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The marbled salamander (''Ambystoma opacum'') is a species of
mole salamander The mole salamanders (genus ''Ambystoma'') are a group of advanced salamanders endemic to North America. The group has become famous due to the presence of the axolotl (''A. mexicanum''), widely used in research due to its paedomorphosis, and t ...
found in the eastern
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...
.


Description

The marbled salamander is a stocky and boldly banded salamander. The marbled salamander exhibits sexual dimorphism with bands of females tending to be gray, while those of males are more white. Juveniles do not have any bands and have white flecks instead. Adults can grow to about 11 cm (4 in), small compared to other members of its genus. Like most of the mole salamanders, it is secretive, spending most of its life under logs or in burrows.


Habitat and range

Marbled salamanders are found in the eastern
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...
, from southern
New England New England is a region comprising six states in the Northeastern United States: Connecticut, Maine, Massachusetts, New Hampshire, Rhode Island, and Vermont. It is bordered by the state of New York to the west and by the Canadian provinces ...
to northern Florida, and west to Illinois and Texas. They have been found as far north as
New Hampshire New Hampshire is a U.S. state, state in the New England region of the northeastern United States. It is bordered by Massachusetts to the south, Vermont to the west, Maine and the Gulf of Maine to the east, and the Canadian province of Quebec t ...
, though only two sightings have been reported there. Their habitats are damp woodlands, forests, and places with soft and wet soil. Seasonally flooded areas are essential for breeding, but the salamanders do not normally enter the water. Like many salamanders, marbled salamanders have poison glands to deter predators. The marbled salamander is the state salamander of
North Carolina North Carolina () is a state in the Southeastern region of the United States. The state is the 28th largest and 9th-most populous of the United States. It is bordered by Virginia to the north, the Atlantic Ocean to the east, Georgia and So ...
.


Lifecycle and reproduction


Lifecycle

The first months that Marbled Salamanders spend living out of the water are the most important in determining how many will survive until the next breeding season.B.B.Rothermel and R.D. Semlitsch. Consequences of forest fragmentation for juvenile survival in spotted (Ambystoma maculatum) and marbled (Ambystoma opacum) salamanders. Canadian Journal of Zoology. 84(6): 797-807. https://doi.org/10.1139/z06-056 Marbled Salamanders are not strong burrowers, therefore they rely on existing holes in the ground for shelter. Desiccation, heat stress, soil moisture, temperature, and pH are all important factors in determining if a Marbled Salamander will survive. Chances of survival are low for Marbled Salamanders who travel through fields, however, they have been observed to traverse fields in order to find other pond areas. Marbled Salamanders survive best in a forest habitat, compared to an open field.TAYLOR, B.E., SCOTT, D.E. and GIBBONS, J.W. (2006), Catastrophic Reproductive Failure, Terrestrial Survival, and Persistence of the Marbled Salamander. Conservation Biology, 20: 792-801. https://doi.org/10.1111/j.1523-1739.2005.00321.x Protecting
wetland A wetland is a distinct ecosystem that is flooded or saturated by water, either permanently (for years or decades) or seasonally (for weeks or months). Flooding results in oxygen-free (anoxic) processes prevailing, especially in the soils. The ...
s is key to the survival of this species. Conservationists recommend leaving a buffer zone of forest around wetlands to increase survivorship of Marbled Salamanders. Marbled Salamanders in the northern portions of their range can also go into a state of torpor to survive the cold months. Adults spend most of their time in their burrows or under logs, as is the case with most mole salamanders. Juvenile marbled salamanders hatch early compared to most salamanders and gain a size advantage by feeding and growing for several months before the Jefferson salamanders and spotted salamanders hatch later in the spring. Larvae typically mature as quickly as two months in the southern part of their range, but take up to six months to mature in the northern part. Marbled salamanders, like other members of this genus, are reported to have relatively long life spans, 8–10 years or more.


Reproduction

Courtship of this species takes place on land. The males will compete by butting heads and blocking another male’s movement with its tail. When courting the female, a male will nudge the vent of a female with its snout, with the intent that the female will respond in kind. This back-and-forth nudging has the appearance of a dance as the two salamanders circle around one another. This display culminates with the male depositing a
spermatophore A spermatophore or sperm ampulla is a capsule or mass containing spermatozoa created by males of various animal species, especially salamanders and arthropods, and transferred in entirety to the female's ovipore during reproduction. Spermatophores ...
and the female moving to take it into her
cloaca In animal anatomy, a cloaca ( ), plural cloacae ( or ), is the posterior orifice that serves as the only opening for the digestive, reproductive, and urinary tracts (if present) of many vertebrate animals. All amphibians, reptiles and birds, a ...
. The female will then lay between 50 and 200 eggs, often remaining with them until the nest floods. One fairly unique parental care behavioral characteristic of Marbled Salamanders is that when the mothers do stay with their eggs, they wrap their bodies around the eggs to form a bowl shape. They do this when rain is coming to collect water over the eggs. For this species, water via rainfall or rising pond water levels must make extended contact with the eggs in order for them to begin hatching. However, it has been observed that females may abandon their eggs before flooding occurs.Petranka, James W. “Observations on Nest Site Selection, Nest Desertion, and Embryonic Survival in Marbled Salamanders.” Journal of Herpetology, vol. 24, no. 3, Society for the Study of Amphibians and Reptiles, 1990, pp. 229–34, https://doi.org/10.2307/1564387. Female Marbled Salamanders have a very low attachment to their eggs, and they will abandon their nest after a disturbance. They have also been observed to abandon undisturbed nests. When the mother leaves the nest, she leaves the eggs vulnerable to predation by other salamanders, frogs, and beetles. Reproductive success is highly variable for the Marbled Salamander. Some years many juveniles will survive, while other years the breeding population may experience a catastrophic failure, and very few juveniles will survive. These catastrophic failures occur randomly, but it has been found that they are mainly influenced by the length of the hydroperiod. A short hydroperiod is the main cause of catastrophic failure. Because Marbled Salamanders have relatively long life spans, their chances of extinction due to catastrophic failure are low. If they do not breed successfully one year, they will be alive the next year to try again. However, if there are other complications affecting their survival, the possibility of a catastrophic failure poses a larger threat to the overall population. Surviving on land, outside of the reproduction season, is very important to keep the population stable. While most Marbled Salamanders return to the pond where they were born to breed, some may travel over 1,000 meters to locate a new pond to breed.Gamble, Lloyd R., Kevin McGarigal, and Bradley W. Compton. "Fidelity and Dispersal in the Pond-Breeding Amphibian, Ambystoma Opacum: Implications for Spatio-Temporal Population Dynamics and Conservation." Biological Conservation, vol. 139, no. 3, 2007., pp. 247-257doi:https://doi.org/10.1016/j.biocon.2007.07.001. ID: 271811. This often occurs when their natal pond has a small population that may not have a large selection of mates. This dispersal helps populations of Marbled Salamanders to avoid genetic problems, by introducing new genes into the population. This dispersal also means that it is important to view these populations as a larger
metapopulation A metapopulation consists of a group of spatially separated populations of the same species which interact at some level. The term metapopulation was coined by Richard Levins in 1969 to describe a model of population dynamics of insect pests in ...
, rather than focusing simply on a single wetland area. Larval salamanders have been found to be positively phototactic until fully developing their rear legs, at which point they switch and become negatively phototactic.


Feeding

Adults take terrestrial invertebrates, such as worms, insects, centipedes, other arthropods, and
mollusk Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is e ...
s (snails, slugs). Larvae take small aquatic animals (
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
, mainly
copepod Copepods (; meaning "oar-feet") are a group of small crustaceans found in nearly every freshwater and saltwater habitat (ecology), habitat. Some species are planktonic (inhabiting sea waters), some are benthos, benthic (living on the ocean floor) ...
s and
cladoceran The Diplostraca or Cladocera, commonly known as water fleas, are a superorder (biology), superorder of small crustaceans that feed on microscopic chunks of organic matter (excluding some predatory forms). Over 1000 species have been recognised s ...
s), but larger individuals will take larger
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s (
isopod Isopoda is an order of crustaceans that includes woodlice and their relatives. Isopods live in the sea, in fresh water, or on land. All have rigid, segmented exoskeletons, two pairs of antennae, seven pairs of jointed limbs on the thorax, an ...
s,
fairy shrimp Anostraca is one of the four orders of crustaceans in the class Branchiopoda; its members are referred to as fairy shrimp. They live in vernal pools and hypersaline lakes across the world, and they have even been found in deserts, ice-covered mou ...
), aquatic insects, snails,
oligochaete worm Oligochaeta () is a subclass of animals in the phylum Annelida, which is made up of many types of aquatic and terrestrial worms, including all of the various earthworms. Specifically, oligochaetes comprise the terrestrial megadrile earthworms ...
s, and eggs and larvae of other
amphibian Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...
s, as well. Marbled Salamanders are considered a
keystone predator A keystone species is a species which has a disproportionately large effect on its natural environment relative to its abundance, a concept introduced in 1969 by the zoologist Robert T. Paine. Keystone species play a critical role in maintaini ...
because they alter the competitive ability of their prey, allowing other species of prey to thrive.


Predator avoidance

When A. opacum is under attack by a predator, they often exhibit tail lashing, head-butting, body coiling, or potentially becoming immobile. These defensive moves are thought to draw attention to the tail, which has granular glands that produce noxious secretions to protect themselves. While some predators have learned to eat the body of Marbled Salamanders and leave the tail, this is still a deterrent for many predators. A problem with the granular glands Marbled Salamanders possess is that secretions are reduced after multiple attacks, making them more vulnerable


References


Further reading

* Gravenhorst JLC. 1807. ''Vergleichende Uebersicht des Linneischen und einiger neuern zoologischen Systeme ... Nebst dem eingeschalteten Verzeichnisse der zoologischen Sammlung des Verfassers und den Beschreibungen neuer Thierarten, die in derselben sind''. Göttingen: Heinrich Dieterich. xx + 476 pp. (''Salamandra opaca'', new species, p. 431). (in
German German(s) may refer to: * Germany (of or related to) **Germania (historical use) * Germans, citizens of Germany, people of German ancestry, or native speakers of the German language ** For citizens of Germany, see also German nationality law **Ger ...
). *Petranka, James W. (1998). ''Salamanders of the United States and Canada''. Washington, District of Columbia: Smithsonian Books. 592 pp. . *Tyning, Thomas F. (1990). ''A Guide to Amphibians and Reptiles''. Stokes Nature Guides. New York: Little, Brown and Company. 416 pp. .


External links


Animal Diversity Web





ITIS
{{Taxonbar, from=Q1407596 Mole salamanders Salamander, Marbled Salamander, Marbled Amphibians described in 1807 Taxa named by Johann Ludwig Christian Gravenhorst Extant Pliocene first appearances Symbols of North Carolina