dcsimg

Description

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A small species of Ambystoma, dark gray to black with a yellow, tan or olive green dorsal stripe often broken up into blotches (Stebbins 1951). The sides have some white speckling. The ventral side is gray or black (Petranka 1998). Ambystoma macrodactylum columbianum A. m. croceum, A. m. krausei, A. m. macrodactylum, A. m. sigillatum are subspecies.See other subspecies accounts at www.californiaherps.com: A. m. croceum (http://www.californiaherps.com/salamanders/pages/a.m.croceum.html) and A. m. sigillatum (http://www.californiaherps.com/salamanders/pages/a.m.sigillatum.html).

References

  • Beneski, J. T. Jr., Zalisko, E. J., and Larsen, J. H. (1986). ''Demography and migratory patterns of the Eastern Long-toed Salamander Ambystoma macrodactylum columbianum.'' Copeia, 1986, 398-408.
  • Blaustein, A. R., Kiesecker, J. M., Chivers, D. P., and Anthony, R. G. (1997). "Ambient UV-B radiation causes deformities in amphibian embryos." Proceedings of the National Academy of Sciences of the United States of America, 94(25), 13735-13737.
  • Sessions, S. K., and Ruth, S. B. (1990). ''Explanation for naturally occurring supernumerary limbs in amphibians.'' Journal of Experimental Zoology, 254, 38-47.

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Lauren M. Chan
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Distribution and Habitat

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Their range extends from south-eastern Alaska south to northern California, and from the Pacific coast east to north-central Idaho and western Montana (Petranka 1998). Found in a variety of habitats from coniferous forests to sagebrush plains to alpine meadows. Found on the ground under bark, rocks, and rotting wood piles. Also found in the quiet water of streams, ponds and lakes (Stebbins 1951).
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Life History, Abundance, Activity, and Special Behaviors

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Eggs exposed to ambient levels of UV-B radiation have been shown to have increased mortality and incidence of deformities than those shielded from UV-B (Blaustein et al 1997). A trematode has been found that disrupts both limb development and regeneration and has been proposed as an explanation of why individuals with supernumerary limbs are found (Sessions and Ruth 1990). Environmental contaminants as well as the introduction of non-native fish predators may also threaten this species. The destruction of wetland habitats may prove to be the greatest threat. The subspecies, A. m. croceum, persists in only a few scattered populations and is threatened with extinction (Petranka 1998).
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Life History, Abundance, Activity, and Special Behaviors

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Life history varies greatly with elevation and climate. Each season individuals migrate to breeding ponds with males arriving earlier and staying longer than females (Beneski et al 1986). At low elevations this migration may be in October or November, but at higher elevations it does not occur until snowmelt in late spring (Petranka 1998). Males deposit spermatophores (packets of sperm) which females pick up after courtship. Single eggs or loose egg clumps are attached to vegetation or detritus. Larvae hatch 2-5 weeks later and metamorphose in about 3 months (Petranka 1998).
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Lifespan, longevity, and ageing

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Maximum longevity: 10 years (wild) Observations: Their life history depends greatly on temperature. In the wild, they live up to 10 years (http://www.fs.fed.us/database/feis/).
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Benefits

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Much of the 139 acres set aside for the species is of high economic value, which is lost if left undeveloped.

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Hilt, M. 1999. "Ambystoma macrodactylum croceum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Ambystoma_macrodactylum_croceum.html
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Megan Hilt, University of Michigan-Ann Arbor
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Life Cycle

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Development - Life Cycle: metamorphosis

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Hilt, M. 1999. "Ambystoma macrodactylum croceum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Ambystoma_macrodactylum_croceum.html
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Megan Hilt, University of Michigan-Ann Arbor
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Conservation Status

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Since being placed on the first Endangered Species List on March 11, 1967, significant steps have been taken in order to ensure their survival, including the reservation of a 139 acre Ellicott Slough National Wildlife Refuge, in San Francisco Bay, California.(U.S. Fish and Wildlife Service 1986, Westphal 1996) Since the loss of habitat is one major source of impending danger to these salamanders' population, the established refuge has been a large step towards saving this rare species.

US Federal List: no special status

CITES: no special status

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Hilt, M. 1999. "Ambystoma macrodactylum croceum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Ambystoma_macrodactylum_croceum.html
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Megan Hilt, University of Michigan-Ann Arbor
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Trophic Strategy

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Both aquatic larvae and terrestrial adults are carnivorous. The larvae primarily eat small aquatic insects and arthropods. The adults also prey upon tree frog tadpoles, earthworms, slugs and various terrestrial insects.(Ferguson 1961)

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Hilt, M. 1999. "Ambystoma macrodactylum croceum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Ambystoma_macrodactylum_croceum.html
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Megan Hilt, University of Michigan-Ann Arbor
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Distribution

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The Santa Cruz long-toed salamander is a relict species, previously widespread throughout California after the Pleistocene era, and is now concentrated in the area of Santa Cruz, California.(Hukill 1997) Upon time of its discovery in 1954 until the present, this species has inhabited four locations around Santa Cruz County. They include the cities of Ellicott, Valencia, Seascape, and Bennett.(U.S. Fish and Wildlife Service 1986)

Biogeographic Regions: nearctic (Native )

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Hilt, M. 1999. "Ambystoma macrodactylum croceum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Ambystoma_macrodactylum_croceum.html
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Habitat

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These salamanders live in ponds, streams or lagoons in the larval stage. As terrestrial adults, the salamanders live in isolated populations in upland and mixed forests and estivate in aquatic habitats. They favor living conditions that allow their skin to stay moist and cool, and residing under forest litter does just this.(Larson 1997)

Terrestrial Biomes: forest

Aquatic Biomes: lakes and ponds; rivers and streams

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Hilt, M. 1999. "Ambystoma macrodactylum croceum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Ambystoma_macrodactylum_croceum.html
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Megan Hilt, University of Michigan-Ann Arbor
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Morphology

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Ambystoma macrodactylum croceum is a small salamander, reaching a full-grown adult length of approximately 127 mm, (5 inches). The body of an adult is a glossy black-charcoal color, with brilliant yellow and orange spots covering its back, decreasing in abundance at the top of the head, and virtually disappearing at the small black eyes. Their head is broad, blunt, and torpedo-shaped.(Duellman and Trueb 1986, Westphal 1996) Two pairs of limbs of approximately equal size are set at right angles from the rib cage.(Ferguson 1961) The four toes on the front legs and five on the back are extremely long. The adult tail is laterally flattened, metamorphosed from the larval stage of a fin. True teeth form a row across the roof of the mouth. In the larval stage, salamanders resemble the adults except that they have caudal fins, four external gill slits on either side of the head, and are a mottled, greenish color.(Duellman and Trueb 1986 Westphal 1996)

Other Physical Features: ectothermic ; bilateral symmetry

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Hilt, M. 1999. "Ambystoma macrodactylum croceum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Ambystoma_macrodactylum_croceum.html
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Reproduction

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The reproductive behavior of Ambystoma macrodactylum croceum is highly unique. After a period of estivation, the salamanders travel great distances to their original breeding site. The males stay for approximately one month, while the females stay for only two weeks. The males spend this time competing with each other for resources in order to increase their fitness, (strength and virility). This increases the brightness of the yellow-orange markings, ultimately augmenting mate sexual preference. In the final courtship ritual, the male nudges the female, and if she is receptive, the male then places his chin on the female's head, rubbing his chin gland in a display of preference. The male then moves away, and the female follows snout-to-tail. The male deposits a mushroom-shaped spermatophore (gelatinous blob of sperm) at the bottom of the lake (pond, or slow-moving stream) for the female. She later takes the spermatophore into her "vent" to fertilize the eggs internally. Approximately 200-400 eggs are laid on a submerged vegetative stalk, and hatch 2-3 weeks later.(Hukill 1997) These jelly-encased eggs are the largest eggs of all salamanders in the area.(Westphal 1996) Larvae live in the pond until about March, when their tails will lose their fin, toes lengthen, and the gills disappear. Sexual maturity is reached at 3-4 years.(U.S. Fish and Wildlife Service 1986)

Key Reproductive Features: gonochoric/gonochoristic/dioecious (sexes separate)

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Hilt, M. 1999. "Ambystoma macrodactylum croceum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Ambystoma_macrodactylum_croceum.html
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Overview

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Ambystoma macrodactylum, or long-toed salamander, is a common amphibian in the family Ambystomatidae, or mole salamanders. Its native range extends from southeastern Alaska through British Columbia, Washington, and much of Oregon to northern California and from the Pacific coast eastward to northern Idaho, western Montana, and southwestern Alberta (Bull 2005).Long-toed salamanders are extremely adaptable amphibians, but they prefer to live in moist environments near ponds or lakes. They are most identifiable by a long greenish-yellow dorsal stripe, with the rest of their backs being black and their undersides dark brown with white flecks. Adults are about three to four inches long and quite slender. Their name comes from the slightly longer fourth toe on their hind feet (Rockney and Wu 2015).

Long-toed salamanders are the earliest breeding amphibians in their range. The timing differs depending on elevation, with those at lower elevations beginning in late fall and winter and those in alpine habitats waiting until summer (Pilliod and Fronzuto 2016). They are considered opportunistic breeders, meaning that they will lay eggs in all sorts of places, in the hope of at least some surviving. Once the eggs hatch, the aquatic larval stage begins, which can last from 1 to 3 years. The larvae feed on aquatic insects and crustaceans. They eventually lose their gills, produce digits, and metamorphose into terrestrial juveniles. The juvenile and adult salamanders stay within a small range of the pond or body of water where they were born, which will become their breeding grounds when they mature. Adult long-toed salamanders feed on arthropods, mollusks and annelids. They can live up to 7 years (Pilliod and Fronzuto 2016).

Ambystoma macrodactylum probably hibernate during winter, particularly at higher elevations, although adults can remain active all year long in lowland areas (Stebbins 2003). Adults hibernate underground, whereas the larvae can hide under logs and on the underside of leaves or sticks in a pond until it becomes warm enough for them to emerge. If and when they hibernate, they usually do so in groups and survive off of stored protein in their bodies, for months if needed. In general, they lead a very secretive life to avoid encountering predators (Pilliod and Fronzuto 2016). When they are threatened by predators, they coil their bodies or lash their tails, produce secretions from their skin, and can vocalize to either scare away the predator or warn other salamanders of danger. Because they absorb water and oxygen through their skin, they are very sensitive to the conditions of their habitats. This makes them good indicators of the health of the environment. They aren’t considered to be endangered, although they are threatened in certain regions.

References

  • Stebbins, Robert C. 2003. A Field Guide to Western Reptiles and Amphibians, 3d ed. Boston: Houghton Mifflin.
  • Bull, Evelyn L. 2005. “Long-Toed Salamander.” In Amphibians of the Pacific Northwest, edited by Lawrence L.C. Jones, William P. Leonard, and Deanna H. Olsen, 34-37. Seattle: Seattle Audubon Society.
  • Pilliod, David S., and Julie A. Fronzuto. 2016. “Ambystoma macrodactylum.” AmphibiaWeb: Information on amphibian biology and conservation. Berkeley, California. http://amphibiaweb.org. Accessed June 1, 2016.
  • Rockney, Heidi, and Karen Wu. 2015. Long-Toed Salamander. Burke Museum of Natural History and Culture. U of Washington. http://www.burkemuseum.org/blog/long-toed-salamander. Accessed May 31, 2016.

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Authors:Natasha Rosenbach,Hadley Reeder,Hadley Reede,Martha Rubardt,Rose Thompson; Editor: Gordon Miller, Seattle University EVST 2100 - Natural History: Theory and Practice
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Associated Plant Communities

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More info for the terms: forest, grassland, woodland

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].
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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.fed.us/database/feis/animals/amphibian/amma/all.html

Common Names

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long-toed salamander
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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.fed.us/database/feis/animals/amphibian/amma/all.html

Conservation Status

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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.
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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.fed.us/database/feis/animals/amphibian/amma/all.html

Cover Requirements

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More info for the term: cover

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].
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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.fed.us/database/feis/animals/amphibian/amma/all.html

Direct Effects of Fire

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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].
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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.fed.us/database/feis/animals/amphibian/amma/all.html

Distribution

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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.
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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.fed.us/database/feis/animals/amphibian/amma/all.html

Food Habits

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More info for the term: density

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].
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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.fed.us/database/feis/animals/amphibian/amma/all.html

Habitat-related Fire Effects

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More info for the term: cover

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.
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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.fed.us/database/feis/animals/amphibian/amma/all.html

Habitat: Cover Types

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This species is known to occur in association with the following cover types (as classified by the Society of American Foresters):

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
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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.fed.us/database/feis/animals/amphibian/amma/all.html

Habitat: Ecosystem

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This species is known to occur in the following ecosystem types (as named by the U.S. Forest Service in their Forest and Range Ecosystem [FRES] Type classification):

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
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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.fed.us/database/feis/animals/amphibian/amma/all.html

Habitat: Plant Associations

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More info on this topic.

This species is known to occur in association with the following plant community types (as classified by Küchler 1964):

More info for the terms: forest, shrub

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
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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.fed.us/database/feis/animals/amphibian/amma/all.html

Habitat: Rangeland Cover Types

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More info on this topic.

This species is known to occur in association with the following Rangeland Cover Types (as classified by the Society for Range Management, SRM):

More info for the terms: forb, forest, grassland, shrubland, woodland

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
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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.fed.us/database/feis/animals/amphibian/amma/all.html

Occurrence in North America

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AK
CA
ID
MT
OR
WA







AB
BC

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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.fed.us/database/feis/animals/amphibian/amma/all.html

Predators

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More info for the terms: cover, forest

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].
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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.fed.us/database/feis/animals/amphibian/amma/all.html

Preferred Habitat

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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].
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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.fed.us/database/feis/animals/amphibian/amma/all.html

Regional Distribution in the Western United States

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This species can be found in the following regions of the western United States (according to the Bureau of Land Management classification of Physiographic Regions of the western United States):

1 Northern Pacific Border

2 Cascade Mountains

3 Southern Pacific Border

4 Sierra Mountains

5 Columbia Plateau

8 Northern Rocky Mountains
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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.fed.us/database/feis/animals/amphibian/amma/all.html

Timing of Major Life History Events

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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].
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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.fed.us/database/feis/animals/amphibian/amma/all.html

U.S. Federal Legal Status

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The Santa Cruz long-toed salamander is federally listed as Endangered [29].
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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.fed.us/database/feis/animals/amphibian/amma/all.html

Use of Fire in Population Management

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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".

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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.fed.us/database/feis/animals/amphibian/amma/all.html

Long-toed salamander

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The long-toed salamander (Ambystoma macrodactylum) is a mole salamander in the family Ambystomatidae.[2] This species, typically 4.1–8.9 cm (1.6–3.5 in) long when mature, is characterized by its mottled black, brown, and yellow pigmentation, and its long outer fourth toe on the hind limbs. Analysis of fossil records, genetics, and biogeography suggest A. macrodactylum and A. laterale are descended from a common ancestor that gained access to the western Cordillera with the loss of the mid-continental seaway toward the Paleocene.

The distribution of the long-toed salamander is primarily in the Pacific Northwest, with an altitudinal range of up to 2,800 m (9,200 ft). It lives in a variety of habitats, including temperate rainforests, coniferous forests, montane riparian zones, sagebrush plains, red fir forests, semiarid sagebrush, cheatgrass plains, and alpine meadows along the rocky shores of mountain lakes. It lives in slow-moving streams, ponds, and lakes during its aquatic breeding phase. The long-toed salamander hibernates during the cold winter months, surviving on energy reserves stored in the skin and tail.

The five subspecies have different genetic and ecological histories, phenotypically expressed in a range of color and skin patterns. Although the long-toed salamander is classified as a species of Least Concern by the IUCN, many forms of land development threaten and negatively affect the salamander's habitat.

Taxonomy

Ambystoma macrodactylum is a member of the Ambystomatidae, also known as the mole salamanders. The Ambystomatidae originated approximately 81 million years ago (late Cretaceous) from its sister taxon Dicamptodontidae.[3][4][5] The Ambystomatidae are also members of suborder Salamandroidea, which includes all the salamanders capable of internal fertilization.[6] The sister species to A. macrodactylum is A. laterale, distributed in eastern North America. However, the species-level phylogeny for Ambystomatidae is tentative and in need of further testing.[7]

Systematics and biogeography

Evolutionary origins

The ancestral origins for this species stem from eastern North America, where species richness of ambystomatids are highest.[8][9] The following biogeographic interpretation on the origins of A. macrodactylum into western North America is based on a descriptive account of fossils, genetics, and biogeography.[10][11] The long-toed salamander's closest living sister species is A. laterale, a native to northeastern North America.[4][9] Ambystomatidae was isolated to the southeast of the mid-Continental or Western Interior Seaway during the Cretaceous (~145.5–66 Ma).[9][12] While three other species of the Ambystomatidae (A. tigrinum, A. californiense, and A. gracile) have overlapping ranges in western North America, the long-toed salamander's closest living sister species is A. laterale, a native to northeastern North America.[4][9] It has been suggested that A. macrodactylum speciated from A. laterale after the Paleocene (~66–55.8 Ma) with the loss of the Western Interior Seaway opening an access route for a common ancestor into the Western Cordillera.[11] Once situated in the montane regions of western North America, species had to contend with a dynamic spatial and compositional ecology responding to the changes in altitude, as mountains grew and the climate changed. For example, the Pacific Northwest became cooler in the Paleocene, paving the way for temperate forest to replace the warmer tropical forest of the Cretaceous.[13] A scenario for the splitting of A. macrodacylum and other western temperate species from their eastern counterparts involves Rocky Mountain uplift in the late Oligocene into the Miocene. The orogeny created a climatic barrier by removing moisture from the westerly air stream and dried the midcontinental area, from southern Alberta to the Gulf of Mexico.[10][14]

Ancestors of contemporary salamanders were likely able to disperse and migrate into habitats of the Rocky Mountains and surrounding areas by the Eocene. Mesic forests were established in western North America by the mid Eocene and attained their contemporary range distributions by the early Pliocene. The temperate forest valleys and montane environments of these time periods (Paleogene to Neogene) would have provided the physiographic and ecological features supporting analogs of contemporary Ambystoma macrodactylum habitats.[10][11][15][16] The Cascade Range rose during the mid Pliocene and created a rain shadow effect causing the xerification of the Columbia Basin and also altered ranges of temperate mesic ecosystems at higher elevations. The rise of the Cascades causing the xerification of the Columbia Basin is a major biogeographic feature of western North America that divided many species, including A. macrodactylum, into coastal and inland lineages.[11][14][16][17]

Subspecies

There are five subspecies of long-toed salamander.[18] The subspecies are discerned by their geographic location and patterns in their dorsal stripe;[19] Denzel Ferguson gives a biogeographic account of skin patterns, morphology; based on this analysis, he introduced two new subspecies: A. macrodactylum columbianum and A. m. sigillatum.[18] The ranges of subspecies are illustrated in Robert Stebbin's amphibian field guides.[19]

Range of A. macrodactylum[10] and ranges of sub-species distributions.[18]
Left: On the dorsal underside under its snout are chin glands that release hormones when males rub against females. These are located under the white flecks. Right: The male vent (excretory and reproductive opening) area has an enlarged anatomy and papillae (pictured). The female vent is not so large and often smoother without papillae folds.[20]

Physical appearance (phenotypes)

Summary of distinguishing skin patterns and morphological features for the subspecies include:[18][21]

A. m. croceum
Orange dorsal color on tail breaking into patches along black body and into tiny dots on head, often absent anterior to eyes. Sides have whitish flecks. Number of costal grooves equals 13.
A. m. columbianum
Yellow to tan dorsal stripe on black body, continuous blotches to spots along body ending in narrowed blotches with spot patterns distributed on the head. White flecks along the sides and underside remaining as separate small flecks. Number of vomerine teeth greater than 35.
A. m. krausei
Yellow to tan dorsal stripe, continuous blotches to spots along body ending in widened blotches with spot patterns distributed on the head. White flecks along the sides and underside remaining as separate small flecks. Number of vomerine teeth equaling 32. Number of costal grooves equals 12.
A. m. sigillatum
Wax yellow to tan dorsal stripe forming spotty to irregular shaped blotches along body ending in dots or specks of dorsal color on head. Number of vomerine teeth equals 44. Number of costal grooves equals 13.
A. m. macrodactylum
Citrine, dull citrine, to tan dorsal stripe that is diffuse and continuous along grayish body. Pattern ending in diffuse specks of stripe color or absent on head and snout. White flecks on sides sometimes coming together to form larger flecks. Number of vomerine teeth equaling 33, forming a distinguished transverse arc. Number of costal grooves equals 13.

Biogeography and genetics

Mitochondrial DNA analysis[11] identifies somewhat different ranges for the subspecies lineages.[11] The genetic analysis, for example, identifies an additional pattern of deep divergence in the eastern part of the range. The spatial distribution of populations and genetics of this species links spatially and historically through the interconnecting mountain and temperate valley systems of western North America.[11][22] The breeding fidelity of long-toed salamanders (philopatry) and other migratory behaviours reduce rates of dispersal among regions, such as within mountain basins. This aspect of their behavior restricts gene flow and increases the degree and rates of genetic differentiation. Genetic differentiation among regions is higher in the long-toed salamander than measured in most other vertebrate groups.[23] Natural breaks in the range of dispersal and migration occur where ecosystems grade into drier xeric low-lands (such as prairie climates) and at frozen or harsher terrain at high elevation extremes (2,200 meters (7,200 ft)).[24]

A. m. columbianum
Genetic evidence for the 'central' subspecies (A. m. columbianum) suggests that it does not extend north into British Columbia, but is restricted to the Blue and Wallowa Mountains of central to northeastern sections of Oregon. Populations are restricted to these areas by the Snake River Canyon (Idaho) to the east and low dry or xeric lands in the Madras basin to the west.[11]
A. m. macrodactylum
The 'coastal' or 'western' subspecies (A. m. macrodactylum) lineage extends north from northeastern California, across the Klamath Siskiyou Range, through the Willamette Valley, along the coastal mountain ranges, including the Cascade Mountains, and continuing north through British Columbia and up into Alaska.[11]
A. m. croceum
The Santa Cruz long-toed salamander (A. m. croceum) is most closely related to the 'coastal' or 'western' subspecies. This conclusion is the most parsimonious biogeographic explanation with nearest populations of A. m. macrodactylum separated by approximately 300 km across the Sacramento-San Joaquin River Delta, California.[10] The isolated endemic populations are listed as an endangered subspecies.[20] Based on the biogeography and molecular clock calibrations, this subspecies may have been separated from the remainder of the distribution since the Miocene, molecular clock calibrations estimating 13.9 million years of separation.[10]
A. m. krausei
The 'eastern' subspecies (A. m. krausei) range is distributed throughout the interior mountains, with the western extent of its range encroaching into the low-land areas of the central interior plateau of Washington and British Columbia and the eastern extent of its range pushing through Rocky Mountain valleys into the lowland foothills and prairies of Montana and Alberta.[25][11]
A. m. sigillatum
The traditional 'southern' subspecies (A. m. sigillatum) does not register a mitochondrial genetic identity.[11] This subspecies was identified by Ferguson as forming an intergrade with A. m. columbianum in south central Oregon.[18]

Thompson and Russell found another evolutionary lineage that originates in a glacially restricted area of the Salmon River Mountains, Idaho.[11] With the arrival of the Holocene interglacial, approximately 10,000 years ago, the Pleistocene glaciers receded and opened a migratory path linking these southern populations to northern areas where they currently overlap with A. m. krausei and co-migrated north into the Peace River (Canada) Valley.[11] Ferguson also noted an intergradation in the same geographic area, but between the morphological subspecies A. m. columbianum and A. m. krausei that run parallel to the Bitterroot and Selkirk ranges.[18] Thompson and Russell suggest that this contact zone is between two different subspecies lineages because the A. m. columbianum lineage is geographically isolated and restricted to the central Oregon Mountains.[11]

Description

The body of the long-toed salamander is dusky black with a dorsal stripe of tan, yellow, or olive-green. This stripe can also be broken up into a series of spots. The sides of the body can have fine white or pale blue flecks. The belly is dark-brown or sooty in color with white flecks. Root tubercles are present, but they are not quite as developed as other species, such as the tiger salamander.[19]

The eggs of this species look similar to those of the related northwestern salamander (A. gracile) and tiger salamander (A. tigrinum).[26] Like many amphibians, the eggs of the long-toed salamander are surrounded by a gelatinous capsule. This capsule is transparent, making the embryo visible during development.[19] Unlike A. gracile eggs, there are no visible signs of green algae, which makes egg jellies green in color. When in its egg, the long-toed salamander embryo is darker on top and whiter below compared to a tiger salamander embryo that is light brown to grey above and cream-colored on the bottom. The eggs are about 2 mm (0.08 in) or greater in diameter with a wide outer jelly layer.[26][27] Prior to hatching—both in the egg and as newborn larvae—they have balancers, which are thin skin protrusions sticking out the sides and supporting the head. The balancers eventually fall off and their external gills grow larger.[28] Once the balancers are lost the larvae are distinguished by the sharply pointed flaring of the gills. As the larvae mature and metamorphose, their limbs with digits become visible and the gills are resorbed.[26][28]

The skin of a larva is mottled with black, brown, and yellow pigmentation. Skin color changes as the larvae develop and pigment cells migrate and concentrate in different regions of the body. The pigment cells, called chromatophores, are derived from the neural crest. The three types of pigment chromatophores in salamanders include yellow xanthophores, black melanophores, and silvery iridiophores (or guanophores).[29][30] As the larvae mature, the melanophores concentrate along the body and provide the darker background. The yellow xanthophores arrange along the spine and on top of the limbs. The rest of the body is flecked with reflective iridiophores along the sides and underneath.[29][18]

As larvae metamorphose, they develop digits from their limb bud protrusions. A fully metamorphosed long-toed salamander has four digits on the front limbs and five digits on the rear limbs.[31] Its head is longer than it is wide, and the long outer fourth toe on the hind limb of mature larvae and adults distinguishes this species from others and is also the etymological origin of its specific epithet: macrodactylum (Greek makros = long and daktylos = toe).[32] The adult skin has a dark brown, dark grey, to black background with a yellow, green, or dull red blotchy stripe with dots and spots along the sides. Underneath the limbs, head, and body the salamander is white, pinkish, to brown with larger flecks of white and smaller flecks of yellow. Adults are typically 3.8–7.6 cm (1.5–3.0 in) long.[26][20]

Habitat and distribution

The long-toed salamander is an ecologically versatile species living in a variety of habitats, ranging from temperate rainforests, coniferous forests, montane riparian, sagebrush plains, red fir forest, semiarid sagebrush, cheatgrass plains, to alpine meadows along the rocky shores of mountain lakes.[19][18][25] Adults can be located in forested understory, hiding under coarse woody debris, rocks, and in small mammal burrows. During the spring breeding season, adults can be found under debris or in the shoreline shallows of rivers, streams, lakes, and ponds. Ephemeral waters are often frequented.[19]

This species is one of the most widely distributed salamanders in North America, second only to the tiger salamander. Its altitudinal range runs from sea level up to 2,800 meters (9,200 ft), spanning a wide variety of vegetational zones.[19][18][33][34][35] The range includes isolated endemic populations in Monterey Bay and Santa Cruz, California.[36] The distribution reconnects in northeastern Sierra Nevada running continuously along the Pacific Coast to Juneau, Alaska, with populations dotted along the Taku and Stikine River valleys. From the Pacific coast, the range extends longitudinally to the eastern foothills of the Rocky Mountains in Montana and Alberta.[37][21][38]

Ecology and life cycle

Long-toed salamander egg mass showing outer jelly layer connecting the mass and two inner capsules separating each egg.

Eggs

Like all amphibians, the life of a long-toed salamander begins as an egg. In the northern extent of its range, the eggs are laid in lumpy masses along grass, sticks, rocks, or the mucky substrate of a calm pond.[39] The number of eggs in a single mass ranges in size, possibly up to 110 eggs per cluster.[40] Females invest a significant amount of resources into egg production, with the ovaries accounting for over 50% of the body mass in the pre-breeding season. A maximum of 264 eggs have been found in a single female—a large number considering each egg is approximately 0.5 millimeters (0.02 in) in diameter.[41] The egg mass is held together by a gelatinous outer layer protecting the outer capsule of individual eggs.[42] The eggs are sometimes laid singly, especially in warmer climates south of the Canada and US border. The egg jellies contribute a yearly supply of biological material that supports the chemistry and nutrient dynamics of shallow-water aquatic ecosystems and adjacent forest ecosystems.[43] The eggs also provide habitat for water molds, also known as oomycetes.[44]

A. m. macrodactylum larva with its eponymous long fourth toes on the rear feet

Larvae

Larvae hatch from their egg casing in two to six weeks.[39] They are born carnivores, feeding instinctively on small invertebrates that move in their field of vision. Food items include small aquatic crustaceans (cladocerans, copepods and ostracods), aquatic dipterans and tadpoles.[45] As they develop, they naturally feed upon larger prey. To increase their chances for survival, some individuals grow bigger heads and become cannibals, and feed upon their own brood mates.[46]

Metamorphosis and juveniles

After the larvae grow and mature, for at least one season (the larval period lasts about four months on the Pacific coast),[37] they absorb their gills and metamorphose into terrestrial juveniles that roam the forest undergrowth. Metamorphosis has been reported as early as July at sea level,[47] for A. m. croceum in October to November and even January.[20] At higher elevations the larvae may overwinter, develop, and grow for an extra season before metamorphosing.[48] In lakes at higher elevations, the larvae can reach sizes of 47 millimeters (1.9 in) snout to vent length (SVL) at metamorphosis, but at lower elevations they develop faster and metamorphose when they reach 35–40 millimeters (1.4–1.6 in) SVL.[23]

Adults

As adults, long-toed salamanders often go unnoticed because they live a subterranean lifestyle digging, migrating, and feeding on the invertebrates in forest soils, decaying logs, small rodent burrows or rock fissures. The adult diet consists of insects, tadpoles, worms, beetles and small fish. Salamanders are preyed upon by garter snakes, small mammals, birds, and fish.[49] An adult may live 6–10 years, with the largest individuals weighing approximately 7.5 grams (0.26 oz), snout to vent lengths reaching 8 cm (3.1 in), and total lengths reaching 14 cm (5.5 in).[50][51]

Behaviour

Seasonal

The life history of the long-toed salamander varies greatly with elevation and climate. Seasonal dates of migration to and from the breeding ponds can be correlated with bouts of sustained rainfall, ice thaw, or snow melt sufficient to replenish the (often) seasonal ponds. Eggs may be spawned at low elevations as early as mid-February in southern Oregon,[52] from early January to July in northwestern Washington,[53] from January to March in southeastern Washington,[54] and from mid-April to early May in Waterton Lakes National Park, Alberta.[55] The timing of breeding can be highly variable; of notable mention, several egg masses in early stages of development were found on July 8, 1999 along the British Columbia provincial border outside Jasper, Alberta.[10] Adults migrate seasonally to return to their natal breeding ponds, with males arriving earlier and staying longer than females, and some individuals have been seen migrating along snow banks on warm spring days.[56] Gender differences (or sexual dimorphism) in this species are only apparent during the breeding season, when mature males display an enlarged or bulbous vent area.

Breeding

Long-toed salamanders gather during the breeding season under a log immediately near the shore of a pond. Notice the range of drab to bright skin colors.

The time of breeding depends on the elevation and latitude of the salamander's habitat. Generally, the lower-elevation salamanders breed in the fall, winter, and early spring. Higher-elevation salamanders breed in spring and early summer. In the higher climates especially, salamanders will enter ponds and lakes that still have ice floating.[19]

Adults aggregate in large numbers (>20 individuals) under rocks and logs along the immediate edge of the breeding sites and breed explosively over a few days.[20] Suitable breeding sites include small fish-free ponds, marshes, shallow lakes and other still-water wetlands.[57] Like other ambystomatid salamanders, they have evolved a characteristic courtship dance where they rub bodies and release pheromones from their chin gland prior to assuming a copulatory mating position. Once in position, the male deposits a spermatophore, which is a gooey stalk tipped with a packet of sperm, and walks the female forward to be inseminated. Males may mate more than once and may deposit as many as 15 spermatophores over the course of a five-hour period.[20][39] The courtship dance for the long-toed salamander is similar to other species of Ambystoma and very similar to A. jeffersonianum.[58][59] In the long-toed salamander, there is no rubbing or head-butting; the males directly approach females and grab on, while the females try to rapidly swim away.[59] The males clasp the female from behind the forelimbs and shake, a behavior called amplexus. Males sometimes clasp other amphibian species during breeding and shake them as well.[53] The male only grabs with the front limbs and never uses his hind limbs during the courtship dance as he rubs his chin side to side pressing down on the female's head. The female struggles but later becomes subdued. Males increase the tempo and motions, rubbing over the female's nostrils, sides, and sometimes the vent. When the female becomes quite docile the male moves forward with his tail positioned over her head, raised, and waving at the tip. If the female accepts the males courtship, the male directs her snout toward his vent region while both move forward stiffly with pelvic undulations. As the female follows, the male stops and deposits a spermatophore, and the female will move forward with the male to raise her tail and receive the sperm packet. The full courtship dance is rarely accomplished in the first attempt.[59] Females deposit their eggs a few days after mating.[20]

Energy storage and defense mechanisms

Long-toed salamander showing an autotomized tail stub
Long-toed salamander showing markings for tail regrowth after loss

In some lowland areas the adult salamanders will remain active all winter long, excluding cold spells. However, during the cold winter months in the northern parts of its range, the long-toed salamander burrows below the frost-line in a coarse substrate to hibernate in clusters of 8–14 individuals.[40][60] While hibernating, it survives on protein energy reserves that are stored in its skin and along its tail.[61] These proteins serve a secondary function as part of a mixture or concoction of skin secretions that is used for defense.[62] When threatened, the long-toed salamander will wave its tail and secrete an adhesive white milky substance that is noxious and likely poisonous.[55][63] The color of its skin can serve as a warning to predators (aposematism) that it will taste bad.[62] Its skin colors and patterns are diverse, ranging from a dark black to reddish brown background that is spotted or blotched by a pale-reddish-brown, pale-green, to a bright yellow stripe.[37][39] An adult may also drop part of its tail and slink away while the tail bit acts as a squirmy decoy; this is called autotomy.[64] The regeneration and regrowth of the tail is one example of the developmental physiology of amphibians that is of great interest to the medical profession.[65]

Conservation status

While the long-toed salamander is classified as least concern by the IUCN,[1] many forms of land development negatively affect the salamander's habitat and have put new perspectives and priorities into its conservation biology. Conservation priorities focus at the population level of diversity, which is declining at rates ten times that of species extinction.[66][67][68][69] Population level diversity is what provides ecosystem services,[70] such as the keystone role that salamanders play in the soil ecosystems, including the nutrient cycling that supports wetland and forested ecosystems.[71]

Two life-history features of amphibians are often cited as a reason why amphibians are good indicators of environmental health or 'canaries in the coal mine'. Like all amphibians, the long-toed salamander has both an aquatic and terrestrial life transition and semipermeable skin. Since they serve different ecological functions in the water than they do in land, the loss of one amphibian species is equivalent to the loss of two ecological species.[72] The second notion is that amphibians, such as long-toed salamanders,[73] are more susceptible to the absorption of pollutants because they naturally absorb water and oxygen through their skin. The validity of this special sensitivity to environmental pollutants, however, has been called into question.[74] The problem is more complex, because not all amphibians are equally susceptible to environmental damage because there is such a diverse array of life histories among species.[75]

Long-toed salamander populations are threatened by fragmentation, introduced species, and UV radiation. Forestry, roads, and other land developments have altered the environments that amphibians migrate to, and have increased mortality.[76] Places such as Waterton Lakes National Park have installed a road tunnel underpass to allow safe passage and to sustain the migration ecology of the species.[2] The distribution of the long-toed salamander overlaps extensively with the forestry industry, a dominant resource supporting the economy of British Columbia and the western United States. Long-toed salamanders will alter migration behaviour and are affected negatively by forestry practices not offering sizable management buffers and protections for the smaller wetlands where salamanders breed.[77][78] Populations near the Peace River Valley, Alberta, have been lost to the clearing and draining of wetlands for agriculture.[79] Trout introduced for the sport fisheries into once fishless lakes are also destroying long-toed salamander populations.[80] Introduced goldfish prey on the eggs and larvae of long-toed salamanders.[81] Increased exposure to UVB radiation is another factor being implicated in the global decline of amphibians and the long-toed salamander is also susceptible to this threat, which increases the incidence of deformities and reduces their survival and growth rates.[82][83][84]

The subspecies Ambystoma macrodactylum croceum (Santa Cruz Long-toed Salamander) is of particular concern and it was afforded protections in 1967 under the US Endangered Species Act.[85] This subspecies lives in a narrow range of habitat in Santa Cruz County and Monterey County, California. Prior to receiving protections, some few remaining populations were threatened by development. The subspecies is ecologically unique, having unique and irregular skin patterns on its back, a unique moisture tolerance, and it is also an endemic that is geographically isolated from the rest of the species range.[18][86][87][88] Other subspecies include A. m. columbianum, A. m. krausei, A. m. macrodactylum and A. m. sigillatum.[21]

See also

References

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Long-toed salamander: Brief Summary

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The long-toed salamander (Ambystoma macrodactylum) is a mole salamander in the family Ambystomatidae. This species, typically 4.1–8.9 cm (1.6–3.5 in) long when mature, is characterized by its mottled black, brown, and yellow pigmentation, and its long outer fourth toe on the hind limbs. Analysis of fossil records, genetics, and biogeography suggest A. macrodactylum and A. laterale are descended from a common ancestor that gained access to the western Cordillera with the loss of the mid-continental seaway toward the Paleocene.

The distribution of the long-toed salamander is primarily in the Pacific Northwest, with an altitudinal range of up to 2,800 m (9,200 ft). It lives in a variety of habitats, including temperate rainforests, coniferous forests, montane riparian zones, sagebrush plains, red fir forests, semiarid sagebrush, cheatgrass plains, and alpine meadows along the rocky shores of mountain lakes. It lives in slow-moving streams, ponds, and lakes during its aquatic breeding phase. The long-toed salamander hibernates during the cold winter months, surviving on energy reserves stored in the skin and tail.

The five subspecies have different genetic and ecological histories, phenotypically expressed in a range of color and skin patterns. Although the long-toed salamander is classified as a species of Least Concern by the IUCN, many forms of land development threaten and negatively affect the salamander's habitat.

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