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MARBLED
SALAMANDER
Ambystoma opacum
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| INTRODUCTION |
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Marbled salamanders are one of two species
of mole (Ambystomid) salamanders
that have been detected in Rhode Island. Mole salamanders are
fossorial, therefore they are rarely observed above ground, with
the exception of their brief breeding season. A much more common
congener, spotted salamander (Ambystoma maculatum), is
widespread and found throughout the state. In contrast, marbled
salamanders have a relatively limited distribution; they are primarily
restricted to western and southern portions of the state.
The marbled salamander is a peculiar member of the amphibian community
in Rhode Island. It is the only species that breeds in the fall,
whereas, all other species breed in the spring or summer (Klemens
1993). Based on our fieldwork in Rhode Island, marbled salamanders
tend to prefer large, undisturbed tracts of forest, although we
also detected this species in a large forested tract of land in
Warwick (Paton and Egan 2001).
Klemens (1993) also found marbled salamanders were more prevalent
in rural areas of Connecticut, however, he also collected some
individuals in suburban and urban areas.
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| IDENTIFICATION |
- Marbled salamanders are the only black and white salamander
in Rhode Island.
- Adults are moderately sized, as their total length is about 7-11
cm [3-4"] (see specific details below).
- Their body is black, with white or light gray crossbands on the
head, back, and tail.
- Dorsal crossband colorations
differs between genders, males have silvery white crossbands,
whereas those of females are silvery gray.
- Gender is best determined just prior
to breeding (August), when adult males have swollen cloacal
glands, which have a squared off appearance. In contrast,
the cloaca of females tends
to be more rounded. After males deposit spermatophores,
the cloacal protuberance of males subsides and is less obvious.
- There are 11-13 costal
grooves.
- The venter is uniformly
bluish-black or brownish-black.
- There is much individual pattern variation. B. Shoop and T. Doty
(pers. comm.) have been photographing marbled salamanders for 10 years
to keep track of individuals at a breeding pond in western Rhode Island.
- Partially albinistic larvae have been reported once in Tiverton,
Rhode Island (Deegan
et al. 1998), which possessed
integumentary xanthophores
and orbital melanophores.
- Hatchlings are blackish and rather drab and have pond-type
morphology with bushy gills and dorsal
fins that extend almost to the front limbs (Petranka
1998a).
- Marbled salamander larvae have a larger head, a higher dorsal tail
fin, and more pointed tail than those of spotted salamanders.
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| SIZE |
| AGE / SEX
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SVL (SNOUT VENT LENGTH) (cm)
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SAMPLE SIZE
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AVERAGE
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RANGE
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Std. Deviation
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ADULT FEMALE
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6.0
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4.4- 7.4
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0.6
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210
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ADULT MALE
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5.4
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4.3- 6.7
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0.5
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158
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JUVENILE
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4.12
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3.0- 5.2
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0.6
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14
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METAMORPH
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3.6
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2.3- 5.0
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0.4
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254
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| AGE / SEX
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TL (TOTAL LENGTH)
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SAMPLE SIZE
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AVERAGE
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RANGE
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Std. Deviation
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ADULT FEMALE
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10.6
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8.2- 13.2
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0.9
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210
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ADULT MALE
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9.7
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8.0- 11.9
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0.8
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158
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JUVENILE
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7.4
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6.2- 8.5
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0.7
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14
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METAMORPH
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6.4
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3.8- 8.0
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0.8
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254
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| AGE/ SEX
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MASS (g)
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SAMPLE SIZE
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AVERAGE
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RANGE |
Std. Deviation
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ADULT FEMALE
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8.0
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3.4- 12.3 |
1.5
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176
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ADULT MALE
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5.5
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3.2- 8.6 |
1.2
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112
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JUVENILE
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3.1
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2- 4.1 |
0.9
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5
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METAMORPH
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1.5
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0.5- 3.1 |
0.42
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239
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| RELATIVE ABUNDANCE |
Rhode Island
During surveys
we conducted at breeding ponds west of Narragansett Bay, marbled
salamanders were somewhat uncommon, as they were detected at only
6 of 119 ponds (Paton and Egan 2001). . Marbled salamanders are
less likely to occur in suburban or urban areas- they are most common
in the rural western part of the state (Deegan et al. 1998). Our
fieldwork suggests they prefer landscapes that are dominated by
forests (Paton and Egan 2000). |
Regional
In Massachusetts,
which is at the northern limits of their range, the species has
been put on the state list of rare and endangered species (Klemens
1993). In contrast, they appear to be widespread and relatively
secure in Connecticut and western Rhode Island (Klemens 1993), although
somewhat localized in their distribution. In addition, they are
still relatively common on eastern Long Island and lower Westchester
County, New York. DeGraaf
and Yamasaki (2001) list them as uncommon in New England.
Marbled salamanders are one of the primary species investigated
at Rainbow Bay, South Carolina, where Semlitsch
et al. (1996) have been studied amphibian community
structure for over 16 years. The study documented tremendous interannual
variation in population sizes and productivity. The number of
breeding females captured annually ranged from none to over 600,
whereas the number of metamorphs emigrating from ponds ranged
from none (4 out of 16 years) to almost 9000.
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| DISTRIBUTION |
Rhode Island
Somewhat uncommon, mainly
restricted to the western and southern portions of the state. We
detected marbled salamanders at 6 of 119 randomly selected ponds
in the mainland area west of Narragansett Bay (Paton and Egan 2001).
Tom Doty and Bob Shoop monitored the population dynamics marbled
salamander at one breeding pond on the Alton Jones Campus in western
Rhode Island throughout the 1970s (Doty
1978) and during the 1990s. The number of breeding adults
has fluctuated throughout the years, but never been extirpated (T.
Doty, pers. comm.).
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Regional
Marbled salamanders
are at the northeastern limit of their range in New England. They
occur from extreme southeastern New Hampshire (Hoopes
1938), southeastern New York, and central Massachusetts
southward through the rest of New England (Klemens 1993). In Connecticut,
they are found throughout the state, but tend to be absent from
high elevations (above 1,100 ft [ 336 m] and most calcareous areas
(Klemens 1993). In New York State, marbled salamanders have been
documented on Long Island, Staten Island, and southeastern mainland
counties northward to the vicinity of Albany (Klemens 1993). Only
one specimen from Vermont and two specimens from New Hampshire have
been collected (Taylor
1993). |
General
Marbled salamanders
occur throughout much of the eastern deciduous forest from southern
New England to northern Florida and westward to the tallgrass prairie
(Petranka 1998a). They appear to primarily be a species of southeastern
North America, as their populations reach very high densities there
(Semlitsch et al. 1996). They are absent from many areas including
portions of the Appalachian Plateau, central Ohio, central Indiana,
most of Florida, southeastern Georgia, and coastal Louisiana (Klemens
1993). Disjunct populations occur along the southern Great Lakes
and in Missouri. |
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| HABITAT NON BREEDING |
| During the non-breeding
season, marbled salamanders are sometimes found under logs and rocks.
They are fossorial, so they probably hibernate in deep burrows,
such as old shrew (Sorex spp.) burrows. We occasionally captured
adult marbled salamanders in early June near breeding ponds, but
they are generally only caught from Aug-Oct. Unfortunately, little
is known about their habitat requirements during the non-breeding
season in Rhode Island, or elsewhere for that matter. |
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| HABITAT BREEDING |
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In Rhode Island, marbled salamanders typically breed in ponds
found in mixed deciduous or coniferous forest stands (e.g., both
white pine (Pinus strobus) and pitch pine (Pinus rigida).
This species appears to prefer dry, friable soils including sand
and gravel deposits, as well as rocky slopes, although they are
sometimes found in low-lying swampy areas (Klemens 1993). Marbled
salamanders are thought to inhabit somewhat drier areas than other
species of Ambystoma (DeGraaf and Yamasaki 2001); however,
in Ohio they are usually found in swampy forests (Ptingsten
and Downs 1989). Breeding occurs in temporary wetlands,
including wooded swamps, kettle ponds, and along river floodplains.
In Rhode Island, such sites generally contain red maple (Acer
rubrum) and pin oak (Quercus palustris) (Raithel unpubl.
data).
In addition to needing extensive forests, the presence of sphagnum
moss in the pond basin is possibly another important habitat characteristic
useful for detecting potential breeding ponds. Egg masses are
generally deposited under sphagnum in dried-up pools and guarded
by females until ponds until water levels inundate the eggs (T.
Doty, pers. comm.).
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| HYDROLOGY |
Marbled salamanders tend to
be most abundant in temporary ponds, which dry out completely each
season. Such sites usually do not have much standing vegetation
and are shared commonly with wood frogs (Rana sylvatica)
and to a lesser extent with spotted salamanders (Ambystoma maculatum)
(Raithel unpubl. data).
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| MOVEMENT CHRONOLOGY |
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In Rhode Island, adults start to immigrate to breeding ponds
around 1 August. Adults are generally only active on rainy nights.
The majority of adults are at breeding ponds by 26 August, with
some stragglers arriving as late as 10 September (Paton
and Crouch In press). Males often court females before
arriving at the nesting sites and males often arrive at the ponds
from a few days to more than 2 weeks before females (Petranka
1998a). Adults in northern populations tend to breed earlier than
those in southern populations (Petranka 1998a). Klemens (1993)
documented peak breeding in Connecticut from mid-September to
early October. Males tend to leave the ponds in early October,
while females remain in ponds for several weeks to brood eggs.
Females generally do not leave ponds until late October or early
November in Rhode Island (Paton and Crouch In press).
Metamorphs emigrate from breeding pond between mid-June and mid-July
(Paton and Crouch In press).
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| REPRODUCTION |
| Marbled salamanders are
one of only two Ambystoma salamanders that mate and oviposits their
eggs on land, all other species use aquatic breeding sites (Petranka
1998a). Marbled salamanders migrate to dried-up pond basins during
August in Rhode Island, with males generally preceding females.
Once females arrive, males initiate courtship by quickly moving
around, bumping into females (and males), and lifting them near
the tail and cloacal region (Petranka 1998a). After a protracted
dance, a male and female pair moves around in circles, while keeping
their cloacal regions in contact. Males eventually deposit a spermatophore
on the ground. A female positions her cloaca over the sperm cap
and picks up seminal fluid from the top of the spermatophore (Petranka
1998a). |
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| EGG MASS |
Eggs are oviposited singly
in small depressions usually beneath a sheltering object such as
logs, bark, leaf mold, or other debris (Green and Pauley 1987).
An average clutch size consists of approximately 100 eggs with a
large range from 50 to 232 (DeGraaf and Yamasaki 2001). Having deposited
the eggs, the females coil around the clutch and broods it. Females
remain with the eggs until hatching, which may take one-half month
to several months. Eggs will not hatch until inundated with water,
so the brooding period is a function of the chronology of pond filling
(Petranka 1998a).
According to Jackson et al. (1989),
females tend to select oviposition sites with high vegetative cover
and tend to concentrate in the lower portions of pond basins in
South Carolina. Nest success was highest for nests in deepest parts
of the basin, where nests were located randomly. In contrast, nests
at mid to high elevations were non-randomly aggregated.
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| LARVAE |
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Typically, eggs hatch in early November as ponds refill
and eggs are flooded. The larvae then overwinter in ponds,
although they do not grow much. As pond temperatures warm
in the spring and invertebrates become more available, metamorphosis
takes place much more rapidly. A. opacum near metamorphosis
in late April and early May when the larvae of spring breeding
species such as A. maculatum hatch. (Klemens 1993).
Larvae are aquatic and primarily nocturnal (DeGraaf and
Yamasaki 2001). As pond temperatures warm in the spring
and invertebrates become more available, metamorphosis takes
places much more rapidly. In New Jersey, the larval period
is 135 days (Hassinger et al. 1970).
Hatchlings and small larvae tend to congregate in leaf
litter in warm, shallow water during the day. We have been
relatively successful dip netting for larvae in the middle
of the day at pond edges (Paton and Egan 2001). At night,
they disperse more evenly throughout the ponds and feed
in the water column (Petranka 1998a). Occasionally, recently
transformed individuals can be found beneath litter and
debris around the margins of breeding ponds, but typically,
metamorphs disperse from ponds during rainy weather soon
after transforming (Petranka 1998a).
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Marbled salamander larva
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| METAMORPHS |
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In Rhode Island, peak emigration of metamorphs takes place
from about mid-June through 1 July, with metamorphs leaving
the pond as late as late July (Paton and Crouch In press).
This is slightly later than reported by Petranka (1998a)
who suggested that metamorphosis occurs primarily in May
and June in central and northern areas of the range, and
in March and April in the south. The length of larval period
varies depending on climatic patterns and the time of hatching
(Petranka 1998a).
A note of caution to biologists working in ponds in Rhode
Island. Not all larval salamanders seen in early spring
(e.g., January-March) are A. opacum, as occasionally A.
maculatum will overwinter in vernal ponds in Rhode Island
(Whitford and Vinegar; S.
Egan, unpubl. data).
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| JUVENILES |
Recently transformed juveniles
are brown or black with light flecks, and the flecks become more
pronounced and form lichen-like patterns within 1-3 weeks after
metamorphosis (Petranka 1998a). Klemens (1993) felt newly transformed
and young salamanders are suffused with a purplish gray pattern,
which later coalesces into crossbands. Juveniles begin developing
the adult pattern within 1-2 months after transformation (Bishop
1941)
Sexual maturity: 15-18 months (Degraaf and Yamasaki 2001).
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| FOOD |
Adults
Adult feed on Arthropods,
including adults and larval insects, and crustaceans. They also
take earthworms and mollusks (DeGraaf and Yamasaki 2001). Bishop
(1941) reported that adults ate earthworms, snails, slugs, staphylinid
beetles, ants, orthoptera, and diptera and that larvae ate small
crustaceans and snails. |
Larvae
Larvae eat small
aquatic insects, crustaceans, and other small invertebrates, and
are cannibalistic. In addition, once wood frog eggs hatch, they
readily feed on larvae wood frogs and spotted salamanders (Klemens
1993), which increases their growth rate dramatically. During that
short period that both species share the pond, the larger A.
opacum larvae exploit different invertebrate food resources
than the minute A. maculatum. |
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| PREDATION |
Adults
{Predation Adults}
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Larvae
{Predation Larvae/Tadpoles} |
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| CONSERVATION CONCERNS |
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Marbled salamanders are listed as a threatened species in Massachusetts,
and Lazell (1979) attributed their
drastic declines in Massachusetts to a combination of factors
including pollution, habitat fragmentation, and acid rain. However,
Klemens (1993) mentioned that Massachusetts is the northern limits
of their range, and declines in the region may be due a to range
retraction. In the southeastern United States, the greatest environmental
threat to this species is loss of bottomland hardwoods and associated
vernal pool habitats (Petranka 1998a). Thousands of local populations
of marbled salamanders have already been eliminated by habitat
loss, and more will be lost in the future. In New England, urban
sprawl has eliminated marbled salamanders from large portions
of their range on western Long Island and lower Westchester County,
and they probably will become scarcer on eastern Long Island and
in the Hudson Valley as urbanization continues (Klemens 1993).
Pechman et al. (1991)
found that marbled salamander populations act as metapopulations,
therefore some contact between sites must be maintained for population
persistence. Isolation of populations is invariably the precursor
of extirpation. Taylor and Scott
(1997) summarized existing demographic data for marbled salamanders
and suggested that at least 70% adult survivorship, or an immigration
event from adjacent populations, was necessary for population
persistence. High adult survivorship is necessary because recruitment
is so sporadic
In Rhode Island, marbled salamander populations appear to be
relatively secure, at least for the short term (Klemens 1993).
However, if suburbanization of the western sections of the state
continues, populations could easily be extirpated. Raithel (unpubl.
data) observed high rates of adult and juvenile mortality caused
by auto traffic, which must negatively affect populations. Other
factors affecting marbled salamanders, including increased environmental
acidity and pollution, may impact populations (Lazell 1979), but
habitat loss and fragmentation is probably the greatest threat
to marbled salamanders in Rhode Island. At the Alton Jones campus,
the marbled salamander population remains relatively stable at
one breeding pond in a relatively unfragmented, mixed deciduous/coniferous
forest over 1000 acres in size (T. Doty and B. Shoop, pers. comm.).
This same area has been aerial sprayed twice with Carbaryl (Sevin)
(Doty 1978), often gets fairly acidic rainfall, and has had outbreaks
of the bacteria Aeromonas hydrophila, which can negatively impact
wood frog productivity (Nyman 1985).
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