wildlife Refuge: 1996-1998. Montana Natural Heritage Program, Helena, MT. 46 p.
- Russell, A. P. and A. M. Bauer. 1993. The amphibians and reptiles of Alberta. University of Calgary Press. Calgary, Alberta. 264 p.
- Russell, A.P., G.L. Powell, and D.R. Hall. 1996. Growth and age of Alberta long-toed salamanders (Ambystoma macrodactylum krausei): a comparison of two methods of estimation. Canadian Journal of Zoology 74: 397-412.
- Russell, R.W. and J.D. Anderson. 1956. A disjunct population of the long-toed salamander from the coast of California. Herpetologica 12: 137-140.
- Ruth, S.B. 1988. The life history and current status of the Santa Cruz long-toed salamander (Ambystoma macrodactylum croceum). Pages 89-110 in H.F. Lisle, P.R. Brown, B.Kaufman, and B.M. McGurty, editors. Proceedings of the Conference on California He
- Ruth, S.B. and K. Tollestrup. 1973. Aspects of the life history and current status of the Santa Cruz long-toed salamander (Ambystoma macrodactylum croceum) at Valencia Lagoon, Santa Cruz County, California. Report for California Division of Highways. 5
- Sage, R.D. Submitted to Conservation Biology. Genetic variability and biogeographical history: implications for the conservation of a relictual salamander (Ambystoma macrodactylum croceum). Conservation Biology (Submitted in 1999-Apparently rejected).
- Schmidt, K.P. 1953. A checklist of North American Amphibians and Reptiles. Sixth edition. American Society of Icthyologists and Herpetologists. 280 p.
- Sessions, S.K. and S.B. Ruth. 1990. Explanation for naturally occurring supernumerary limbs in amphibians. Journal of Experimental Zoology 254(1): 38-47.
- Sheppard R.F. 1977. The ecology and home range movements of Ambystoma macrodactylum krausei (Amphibia : Urodela). M.S. Thesis, University of Calgary. 138pp.
- Sherratt, J. 2003. The effects of hydroperiod on the morphology of the long-toed salamanser Ambystoma macrodatylum. Undergraduate Honors Thesis. Carroll College, Helena, MT. 26 p.
- Shoop, C.R. 1968. Migratory orientation of Ambystoma maculatum: movements near breeding ponds and displacements of migrating individuals. Biology Bulletin 135: 230-238.
- Slater, J.R. 1936. Notes on Ambystoma gracile Baird and Ambystoma macrodactylum Baird. Copeia 1936(4): 234-236.
- Spohr, S.M. 2005. Ambystoma maculatum (Spotted Salamander). Predation. Herpetological Review 36:294-295.
- Stebbins, R. C. 2003. A field guide to western reptiles and amphibians. 3rd Edition. Houghton Mifflin Company, Boston and New York. 533 p.
- Talent, L.G. and C.L. Talent. 1980. A population of the endangered Santa Cruz long-toed salamander, Ambystoma macrodactylum croceum, from Monterey County, California. California Fish and Game 66(3):184-186.
- Tallmon, D.T., W.C. Funk, W.W. Dunlap, and F.W. Allendorf. 2000. Genetic differentiation of long-toed salamanders (Ambystoma macrodactylum) populations. Copeia 2000(1): 27-35.
- Test, F.C. 1893. Annotated list of reptiles and batrachians collected. In B.W. Evermann. A reconnaisance of the streams and lakes of western Montana and northwestern Wyoming. Bulletin of United States Fish Commission 11(1891): 57-59.
- Thompson, M.D. 2000. Long-toed phylogeography a 'neo-perspective' on declines. The Boreal Dip Net 5(1): 14-16.
- Thompson, M.D. 2001. An unusually adept ambystomatid, the long toed salamander, coping at northern extremes. The Boreal Dip Net. 5(2):8-10.
- Thompson, M.D. and A.P. Russell. 2000. Phylogeography of Ambystoma macrodactylum: post glacial range expansion and resultant genetic diversity. Field Summary Report No. 1. University of Calgary. Calgary, Canada. 39 p.
- Thompson, M.D. and A.P. Russell. 2001. Phylogeography of Ambystoma macrodactylum: post glacial range expansion and resultant genetic diversity. Field Summary Report No. 2. University of Calgary, Calgary, Canada. 65 p.
- Thompson, Richard W., Western Resource Dev. Corp., Boulder, CO., 1996, Wildlife baseline report for the Montana [Montanore] Project, Lincoln and Sanders counties, Montana. In Application for a Hard Rock Operating Permit and Proposed Plan of Operation, Montanore Project, Lincoln and Sanders Counties, Montana. Vol. 5. Stroiazzo, John. Noranda Minerals Corp., Libby, MT. Revised September 1996.
- Thoms, C., and C.C. Corkran. 1994. Identifying Oregon amphibians in the field: distinctions between long-toed and northwestern salamanders. Northwest Science 68(2): 154.
- Tihen, J.A. 1969. Ambystoma. Catalogue of American Amphibians and Reptiles 75.1-75.4.
- Timken, R. No Date. Amphibians and reptiles of the Beaverhead National Forest. Western Montana College, Dillon, MT. 16 p.
- Turtle, S.L. 2000. Embryonic survivorship of the spotted salamander (Ambystoma maculatum) in roadside and woodland vernal pools in southeastern New Hampshire. Journal of Herpetology 34(1): 60-67.
- Tyler, T., W.J. Liss, L.M. Ganio, G.L. Larson, R. Hoffman, E. Deimling, and G. Lomnicky. 1998a. Interaction between introduced trout and larval salamanders (Ambystoma macrodactylum) in high elevation lakes. Conservation Biology 12: 94-105.
- Tyler, T.J. 1996. Interactions between stocked trout larval salamanders (Ambystoma macrodactylum) in high-elevation lakes. Masters Thesis, Oregon State University. 55p.
- Tyler, T.J., W.J. Liss, R.L. Hoffman, and L.M. Ganio. 1998. Experimental analysis of trout effects on survival, growth, and habitat use of two species of Ambystomid salamanders. Journal of Herpetology 32(3): 345-349.
- Vasconcelos, D. and A.J.K. Calhoun. 2004. Movement patterns of adult and juvenile Rana sylvatica (LeConte) and Ambystoma maculatum (Shaw) in three restored seasonal pools in Maine. Journal of Herpetology 38(4):551-561.
- Verrell, P. and J. Pelton. 1996. The sexual strategy of the central long-toed salamander, Ambystoma macrodactylum columbianum, in south-eastern Washington. Journal of the Zoological Society of London 240: 37-50.
- Verrell, P. and K. Davis. 2003. Do non-breeding, adult long-toed salamanders respond to conspecifics as friends or as foes? Herpetologica 59(1): 1-7.
- Verrell, P., N. Strand, and E.Hanson. Sexual dimorphism of mate location in the long-toed salamander Ambystoma macrodactylum columbianum. Ethology 107(8): 677-684.
- Verrell, P.A. 2004. The male reproductive cycle of the North American salamander Ambystoma macrodactylum columbianum. Amphibia-Reptilia 25(3):349-356.
- Walker, R.S. 1984. Distribution of the northern long-toed salamander (Ambystoma macrodactylum) east of the continental divide in Montana. Honors thesis, Carroll College. Helena, MT. 37 pp.
- Walls, S.C. and A.R. Blaustein. 1994. Does kinship influence density dependence in a larval salamander? Oikos 71(3):459-468.
- Walls, S.C., J.J. Beatty, B.N. Tissot, D.G. Hokit, and A.R. Blaustein. 1993. Morphological variation and cannibalism in a larval salamander (Ambystoma macrodactylum columbianum). Canadian Journal of Zoology 71: 1543-1551.
- Walls, S.C., S.S. Belanger, and A.R. Blaustein. 1993. Morphological variation in a larval salamander: dietary induction of plasticity in head shape. Oecologica 96:162-168.
- Walsh, R. 1998. An extension of the known range of the long-toed salamander, Ambystoma macrodactylum, in Alberta. Canadian Field Naturalist 112: 331-333.
- Watson S. 1997. Food level effects on metamorphic timing in the long-toed salamander, Ambystoma macrodactylum krausei. Masters Thesis, Univesisty of Calgary. pp. 206.
- Weisel, G.F. 1952. Animal names, anatomical terms, and some ethnozoology of the Flathead Indians. Journal of the Washington Academy of Sciences 42(11): 345-355.
- Werner, J.K. and J.D. Reichel. 1996. Amphibian and reptile monitoring/survey of the Kootenai National Forest: 1995. Montana Natural Heritage Program. Helena, MT. 115 pp.
- Werner, J.K. and J.D. Reichel. 1994. Amphibian and reptile survey of the Kootenai National Forest: 1994. Montana Natural Heritage Program. Helena, MT. 104 p.
- Werner, J.K. and T. Plummer. 1995. Amphibian and reptile survey of the Flathead Indian Reservation 1993-1994. Salish Kootenai College, Pablo, MT. 55 pp.
- Werner, J.K. and T. Plummer. 1995. Amphibian monitoring program on the Flathead Indian Reservation 1995. Salish Kootenai College, Pablo, MT. 46 p.
- Werner, J.K., B.A. Maxell, P. Hendricks and D.L. Flath. 2004. Amphibians and Reptiles of Montana. Mountain Press Publishing Company: Missoula, MT. 262 pp.
- Werner, J.K., T. Plummer, and J. Weaselhead. 1998b. The status of amphibians on the Flathead Reservation, Montana. Intermountain Journal of Sciences 4(3-4): 88.
- Werner, J.K., T. Plummer, and J. Weaslehead. 1998. Amphibians and reptiles of the Flathead Indian Reservation. Intermountain Journal of Sciences 4(1-2): 33-49.
- Wildy, E.L., and A.R. Blaustein. 2001. Learned recognition of intraspecific predators in larval long-toed salamanders Ambystoma macrodactylum. Ethology 107(6): 479-494.
- Wildy, E.L., D.P. Chivers, and A.R. Blaustein. 1999. Shifts in life history trairs as a response to cannibalism in larval long-toed salamanders (Ambystoma macrodactylum). Journal of Chemical Ecology 25: 2337-2346.
- Wildy, E.L., D.P. Chivers, J.M. Kiesecker, and A.R. Blaustein. 1998. Cannibalism enhances growth in larval long-toed salamanders, (Ambystoma macrodactylum). Journal of Herpetology 32(2) 286-289.
- Wildy, E.L., D.P. Chivers, J.M. Kiesecker, and A.R. Blaustein. 2001. The effects of food level and conspecific density on biting and cannibalism in larval long-toed salamanders, Ambystoma macrodactylum. Oecologial 128(2): 202-209.
- Williams, T.A., and C.D. Anthony. 1994. Technique to isolate salamander granular gland products with a comment on the evolution of adhesiveness. Copeia 1994(2): 540-541.
- Williams, T.A., and J.H. Larsen, Jr. 1986. New function for the granular skin glands of the eastern long-toed salamander, Ambystoma macrodactylum columbianum. Journal of Experimental Zoology 239(3): 329-334.
- Wirsing, A.J., J.D. Roth, and D.L. Murray. 2005. Can prey use dietary cues to distinguish predators? A test involving three terrestrial amphibians. Herpetologica 61(2):104-110.
- Wright, A.N. and K.R. Zamudio. 2002. Color pattern asymmetry as a correlate of habitat disturbance in spotted salamanders (Ambystoma maculatum). Journal of Herpetology 36(1):129-133.
- Zackheim, Karen, 1973?, Exhibit H: Wildlife Study. In Ash Grove Cement Co. files.
- Zalisko, E.J. 1987. Morphology, histochemistry, and seasonal changes in the vasa deferentia of Rhyacotriton olympicus olympicus (Dicamptodontidae) and Ambystoma macrodactylum columbianum (Ambystomatidae), (Amphibia, Urodela). Ph.D. Dissertation. Washi
- Zalisko, E.J. and J.H. Larsen Jr. 1990. Breeding and post-breeding structure of the vas deferens of the long-toed salamander Ambystoma macrodactylum. Journal of Morphology 203(3): 321-330.
- Zalisko, E.J. and J.H. Larsen, Jr. 1989. Fate of unused sperm in post-breeding male Ambystoma macrodactylum columbianum. Journal of Herpetology 23(4): 463-464.
- Zamudio, K.R. and W.K. Savage. 2003. Historical isolation, range expansion, and secondary contact of two highly divergent mitochondrial lineages in spotted salamanders (Ambystoma maculatum). Evolution 57:1631-1652.
- Zisook, R., K. Almond, and B. Sharpe. 1996. Amphibian survey of the Birch Creek drainage, Beaverhead County. Wildland Studies Project. San Francisco State University, San Francisco, CA. 9 p.