Ambystoma macrodactylum
|
|
 |
| Creative Commons image © 2013 John P. Clare. |
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: www.fs.usda.gov/database/feis/animals/amphibian/amma/all.html
[].
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:
Ambystoma macrodactylum columbianum Ferguson, eastern long-toed salamander [5]
Ambystoma macrodactylum croceum Ferguson, Santa Cruz long-toed salamander [9,27]
Ambystoma macrodactylum krausei Peters, northern long-toed salamander [5]
Ambystoma macrodactylum macrodactylum Baird, western long-toed salamander
Ambystoma macrodactylum 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.
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:
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].
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
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
FIRE REGIMES : Find fire regime information for the plant communities in which this
species may occur by entering the species name in the FEIS home page under
"Find Fire Regimes".
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. Department of the Interior, Fish and Wildlife Service. 2016. Endangered Species Program, [Online]. Available: http://www.fws.gov/endangered/. [86564]
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]
FEIS Home Page
https://www.fs.usda.gov/database/feis/animals/amphibian/amma/all.html
