Ambystoma opacum (Gravenhorst, 1807)
Marbled Salamander (Ambystoma
opacum gallery)
Introduction & Physical Description
Ambystoma
opacum is a small, robust salamander, with thick
limbs, head, and tail. The dorsal coloration and pattern is
striking, consisting of a black background overlain with silver or
white solid and broken crossbars. The crossbars of males are
typically a brighter white color, while those of females are duller,
silver-gray. Crossbars number from 3-8 on the body, and 4-8 on the
tail, and costal grooves number 11-12 (Trauth et al., 2004). Adults can measure 3.0-5.0 inches in total
length.
Newly metamorphosed young are dark brown or black with white
flecks, and develop web-like whitish coloration about the dorsum
within 3 weeks of transformation. Adult coloration begins to appear
within 3 months after transformation.
Hatchling larvae are pond type, with short, robust bodies, long
gills, and high tail fins that extend past the limb insertions.
Hatchlings typically measure 10-14 mm (.40-.50 inches) in total
length, with more advanced embryos reaching up to 19 mm (.75 inches)
(Petranka, 1998). Larvae are black or brown dorsally, with a series
of dots along the sides of the abdomen, just below the level of the
limb bases (Petranka, 1998). Mottling is sometimes present in older
larvae, as well as various amounts of light green or yellowish
coloration. The ventral surface is mildly mottled with black
coloration, as is the throat.
When disturbed, A. opacum will assume a defensive posture
consisting of lowering the head, and raising and lashing the tail. A
noxious, milky secretion may also be produced from the tail.
There are no subspecies recognized, but pattern and coloration
vary immensely among populations. A. opacum is considered a
rather stable taxon, whose life history has been thoroughly examined
in previous years.
Synonymy & Etymology
Taxonomic Synonymy: Salamandra opaca (Gravenhorst, 1807),
Salamandra fasciata (Green, 1818), Salamandra gravenhorstii (Leuckard,
1826), Ambystoma opaca (Baird, 1850), Amblystoma opacum (Gray,
1850), Ambystoma faciatum (Dumeril, Bibron, and Dumeril, 1854),
Salamandra armigera (Dumeril, Bibron, and Dumeril, 1854), Ambistoma
fasciatum (Jan, 1857), Salamandroides gravenhorstii (Fitzinger,
1864), Ambystoma opacum (Dunn, 1918), Ambystoma opacum (Tihen, 1958)
Vernacular Synonymy: Marbled Salamander, Blotched
Salamander, Banded Salamander, Opaque Salamander
Etymology: The genus name Ambystoma is thought to
be Latin in origin, derived from the word anabystoma, and translating
to "to cram into mouth" (Beltz, 2004). It has also been
proposed that Ambystoma is simply a misprint of the Greek term
Amblystoma, which breaks down to amlys = blunt, and stoma = mouth (Beltz,
2004). The species name "opacum" is Latin, and means
shaded, dark, or obscure, and refers to the dark color pattern of
A. opacum (Beltz, 2004).
Distribution & Habitat
A. opacum is found on much of the eastern coast of the United
States, excluding most of New Hampshire and Florida, and all of
Maine. The known range includes southern New England, south to
northwestern Florida, and inland to eastern Texas, southeastern
Oklahoma, most of Arkansas, southeastern Missouri, southern
Illinois, and southwestern Indiana. Populations are absent from most
of western North Carolina, northeastern Tennessee, northeastern
Kentucky, central and western Virginia, eastern West Virginia, most
of northern Pennsylvania, and most of northern Ohio (Petranka,
1998). A. opacum may inhabit deciduous
forests, tallgrass prairie (in the western range), and floodplain
forests. New York populations have been observed in
uncharacteristically dry areas outside the breeding season (Petranka,
1998). Habitats may also include swamp areas, as well as sandy
woods and rocky hillsides (Trauth et al., 2004). For the most
part, adults and juveniles are fossorial, but often emerge after
rains during non-breeding times. A. opacum cannot create a
new burrow from scratch, but enlarge and occupy rodent burrows and
other suitable sites. A. opacum are territorial, and
aggressively attack conspecifics in their vicinity.
Breeding Behavior
A. opacum are biphasic, but court and
oviposit on land. Females deposit eggs in dried or partially dried
areas of temporary water sources, where they are later submerged by
rain waters. Eggs are brooded by females, and do not hatch until
flooded with water.
Migration to the breeding ponds occurs in later summer and fall,
and primarily on rainy nights. Northern populations tend to breed
earlier than southern populations, which is reversed in other
Ambystomid species.
Migrating males arrive at the ponds several days, to two weeks
prior to females. Adults in North Carolina and Rhode Island were
observed leaving the ponds at nearly the same point of entry,
indicating homing behavior (Petranka, 1998). The sex ratio varies
highly, but males always outnumber females. Evidence shows that
males often court females before reaching the nesting site (Petranka,
1998), after which both continue to the breeding ponds.
Courtship consists of the male nudging and lifting up other
adults of both sexes near the tail and cloacal region. Females
respond by nudging the cloacal region of males (Petranka, 1998).
Mating pairs move away from the other adults, and engage in a
circular dance, both nudging the other about the cloacal region.
After a time, the male will move along the body of the female,
undulating the tail and raising the body, and deposit a
spermatophore. The male will then attempt to lead the female over
the spermatorphore, where she will collect the sperm cap into her
cloaca. Females sometimes do not follow the male, instead nosing
about until a spermatophore is found, and positioning herself to
collect the sperm cap (Petranka, 1998).
Fertilized females select shallow nest sites in the dried or
partially dried floor areas of temporary ponds and ditches. Nests
may consist of rodent burrows, or shallow constructions just below
the leaf litter or other protective cover (Petranka, 1998). In
certain habitats, females may create communal nests with several
clutches (Petranka, 1998). At a site in Indiana, more than half of
adults observed nested communally, a behavior thought to result from
the unique aquatic habitat, which lacked leaf litter or other
protective materials that would normally cover the eggs (Palis,
1996).
Clutches vary in size from 50-200 eggs, and clutch size is
directly correlated to the snout-to-vent length of the female. Eggs
are deposited singly, but in clusters (Trauth et al., 2004), and the
female remains to brood, coiling around the eggs, and moving and
turning them over every now and then. Females may stay with their
clutches until the nest is flooded, although some may leave prior to
flooding. Fresh eggs measure between 1.9-2.6 mm, and increase in
size with development (Petranka, 1998). Larger eggs generally result
in larger hatchlings, but at the cost of the female producing a
smaller clutch. Eggs are ready to hatch at 9-15 days after
oviposition, but hatch only after inundated. The eggs become rather camouflaged
as they pick up dirt and debris from the surrounding environment.
The thick, sticky outer egg capsule protects the embryos from
dehydration prior to hatching, and may shrink or swell markedly
depending on the state of hydration (Petranka, 1998).
Hatching may occur 1-2 days after inundation in advanced-stage
eggs (Petranka, 1998). Hatching is stimulated by the reduction in
oxygen absorption experienced when covered in water. The lack of
oxygen triggers the release of digestive enzymes, from hatching
glands on the snout, that dissolve the egg capsules (Petranka,
1998). Hatching is based solely on environmental cues, which can
result in larvae hatching in both underdeveloped or advanced states,
depending on the amount of time allowed for development.
Larvae spend the winter developing, and reach maximum growth
rates in late winter and early spring, when the waters begin to
warm. Southern populations metamorphose primarily in March and
April, while the central and northern populations metamorphose a few
months later, in May and June (Petranka, 1998). The longer larval
period of the central and northern range is a result of slower
growth rates in the colder climates (Petranka, 1998). Larvae may
reach a maximum length of 30-72 mm at metamorphosis, depending on
the exact location and seasonal characteristics.
Metamorphosed juveniles can be found around breeding ponds
shortly after transforming, before dispersing on land. Sexual
maturity is reached within 1-5 years, at which point the salamanders
return to the same water source to reproduce.
The chosen depth of the nest in the pond varies among geographic
ranges, and depends on the annual rainfall characteristics in the
particular area. In ponds that fill slowly, the time of hatching is
related to the placement of the nests (Petranka, 1998). Those that
are placed in deeper areas hatch first, when early rains fill the
bottom of the pond, giving them a size advantage over later
hatchlings. However, such larvae run the risk of desiccation if the
following rains do not occur before the shallow water dries up.
Those placed at higher areas hatch last and will be the smallest
larvae in the pond, subjecting them to predation by older larvae.
Higher nest sites may also be subject to freezing, or desiccation if
the water level does not rise high enough to inundate the eggs (Petranka,
1998).
Studies have shown that females require a minimum moisture level
to stimulate breeding, and will not breed below the minimum
threshold (Petranka, 1998).
Again, females may brood clutches until the nests are flooded,
although deserted nests are often found. Females may also brood dead
eggs normally (Petranka, 1998). In some populations, the survival
rate of young is directly correlated to the time the female spent
brooding the clutch (Petranka, 1998). In general, the length of the
brooding period is related to the seasonal characteristics of the
particular habitat, and may last anywhere from a few weeks to
several months. Females that brood for several months often incur
significant loss of food reserves and energy.
References
Beltz, Ellin (2003). North American Reptile and Amphibian Names. Ellin
Beltz. http://ebeltz.net/herps/etyhome.html
(Accessed: 2004).
Frost, Darrel & AMNH. Amphibian Species of the World 3.0, an Online
Reference. American Museum of Natural History.
http://research.amnh.org/herpetology/amphibia/index.php.
(Accessed: 2004).
Obst, Fritz Jugen, Udo Jacob, and K. Richter. Completely Illustrated Atlas of Reptiles and Amphibians for the Terrarium.
Neptune City, NY: T.F.H. Publications, Inc., 1989.
Palis, J. G. 1996. Ambystoma opacum (marbled salamander). Communal
nesting. Herpetological Review 27:134.
Petranka, James W. Salamanders of the United States and Canada.
Smithsonian Institution Press, 1998.
Stebbins, Robert C., and Nathan W. Cohen. A Natural History of
Amphibians. Princeton Paperbacks, 1997.
Trauth, S. E., H. W. Robinson, and M. V. Plummer. The Amphibians and
Reptiles of Arkansas. Fayetteville, AK: University of Arkansas Press,
2004.
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