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Montana Field Guides

Long-toed Salamander - Ambystoma macrodactylum

Image Copyright and Usage Information
Native Species

Global Rank: G5
State Rank: S5
(see State Rank Reason below)


Agency Status
USFWS:
USFS:
BLM:


 

External Links

State Rank Reason (see State Rank above)
Species is relatively common within suitable habitat and widely distributed across portions of the state
  • Details on Status Ranking and Review
    Long-toed Salamander (Ambystoma macrodactylum) Conservation Status Review
    Review Date = 05/03/2018
    Range Extent

    ScoreF - 20,000-200,000 km squared (about 8,000-80,000 square miles)

    Comment84,839 square Kilometers from Natural Heritage Program Range Maps

    Long-term Trend

    ScoreU - Unknown. Long-term trend in population, range, area occupied, or number or condition of occurrences unknown

    CommentUnknown, little historic data are available. Habitat has remained stable since European arrival

    Short-term Trend

    ScoreE - Stable. Population, range, area occupied, and/or number or condition of occurrences unchanged or remaining within ±10% fluctuation

    CommentGiven consistent detections during lentic surveys across the species range, occupancy of breeding sites is likely stable within 10%

    Threats

    ScoreH - Unthreatened. Threats if any, when considered in comparison with natural fluctuation and change, are minimal or very localized, not leading to significant loss or degradation of populations or area even over a few decades’ time. (Severity, scope, and/or immediacy of threat considered Insignificant.)

    CommentNo operational or potential threats are likely to impact the species in the future

    Intrinsic Vulnerability

    ScoreC - Not Intrinsically Vulnerable. Species matures quickly, reproduces frequently, and/or has high fecundity such that populations recover quickly (< 5 years or 2 generations) from decreases in abundance; or species has high dispersal capability such that extirpated populations soon become reestablished through natural recolonization (unaided by humans).

    CommentSpecies is mature in 1-2 years, produces thousands of eggs.

    Environmental Specificity

    ScoreC - Moderate. Generalist. Broad-scale or diverse (general) habitat(s) or other abiotic and/or biotic factors are used or required by the species but some key requirements are scarce in the generalized range of the species within the area of interest.

    CommentSpecies is found in forested habitats across western and some of central Montana

    Raw Conservation Status Score

    Score 3.5 + 0 (geographic distribution) + 0 (environmental specificity) + 0 (short-term trend) + 1 (threats) = 4.5

 
General Description
EGGS
Typically laid in ponds in small clusters of 9-81 (X = 23, SD = 14.5, N = 36 across 5 sites in northwest Montana). Each ovum is black or brown above, white to gray below, and surrounded by two jelly layers. Ovum diameters are approximately 2.5 mm (0.1 in), but total egg diameters, including the two jelly layers, are usually 12-17 mm (0.47-0.67 in) (Slater 1936).

LARVAE
Translucent, light tan, or black dorsally and laterally with black and gold flecks. White to
pinkish ventrally. Three pairs of external feathery gills emanate from the sides of the head with 9-13 gill rakers on their anterior surface (Russell and Bauer 2000). Snout-vent length (SVL) of 10-60 mm (0.4-2.4 in).

JUVENILES AND ADULTS
Fourth toe on the hind foot is elongate and longer than the sole of the foot. Incomplete or fully formed yellow, orangish, or reddish dorsal stripe may extend from the tip of the snout to the tip of the tail. Eyelids are the same color as the dorsal stripe. White flecking present on the lateral and ventral surfaces over a black lateral and pink ventral base color. 12-13 costal grooves are present. SVL of 25-80 mm (1-3.1 in) (Russell et al. 1996).

Diagnostic Characteristics
Adult Coeur d’Alene Salamanders (Plethodon idahoensis) have nasolabial grooves and their toes are webbed and shorter than the soles of their feet. See sections on habitat use for differences in habitat used by Long-toed and Coeur d’Alene Salamanders. Western Tiger Salamander (Ambystoma mavortium) eggs have 3 jelly layers and have total diameters less than 10 mm (0.4 in), including the jelly layers. Larval Western Tiger Salamanders have larger heads are usually olive green to silvery white in base color and have 15-25 gill rakers on the anterior surface of their gills (Russell and Bauer 2000). See sections on distribution for geographic areas of possible overlap for Long-toed and Western Tiger Salamanders.

Species Range
Montana Range Range Descriptions

Native

Western Hemisphere Range

 


Range Comments
Five subspecies are recognized and range from central California through the Pacific Northwest
to southeast Alaska at elevations up to or above 2,700 m (8,859 ft) (Ferguson 1961; Petranka
1998). Only a single subspecies, the Northern Long-toed Salamander (A. m. krausei), occurs in Montana. Their known range in Montana extends west of the Rocky Mountain Front and the Missouri, Jefferson, and Beaverhead Rivers.

Maximum elevation: 2,774 m (9,100 ft), Beaverhead County K. Storrar (Werner et al. 2004).

Observations in Montana Natural Heritage Program Database
Number of Observations: 2817

(Click on the following maps and charts to see full sized version) Map Help and Descriptions
Relative Density

Recency

 

(Observations spanning multiple months or years are excluded from time charts)



Migration
Although individual animals tend to use the same migration routes, no preference in habitat, relative soil moisture or vegetation is evident for the species’ movements to and from breeding pools (Beneski et al. 1986).

Habitat
Adults are found in a wide variety of habitats including semi-arid sagebrush, alpine meadows, dry woodlands, humid forests, rocky shores of mountain lakes, and disturbed agricultural areas (Nussbaum et al. 1983). Outside of the breeding season adults are primarily subterranean and have been documented to commonly move at least 600 m (1,969 ft) from the nearest breeding site on the University of Montana’s Lubrecht Experimental Forest (Maxell et al. 2009). During the breeding season adults may be found in shallow waters or under logs, rocks, and other debris near water.

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 (common or occasional) of each of the 82 ecological systems mapped in Montana for vertebrate animal species that regularly breed, overwinter, or migrate through the state by:
    1. 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 2012, Adams 2003, and Werner et al. 2004);
    2. Evaluating structural characteristics and distribution of each ecological system relative to the species' range and habitat requirements;
    3. Examining the observation records for each species in the state-wide point observation database associated with each ecological system;
    4. 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 listed as associated with 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 listed as 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.  Common versus occasional association with an ecological system was assigned based on the degree to which the structural characteristics of an ecological system matched the preferred structural habitat characteristics for each species as represented in scientific 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 assignment of common versus occasional association.  If you have any questions or comments on species associations with ecological systems, please contact the Montana Natural Heritage Program's Senior Zoologist.

    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: mtnhp.org/requests) 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.

    Literature Cited
    • 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.  2012.  Mammals of Montana.  Second edition.  Mountain Press Publishing, Missoula, Montana.  429 pp.
    • 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.

Food Habits
Larvae and adults feed on a variety of aquatic and terrestrial invertebrates and larvae feed on other amphibian larvae including conspecifics (Farner 1947, Anderson 1968b, Walls et al. 1993, Maxell et al. 2009). Franz (1997) identified that larvae diets consisted of ostracods/cyclops; also red water mites, insect egg masses, algae. Farner (1947) determined adult diets contained terrestrial arthropods (mostly formicid coleop, diptera) 74% and aquatic insect larvae (mostly trichop) 37%.

Reproductive Characteristics
Adults go to the breeding ponds immediately after snowmelt. In the Pacific Northwest and western Montana, it is the earliest amphibian to breed (Nussbaum et al. 1983). Breeding takes place in temporary or permanent ponds or in quiet water at the edge of lakes and streams, usually those without fish present (Funk and Dunlap 1999, Monello and Wright 1999). Like all salamanders, they have internal fertilization. Following breeding, they move to adjacent uplands.

Eggs are attached to vegetation or loose on the bottom at depths up to 0.8 m (31.5 in) and hatch in 3 to 6 weeks. Paedogenesis is unknown in this species. Larvae usually transform at the end of their first summer at low elevations or at the end of their second, third or fourth summer at high elevations and in cold waters at lower elevations (Howard and Wallace 1985, Maxell et al. 2009).

In Idaho, they have been documented breeding in February to May below 2,100 m (6,890 ft) and June to July above 2,100 m (6,890 ft). Clutch sizes are typically around 167 at lower elevations and 90 at high elevations. Metamorphose below 2,100 m (6,890 ft) occurs when SVL is 35-40 mm (1.4-1.6 in) in year 1 and above 2,100 m (6,890 ft), during year 2 to 4 when SVL 47 mm (1.9 in) (Howard and Wallace 1985). Metamorphs are typically seen in August to September (Brunson and Demaree 1951, Franz 1971). Salamanders become sexually mature when SVL is 50 mm (2.0 in).

Management
The following is taken from the Status and Conservation section for the Long-toed Salamander account in Maxell et al. 2009

Long-toed Salamanders are the most widely distributed and common amphibian species west of the Continental Divide with larvae being found in most fishless standing waterbodies with adjacent soils that provide suitable terrestrial habitat (i.e., sites that are not surrounded by extensive areas of bare rock). Their status in the front ranges east of the Continental Divide is uncertain. Risk factors relevant to the viability of populations of this species are likely to include all the general risk factors described above with the exception of harvest and commerce. Individual studies that specifically identify risk factors or other issues relevant to the conservation of Long-toed Salamanders include the following. (1) A number of studies have found adverse impacts of introduced fish on Long-toed Salamanders. Funk and Dunlap (1999) found that trout effectively excluded salamander populations from lakes in the Bitterroot Mountains. However, when fish went extinct in lakes that did not have spawning habitat salamanders were able to recolonize some of them over a twenty-year time period. In the Palouse region of northern Idaho, Monello and Wright (1999) found the presence of Long-toed Salamanders to be highly negatively correlated with the presence of a variety of fish species, including Largemouth Bass (Micropterus salmoides), Bluegill (Lepomis macrochirus), Channel Catfish (Ictalurus punctatus), and Goldfish. Tyler et al. (1998a) found a similar pattern in North Cascades National Park and found that nitrogen levels were positively correlated with salamander densities in fishless lakes, apparently an indication of bottom up limitations on the food web. Similarly, Long-toed Salamanders in the central and northern portion of the Sierra Nevada Mountains are largely restricted to fishless lakes (Bradford and Gordon 1992 as cited in Knapp 1996). Tyler et al. (1998b) found that when Rainbow Trout (Oncorhynchus mykiss) were stocked in experimental ponds with Long-toed Salamanders, larval survivorship was lower and larval body lengths were smaller than in control ponds without fish, supporting the theory that introduced trout not only impact salamanders through direct predation, but also indirectly by increasing refuge use and, thereby, reducing foraging time. (2) In a study of the Long-toed Salamander in Douglas-fir (Pseudotsuga menziesii) forests in the Swan River Valley, McGraw (1997) found that areas where overstory removal (250-300 trees harvested per hectare) and new forestry (leave 13-25 dominant tree species per hectare and retain all snags and hardwoods) harvest techniques were applied had less ground cover, higher soil temperatures, and 75% fewer terrestrial salamanders than control plots. Interestingly, he also found that larvae were more abundant in ponds where a fraction of the pond margin was harvested than either ponds whose margins were completely harvested or ponds whose forest margins were completely intact. (3) Tallmon et al. (2000) found that gene flow among populations in the Bitterroot Mountains is greater between populations on the same mountain ridge than between populations on adjacent mountain ridges, indicating that drainages between ridges act as more of a barrier to dispersal than the steep terrain on the ridge itself. The dominance of terrestrial dispersal is also supported by the genetic analyses of Howard and Wallace (1981). (4) Fukumoto and Herrero (1998) documented mortality of a minimum of 1-2% of the adult breeding population as they crossed a roadway to the breeding site in Waterton Lakes National Park in Alberta. However, the authors suggest that actual mortality may have been considerably higher, contributing to the unusual 3:1 female biased sex ratio observed at the breeding site. (5) Blaustein et al. (1994d) found that the species has low levels of photolyase, an enzyme that repairs UV-B radiation damage to DNA. Blaustein et al. (1997) subsequently found that only 14.5% of embryos exposed to 94% of ambient UV-B radiation survived to hatching as compared to 95% survival for larvae exposed to only 10% of ambient UV-B radiation. Together these findings suggest that enhanced UV-B radiation from thinning of the ozone layer may be impacting salamander populations now or will impact them at some time in the future. (6) Sessions and Ruth (1990) found that cysts of a trematode parasite apparently caused limb deformities at the site of the cyst. This parasite has now been found in larvae collected in Montana (Pieter Johnson, Claremont Mckenna College, personal communication). (7) The extent of the use of larval Long-toed Salamanders as fishing bait is unknown in Montana, however the species is known to be used in large numbers in other states (Collins 1981) so this practice does have the potential to impact populations in Montana. Accidental introduction of larvae being used for bait may result in hybridization and genetic introgression, possibly leading to the elimination of distinct genetic makeups (Collins 1981). (8) Bradford et al. (1994) found that the LC50 pH for Pacific treefrog embryos and hatchlings exposed for 7 days averaged 4.3 and that pH levels greater than or equal to 5.0 had no significant lethal or sublethal effects.

References
  • Literature Cited AboveLegend:   View Online Publication
    • 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.
    • 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.
    • 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.
    • Blaustein, A.R., P.D. Hoffman, D.G. Hokit, J.M. Kiesecker, S.C. Walls, and J.B. Hays. 1994d. UV repair and resistance to solar UV-B in amphibian eggs: a link to population declines? Proceedings of the National Academy of Sciences 91: 1791-1795.
    • 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.
    • Brunson, R.B. and H.A. Demaree, Jr. 1951. The herpetology of the Mission Mountains, Montana. Copeia (4):306-308.
    • Collins, J.P. 1981. Distribution, habitats and life history variation in the tiger salamander (Ambystoma tigrinum) in east-central and southeast Arizona. Copeia 1981(3): 666-675.
    • Farner, D.S. 1947. Notes on the food habits of the salamanders of Crater Lake, Oregon. Copeia 1947(4): 259-261.
    • Ferguson, D.E. 1961. The geographic variation of Ambystoma macrodactylum Baird, with the description of two new subspecies. American Midland Naturalist 65: 311-338.
    • Franz, R. 1971. Notes on the distribution and ecology of the herpetofauna of northwestern Montana. Bulletin of the Maryland Herpetological Society 7: 1-10.
    • 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.
    • 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.
    • 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.
    • Knapp, R.A. 1996. Non-native trout in natural lakes of the Sierra Nevada: an analysis of their distribution and impacts on native aquatic biota. In: Sierra Nevada ecosystem project: final report to Congress, volume III, assessments, commissioned reports, and background information. Davis, CA: University of California Centers for Water and Wildland Resources. Wildland Resources Center Report 38. p 363-407.
    • Maxell, B.A., P. Hendricks, M.T. Gates, and S. Lenard. 2009. Montana amphibian and reptile status assessment, literature review, and conservation plan, June 2009. Montana Natural Heritage Program. Helena, MT. 643 p.
    • 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.
    • 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.
    • Petranka, J.W. 1998. Salamanders of the United States and Canada. Smithsonian Institution Press, Washington D.C. 587 pp.
    • Russell, A. P. and A. M. Bauer. 2000. The Amphibians and Reptiles of Alberta: A field guide and primer of boreal herpetology. University of Calgary Press, Toronto, Ontario. 279 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.
    • Sessions, S.K. and S.B. Ruth. 1990. Explanation for naturally occurring supernumerary limbs in amphibians. Journal of Experimental Zoology 254(1): 38-47.
    • Slater, J.R. 1936. Notes on Ambystoma gracile Baird and Ambystoma macrodactylum Baird. Copeia 1936(4): 234-236.
    • 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.
    • 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., W.J. Liss, R.L. Hoffman, and L.M. Ganio. 1998b. Experimental analysis of trout effects on survival, growth, and habitat use of two species of Ambystomid salamanders. Journal of Herpetology 32(3): 345-349.
    • 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.
    • 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.
  • Additional ReferencesLegend:   View Online Publication
    Do you know of a citation we're missing?
    • [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. 1968a. Thermal histories of two populations of Ambystoma macrodactylum. Herpetologica 24(1): 29-35.
    • 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. 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.
    • 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.
    • 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.
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    • 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, K. 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.
  • Web Search Engines for Articles on "Long-toed Salamander"
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Citation for data on this website:
Long-toed Salamander — Ambystoma macrodactylum.  Montana Field Guide.  .  Retrieved on , from