Ambystoma macrodactylum



INTRODUCTORY

AUTHORSHIP AND CITATION:
Howard, Janet L. 1997. Ambystoma macrodactylum. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/ [].

ABBREVIATION:
AMMA

SYNONYMS:
No entry

COMMON NAMES:
long-toed salamander

TAXONOMY:
The currently accepted scientific name of the long-toed salamander is Ambystoma macrodactylum Baird. It is in the mole salamander family, Ambystomatidae [10,23,27]. Subspecies of the long-toed salamander are:

A. m. columbianum Ferguson, eastern long-toed salamander [5]
A. m. croceum Ferguson, Santa Cruz long-toed salamander [9,27]
A. m. krausei Peters, northern long-toed salamander [5]
A. m. macrodactylum Baird, western long-toed salamander
A. m. sigillatum, southern long-toed salamander [27]

When the species as a whole is discussed in this report, it will be referred to by its common name, long-toed salamander. Subspecies will be referred to by the subspecies common names listed above.

ORDER:
Caudata (salamanders)

CLASS:
Amphibian

FEDERAL LEGAL STATUS:
The Santa Cruz long-toed salamander is federally listed as Endangered [29].

OTHER STATUS:
Information on state- and province-level protection status of animals in the United States and Canada is available at NatureServe, although recent changes in status may not be included.


WILDLIFE DISTRIBUTION AND OCCURRENCE

SPECIES: Ambystoma macrodactylum
GENERAL DISTRIBUTION:
The long-toed salamander is distributed from southeastern Alaska and northwestern British Columbia south through Washington and Oregon to northeastern California and east to extreme western Alberta, the Idaho panhandle, and western Montana [2,27]. Distribution of subspecies is as follows[27]:

western long-toed salamander: Vancouver Island, British Columbia; Coastal Ranges of Washington and Oregon

eastern long-toed salamander: southeastern Alaska and northern British Columbia; central and eastern Washington; north-central and northeastern Oregon; western half of the Idaho panhandle

northern long-toed salamander: eastern British Columbia; extreme western Alberta; western Montana; eastern half of the Idaho panhandle

southern long-toed salamander: southwestern Oregon; northeastern California

Santa Cruz long-toed salamander: a disjunct population known from only 3 locations in California - Ellicott Pond State Wildlife Reserve, Santa Cruz Co.; Valencia Lagoon, Santa Cruz Co.; Elkhorn Slough, Monterey Co.

ECOSYSTEMS:
FRES20 Douglas-fir
FRES21 Ponderosa pine
FRES22 Western white pine
FRES23 Fir-spruce
FRES24 Hemlock-Sitka spruce
FRES25 Larch
FRES26 Lodgepole pine
FRES28 Western hardwoods
FRES29 Sagebrush
FRES36 Mountain grasslands
FRES37 Mountain meadows
FRES41 Wet grasslands
FRES44 Alpine

STATES:
AK CA ID MT OR WA

AB BC

BLM PHYSIOGRAPHIC REGIONS:
1 Northern Pacific Border
2 Cascade Mountains
3 Southern Pacific Border
4 Sierra Mountains
5 Columbia Plateau
8 Northern Rocky Mountains


KUCHLER PLANT ASSOCIATIONS:
K001 Spruce-cedar-hemlock forest
K002 Cedar-hemlock-Douglas-fir forest
K003 Silver fir-Douglas-fir forest
K004 Fir-hemlock forest
K005 Mixed conifer forest
K007 Red fir forest
K008 Lodgepole pine-subalpine forest
K009 Pine-cypress forest
K010 Ponderosa shrub forest
K011 Western ponderosa forest
K012 Douglas-fir forest
K013 Cedar-hemlock-pine forest
K014 Grand fir-Douglas-fir forest
K015 Western spruce-fir forest
K018 Pine-Douglas-fir forest
K025 Alder-ash forest
K026 Oregon oakwoods
K028 Mosaic of K002 and K026
K029 California mixed evergreen forest
K030 California oakwoods
K049 Tule marshes
K052 Alpine meadows and barren
K055 Sagebrush steppe

SAF COVER TYPES:
205 Mountain hemlock
206 Engelmann spruce-subalpine fir
207 Red fir
208 Whitebark pine
210 Interior Douglas-fir
211 White fir
212 Western larch
213 Grand fir
215 Western white pine
217 Aspen
218 Lodgepole pine
221 Red alder
222 Black cottonwood-willow
223 Sitka spruce
224 Western hemlock
225 Western hemlock-Sitka spruce
226 Coastal true fir-hemlock
227 Western redcedar-western hemlock
228 Western redcedar
229 Pacific Douglas-fir
230 Douglas-fir-western hemlock
233 Oregon white oak
234 Douglas-fir-tanoak-Pacific madrone
235 Cottonwood-willow
237 Interior ponderosa pine
243 Sierra Nevada mixed conifer
244 Pacific ponderosa pine-Douglas-fir
245 Pacific ponderosa pine
246 California black oak
247 Jeffrey pine
255 California coast live oak
256 California mixed subalpine

SRM (RANGELAND) COVER TYPES:
109 Ponderosa pine shrubland
110 Ponderosa pine-grassland
202 Coast live oak woodland
203 Riparian woodland
213 Alpine grassland
216 Montane meadows
217 Wetlands
401 Basin big sagebrush
409 Tall forb
411 Aspen woodland
422 Riparian
906 Broadleaf forest
921 Willow

PLANT COMMUNITIES:
The long-toed salamander occupies a wide variety of habitats including grassland, sagebrush (Artemisia spp.)-grassland, pinyon-juniper (Pinus-Juniperus spp.) woodland, coniferous forest, and coast live oak (Quercus agrifolia) woodland communities [1,5,6,7,30].


BIOLOGICAL DATA AND HABITAT REQUIREMENTS

SPECIES: Ambystoma macrodactylum
TIMING OF MAJOR LIFE HISTORY EVENTS:
Migration to breeding waters begins in winter for southerly subspecies and during spring snowmelt for northerly subspecies. Sustained temperatures above freezing and abundant, free soil water apparently trigger migration. Migration almost always occurs at night. Long-toed salamanders in cold climates crawl on top of and beneath snow to reach breeding ponds; the ponds are often still partially frozen during mating [2,5,12]. Mating behaviors are described in Anderson [1]. In all locales, males arrive at ponds before females and generally stay for longer periods of time. Females leave soon after depositing their eggs [2,5,12]. They attach eggs to vegetation, submerged wood, or rocks. Santa Cruz long-toed salamanders lay single eggs; southern long-toed salamanders lay eggs in small clusters. Other subspecies tend to lay eggs in masses [2]. The breeding period lasts about a month [2,5,12]; high-elevation populations generally have longer breeding periods than low-elevation populations [12].

Life span: A sampling of adults and subadults (individuals that have metamorphosed but not yet reached sexual maturity) in a northern long-toed salamander population in Alberta showed an age distribution from 1 year to 10 years of age, with most individuals in the 2- to 3-year-old age bracket [20].

Life History - General: Life histories of long-toed salamanders vary with temperature and moisture conditions. Several life history patterns are evident: a one-season larval period (in warm climates); either a short facultative one-season larval period or a two-season larval period (moderate climates); and a three- to four-season larval period (cold climates). In warm climates, period of development is limited by precipitation. Breeding cannot take place until temporary ponds fill. Variation in rainfall determines the length of time water remains and, therefore, period of larval development. Metamorphosis occurs when ponds begin to shrink [2]. In cold climates, development time extends to several years due to short growing season. Regardless of subspecies, long-toed salamander larvae do not transform until attaining a snout-to-vent length of at least 33 mm. In cold climates, it may take 4 years to reach that size [12].

Life History of Subspecies:

Eastern long-toed salamanders - In ponderosa pine forest near Moscow, Idaho, migration to breeding ponds began in late February. By late April, most adults had left the ponds [22]. Howard and Wallace [13] reported that low-elevation (1,390 feet (420 m)) populations in Nez Perce County, Idaho, bred in early February; mid-elevation (3,760 feet (1140 m)) populations in Baker County, Oregon, bred in April; and high-elevation (8,150 feet (2470 m)) populations in Wallowa County, Oregon, bred in June and July. Females at high-elevation sites laid fewer, larger eggs than females at lower-elevation sites. Number of eggs per female averaged 166 (SD +/- 60) at the 1,390-foot site and 90 (SD +/-49) at the 8,150-foot site. Larvae from populations below 6,930 feet (2100 m) metamorphosed in their first summer, while larvae from higher elevations metamorphosed in late summer of their third or fourth year.

Northern long-toed salamanders - A sample of wild individuals in Alberta reached sexual maturity at 47 mm in length, a length attained at about 3 years of age. Well-fed, captive individuals, raised in aquaria from eggs, exceeded 47 mm in length by their first year but did not reach sexual maturity until their second year [20].

Southern long-toed salamanders - Populations in the Sierra Nevada have facultative one-season and two-season larval periods. In Calaveras County, California, at 6,530 feet (1980 m) elevation, time from egg deposition to metamorphosis was 80 to 90 days in temporary ponds. Larval period is probably longer at that elevation in large, permanent ponds. At higher elevations, southern long-toed salamanders do not reach the critical size for metamorphosis in a single season. In Alpine County, California, (elevation 8,085 feet (2450 m)), mating and egg deposition occurs from late May to late June, as soon as ponds partially thaw. Larvae develop in summer and spend the winter beneath ice, transforming in August or September of their second year. Adults first reproduce at age 2 or 3 [2].

Santa Cruz long-toed salamanders - Living in a mediterranean climate, Santa Cruz long-toed salamanders experience one of the driest environments of the species. Larval development is completed within one season. In drought years, rainfall is sometimes insufficient to allow normal breeding and larval development to occur. In wetter years, migration to breeding ponds begins with late fall and winter rains. Santa Cruz long-toed salamanders only migrate on rainy nights. Subadults move to ponds after light rains, but adults migrate only after heavy, ground-soaking rains. Breeding occurs from January to mid-February and eggs hatch from late February to mid-March. Time from breeding to larval transformation and pond shrinkage varies from about 90 to 140 days. Santa Cruz long-toed salamanders are sexually mature at age 2 [2].

PREFERRED HABITAT:
General: Long-toed salamanders occur in diverse habitats including coniferous forest, oak (Quercus spp.) woodland, alpine, sagebrush (Artemisia spp.), and marshland communities [2,26]. They use springs, ponds, small lakes, slow-moving streams, and marshlands for breeding and larval development [2,5].

Habitat of Subspecies:

Eastern long-toed salamanders occur in ponderosa pine, lodgepole pine, and subalpine fir-Engelmann spruce (Abies lasiocarpa-Picea engelmannii) zones. A population near Moscow, Idaho, used artificial ponds within ponderosa pine-grand fir (A. grandis) forest for breeding [22]. Eastern long-toed salamanders have also been documented in wheatfields (Triticum aestivus) with irrigation ponds, ponderosa pine-big sagebrush (Artemisia tridentata) woodlands with temporary ponds, and sparsely vegetated whitebark pine-mountain heather (Pinus albicaulis-Phyllodoce empetriformis) communities with permanent lakes [12,13]. Long-toed salamander larvae, presumably eastern long-toed salamanders, were found in a spring within a cottonwood-quaking aspen (Populus spp.-P. tremuloides) riparian community on the Bruneau Resource Area of southern Idaho [18].

The southern long-toed salamander occurs in mixed Sierra Nevada coniferous forest and alpine communities. It has been noted at 8,075 feet (2,750 m) elevation in Alpine County, California. A population at 6,534 feet (1980 m) elevation in Calaveras County, California, occurred in and near a temporary pond formed from snowmelt. The pond was shaded by large trees, including white fir (Abies concolor), ponderosa pine (Pinus ponderosa), lodgepole pine (P. contorta), and quaking aspen, that provided shade for most of the day. The pond was clear and moderately acidic (pH 5.9). It lacked aquatic vegetation and was littered with needles and small woody debris. Further east, a population occurring at 8,085 feet (2,450 m) in Alpine County, California, occupied permanent ponds fed by snowmelt and springs. Lodgepole pine, western white pine (Pinus monticola), and mountain hemlock (Tsuga mertensiana) were sparse to numerous around pond margins but always provided at least some shade. The pond waters were very clear, lacking live vegetation but with considerable downed woody debris including floating and submerged logs [2].

Santa Cruz long-toed salamanders in the two Santa Cruz County populations occur in and near temporary ponds in coast live oak (Quercus agrifolia) woodlands [2,21]. Pond waters are often turbid and aquatic plant growth is extensive. In summer, adults seek moist areas such as seeps and willow (Salix spp.) thickets near pond shores [2]. The Monterey County population occurs in a cattail-bulrush (Typha-Scirpus spp.) marsh [26].

COVER REQUIREMENTS:
Since they are highly susceptible to desiccation, adult and subadult long-toed salamanders spend most of their lives underground or beneath objects. Larvae use submerged objects and aquatic vegetation for cover [2].

Southern long-toed salamander larvae generally remain hidden under bark, logs, or other submerged objects. They overwinter beneath such objects, in water more than 12 inches (30 cm) deep. In mid-summer in Calaveras County, California, subadults sought cover beneath objects in dried temporary ponds; they were never found outside pond perimeters. In late summer, subadults were still beneath objects in the dried ponds but had formed ball-shaped aggregations of 15 to 43 individuals. Adults used large, rotting logs for cover most of the year [2].

Santa Cruz long-toed salamander larvae in Santa Cruz County use dense aquatic vegetation and turbid water for cover. Subadults cannot disperse to coast live oak woodlands immediately after transformation due to arid summer climate. After summer metamorphosis, they retreat to willow thickets at shore edges or beneath matted vegetation or other debris at the bottoms of drying ponds. When these substrates dry, subadults seek the same substrates used by adults in summer: rodent burrows, buried logs, dense tule (Scirpus acutus) mats, or other microhabitats where moisture is retained throughout the dry season. Subadults often aggregate at these sites, tightly entwined in groups of three to nine individuals. With onset of autumn rains, subadults move into coast live oak woodlands [2]. Adult Santa Cruz salamanders in Monterey County have been found in willow thickets and beneath wooden boxes and other urban debris during the dry season [26].

FOOD HABITS:
Adult long-toed salamanders hunt terrestrial and aquatic arthropods. They also scavenge dead arthropods [19,30]. The diet of larvae is similar: larvae consume aquatic arthropods and terrestrial arthropods that fall into the water, and scavenge arthropod remains. In addition, some long-toed salamander larvae are cannibalistic. Cannibal larvae are morphologically different from "normal" larvae, having larger heads and jaws, reduced gills, and a more slender body. Larvae may become cannibalistic in response to either high larval population density or a scarcity of other food sources [31].

In summer, proteins and fats are stored in the tails of long-toed salamanders. These nutrients are metabolized during long periods of dormancy [32].

Eastern long-toed salamander larvae in Oregon have been observed feeding on hatchling Pacific treefrog (Hyla regilla) larvae. Cascades frog (Rana cascadae) larvae and fairy shrimp (Anostraca) were other potential prey in the breeding pond [31].

PREDATORS:
Adult long-toed salamanders are probably not highly vulnerable to predation. Except during migration, they are secretive in habit. Even then, they migrate to and from breeding ponds at night, in winter or during spring snowmelt, when most predators that would potentially prey on long-toed salamander are relatively inactive [5]. Additionally, long-toed salamanders secrete a toxin from glands in their tails when captured; the toxin often prompts predators to drop and abandon the long-toed salamanders [32].

Near Moscow, Idaho, a common garter snake (Thamnophis sirtalis) was observed in the process of swallowing an eastern long-toed salamander. Other potential predators captured near breeding ponds were western terrestrial garter snakes (T. elgans) and shrews (Sorex spp.). However, these predators were not active until late April, when all but a few male long-toed salamanders had already departed from breeding ponds and returned to forest cover [5].

Long-toed salamander larvae prey upon each other [2,5].

MANAGEMENT CONSIDERATIONS:
No entry


FIRE EFFECTS AND USE

SPECIES: Ambystoma macrodactylum
DIRECT FIRE EFFECTS ON ANIMAL:
There are no data available on fire effects on salamanders. It is likely that fire has little direct effect on long-toed salamanders. Adults are rarely active above ground; when they are active, it is usually under cool, moist weather conditions [13] that occur outside the fire season. During the dry season, adult long-toed salamanders are generally either in burrows, where they are relatively safe from fire, or under moist rotten logs or moist vegetation mats that are not likely to burn except except under extreme fire weather conditions.

If caught in the open during a fire, long-toed salamanders would probably be killed. They are very slow-moving [2], and probably cannot escape even slow-moving fire. Even if missed by fire, they probably could not survive the heat. High temperatures are lethal to long-toed salamanders. In the laboratory, adults from northeastern Oregon and western Idaho were killed by water temperatures that ranged from above 91 to 96 degrees Fahrenheit (33-36 oC) [13].

HABITAT RELATED FIRE EFFECTS:
Adult and subadult long-toed salamanders use logs and large branches for cover, and larvae use floating and submerged downed woody debris of all size classes for cover [2]. Fire that increases downed woody debris while retaining some overhead shade probably improves habitat structure of long-toed salamanders.

FIRE USE:
No entry


REFERENCES

SPECIES: Ambystoma macrodactylum
1. Anderson, James D. 1961. The courtship behavior of Ambystoma macrodactylum croceum. Copeia. 2: 132-139. [27244]

2. Anderson, James D. 1967. A comparison of the life histories of coastal and montane populations of Ambystoma macrodactylum in California. The American Midland Naturalist. 77(2): 323-355. [23174]

3. Anderson, James D. 1968. A comparison of the food habits of Ambystoma macrodactylum sigillatum, Ambystoma macrodactylum croceum, and Ambystoma tigrinum californiense. Herpetologica. 24(4): 273-284. [28328]

4. Beneski, John T., Jr.; Larsen, John H., Jr.; Miller, Brian T. 1995. Variation in the feeding kinematics of mole salamanders (Ambystomatidae: Ambystoma) Canadian Journal of Zoology. 73(2): 353-366. [28330]

5. Beneski, John T., Jr.; Zalisko, Edward J.; Larsen, John H., Jr. 1986. Demography and migratory patterns of the eastern long-toed salamander, Ambystoma macrodactylum columbianum. Copeia. 1986(2): 398-408. [23175]

6. Bernard, Stephen R.; Brown, Kenneth F. 1977. Distribution of mammals, reptiles, and amphibians by BLM physiographic regions and A.W. Kuchler's associations for the eleven western states. Tech. Note 301. Denver, CO: U.S. Department of the Interior, Bureau of Land Management. 169 p. [434]

7. Bradford, David F.; Swanson, Christina; Gordon, Malcolm S. 1994. Effects of low pH and aluminum on amphibians at high elevation in the Sierra Nevada, California. Canadian Journal of Zoology. 72(7): 1272-1279. [28331]

8. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. [905]

9. Ferguson, Denzel E. 1961. The geographic variation of Ambystoma macrodactylum Barid, with the description of two new subspecies. The American Midland Naturalist. 65(2): 311-338. [27245]

10. Frost, Darrel R., ed. 1985. Amphibian species of the world: a taxonomic and geographic reference. Lawrence, KS: Allen Press, Inc. 732 p. [26535]

11. Garrison, George A.; Bjugstad, Ardell J.; Duncan, Don A.; [and others]. 1977. Vegetation and environmental features of forest and range ecosystems. Agric. Handb. 475. Washington, DC: U.S. Department of Agriculture, Forest Service. 68 p. [998]

12. Howard, James H.; Wallace, Richard L. 1985. Life history characteristics of populations of the long-toed salamander (Ambystoma macrodactylum) from different altitudes. The American Midland Naturalist. 113(2): 361-373. [21751]

13. Howard, James H.; Wallace, Richard L.; Stauffer, Jay R., Jr. 1983. Critical thermal maxima in populations of Ambystoma macrodactylum from different elevations. Journal of Herpetology. 17(4): 400-402. [27246]

14. Kezer, James; Farner, Donald S. 1955. Life history patterns of the salamander Ambystoma macrodactylum in the high Cascade Mountains of southern Oregon. Copeia. 2: 127-131. [28332]

15. Knudsen, Jens W. 1960. The courtship and egg mass of Ambystoma gracile and Ambystoma macrodactylum. Copeia. 1: 44-46. [28333]

16. Kuchler, A. W. 1964. United States [Potential natural vegetation of the conterminous United States]. Special Publication No. 36. New York: American Geographical Society. 1:3,168,000; colored. [3455]

17. Leonard, William P.; Richter, Klaus O. 1994. Western long-toed salamander demographics and oviposition in a small vernal wetland of the Puget Sound lowlands. Northwest Science. 68(2): 135. [28334]

18. Munger, James C.; Peterson, Charles. 1993. A preliminary survey of the herpetofauna of Bruneau Resource Area, Boise District. Tech. Bulletin No. 93-2. Boise, ID: U.S. Department of the Interior, Bureau of Land Management, Idaho State Office. 34 p. [23681]

19. Oliver, James A. 1955. The natural history of North American amphibians and reptiles. Princeton, NJ: D. Van Nostrand Company, Inc. 359 p. [28336]

20. Russell, Anthony P.; Powell, G. Lawrence; Hall, Dean R. 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. [27247]

21. Russell, R. W.; Anderson, James D. 1956. A disjunct population of the long-nosed salamander from the coast of California. Herpetologica. 12: 137-140. [23227]

22. Schaub, David L.; Larsen, John H., Jr. 1978. The reproductive ecology of the Pacific treefrog (Hyla regilla). Herpetologica. 34(4): 409-416. [27248]

23. Schmidt, Karl P. 1953. A checklist of North American amphibians and reptiles. 6th ed. Chicago, IL: University of Chicago Press; American Society of Icthyologists and Herpetologists. 280 p. [24389]

24. Semlitsch, Raymond D. 1983. Burrowing ability and behavior of salamanders of the genus Ambystoma. Canadian Journal of Zoology. 61: 616-620. [28335]

25. Shiflet, Thomas N., ed. 1994. Rangeland cover types of the United States. Denver, CO: Society for Range Management. 152 p. [23362]

26. State of California, Department of Fish and Game, Natural Heritage Division, Natural Diveristy Data Base. 1995. Endangered and threated animals of California. [Sacramento, CA]. 13 p. [24904]

27. Stebbins, Robert C. 1985. Western reptiles and amphibians. 2nd ed. Peterson Field Guides No. 16. Boston: Houghton Mifflin Company. 336 p. [22647]

28. The Network of Natural Heritage Programs and Conservation Data Centers and The Nature Conservancy. 1994. Element distribution - North America, vertebrates. Arlington, VA: The Nature Conservancy, Central Conservation Databases. 31 p. [23374]

29. U.S. Fish and Wildlife Service. 2013. Listed animals. In: Environmental Conservation Online System, [Online]. In: Species reports. Available: http://ecos.fws.gov/tess_public/pub/listedAnimals.jsp. [86534]

30. Verner, Jared; Boss, Allan S., tech. coords. 1980. California wildlife and their habitats: western Sierra Nevada. Gen. Tech. Rep. PSW-37. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station. 439 p. [10237]

31. Walls, Susan C.; Beatty, Joseph J.; Tissot, Brian N.; [and others]. 1993. Morphological variation and cannibalism in a larval salamander (Ambystoma macrodactylum columbianum). Canadian Journal of Zoology. 71: 1543-1551. [27249]

32. Williams, Thomas A.; Larsen, John H., Jr. 1986. New function for the granular skin glands of the eastern long-toed salamander, Ambystoma macrodactylum columbianum. Journal of Experimental Zoology. 239: 329-333. [23176]


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