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Wildlife, Animals, and Plants
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BIOLOGICAL DATA AND HABITAT REQUIREMENTS
SPECIES: Ambystoma macrodactylum | Long-Toed Salamander
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].
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].
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].
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].
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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SPECIES: Ambystoma macrodactylum | Long-Toed Salamander
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