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Ambystoma cingulatumFlatwoods Salamander(Also: Frosted Flatwoods Salamander)

By Diana Nickle

Geographic Range

Flatwoods salamanders, Ambystoma cingulatum, inhabit the southern Atlantic coast of the United States ranging from the middle of South Carolina to the panhandle of Florida. They span as far west as the Apalachicola-Flint Rivers in northern Florida, but not crossing the Appalachians in South Carolina or Georgia. Flatwoods salamander individuals may travel up to 1700 meters within their geographic range to breed in the winter, but otherwise will never stray more than a couple meters from their underground burrows. (Bishop, et al., 2006; Jones, et al., 2012; Lannoo, 2005; Palis and Aresco, 2007; Semlitsch, et al., 2017)

Habitat

Flatwoods salamanders inhabit wet terrestrial environments with breeding sites that include vernal pools, roadside ditches, cypress or other forest swamps, marshes, and sphagnum patches. While mainly staying in/near freshwater, these salamanders can tolerate low salt concentrations. Non-breeding sites include heavily fire-dependent pine flatwoods and include a range of grasses and trees like wiregrass Aristida stricta, sawgrass Cladium, longleaf pines Pinus palustris, sand pines Pinus clausa and slash pines Pinus elliottii.

Flatwoods salamanders have been found up to 125 meters in elevation and below sea level in coastal areas. They live in a moderately temperate environment and can survive in temperatures ranging from 2 degrees Celsius to 27 degrees Celsius; if temperatures reach extremes, they will burrow into the wet sandy soil. While they spend most of the year underground, flatwoods salamanders will migrate to areas of standing water in the winter and will emigrate back in the early spring to their burrows. These burrows may be former crayfish burrows (Anderson and Williamson, 1976; Haas, 2005; Haas, et al., 2009; Haas, et al., 2013; Jones, et al., 2012; Lannoo, 2005; Means, et al., 1996; Semlitsch, et al., 2017; Westervelt, et al., 2013)

  • Aquatic Biomes
  • lakes and ponds
  • temporary pools
  • Range elevation
    0 to 125 m
    0.00 to 410.10 ft

Physical Description

Flatwoods salamanders average 15 costal grooves (range 13-16). They have a black or dark gray base color with white or silver reticulations, spots and/or stripes ranging from their nose to the tip of their tail. The underside is less striking by only having small white specks on the black or dark gray base. These salamanders have a long, rounded tail which is used for defense and reproductive purposes. As adults, they range from 9 to 13.5 centimeters in total length. Weight of adults vary from 2.5 - 10g. Palis et al. (2006) reported adult males to weigh 4.4 +/- 1.1g, gravid (with eggs) females to weigh 8.0 +/- 2.5 g, and non-gravid females to weigh 3.9 +/- 1.3 g. Females, on average, are larger in size while males are more colorful with brighter flecks of white or silver. Like most salamanders, flatwoods salamanders have skin that secretes mucus.

In the larval stage, they hatch at a total length of 10 mm and are dark brown with a greenish tint, with a lighter brown horizontal strip down the ventricle of its body. Flatwoods salamanders have large external bushy gills that assist in oxygen consumption while they are in the larvae stage. At lengths of 35-45 mm, they begin their metamorphosis to their adult stage.

Flatwoods salamanders, also know as frosted flatwoods salamanders, used to be part of the same species with reticulated flatwoods salamanders (Ambystoma bishopi). Flatwoods salamanders can be differentiated by having more costal grooves and having relatively longer limbs and a relatively bigger head. Males of the species have been reported to have relatively longer tails than male reticulated salamanders. ("Federal Register: Endangered and threatened wildlife and plants; Determination of endangered status for reticulated flatwoods salamander; Designation of critical habitat for frosted flatwoods salamander and reticulated flatwoods salamander; Final rule", 2009; Means, et al., 1996; Palis, et al., 2006; Pauly, et al., 2011; Whiles, et al., 2004)

  • Sexual Dimorphism
  • female larger
  • male more colorful
  • Range mass
    2.5 to 10 g
    0.09 to 0.35 oz
  • Range length
    9 to 13.5 cm
    3.54 to 5.31 in

Development

Female flatwoods salamander lays the group of eggs in a dry pond or under Sphagnum moss, but they will not hatch until the area is inundated. They can stay inactive for a matter of weeks if flooding does not occur. Then the salamander larvae will emerge within an hour. Measured to be as small as 9.6 mm and as large as 14.8 mm immediately after hatching they average around 10-12 mm long. They are dark brown on the dorsal side and a lighter brown on the ventral side. The larvae have a faint tan lateral stripe when first hatched and develop a gold stripe that runs vertical from their gills to their tail as they mature. They begin to form limbs and the torso elongates about a month into development. They lose their external gills and their recognizable silver specks begin to appear as they reach full maturity in the spring and completely metamorphose into an adult. Some larva will metamorphose in a year, but some populations are known to take 2 years to reach adulthood. Only one female was identified in 8 groups of salamander eggs (Anderson and Williamson 1976), one might conclude that the male gamete is dominant and that sex could be determined by the male parent. (Anderson and Williamson, 1976; Haas, et al., 2009; Lannoo, 2005)

Reproduction

Flatwoods salamanders are polygynandrous and will migrate to common terrestrial areas for breeding from October to early December. Males will secrete pheromones using their mental glands located under their chin to attract females. After mating the males and females will remain in the area until late December to early January when they emigrate back to their habitats that they inhabit for the remainder of the year. While little is known about the mating habits because of their solitary nature, it can be concluded that males are non-territorial because of their migration to neutral areas for reproduction purposes. (Anderson and Williamson, 1976; Haas, et al., 2009; Lannoo, 2005)

Although some female flatwoods salamanders can mature at one years of age, some populations take two years. Males mature at one. They will mate only once a year trekking to their yearly breeding grounds from late October to early December. Eggs are laid in herbaceous areas in roadside ditches, dry pond basins, under logs or in sphagnum mats. These areas are known to be covered in Liquidambar stryaciflua (sweetgum trees), Pinus eliotti (slash pines), Pinus palustris (longleaf pines) and Aristida stricta (wiregrass). Females will lay small groups of eggs ranging from 1-34 and averaging at 7.2; the eggs are usually laid in a linear or clumped fashion. It takes 2 weeks for the embryos to develop, but most of the time they won't emerge as soon as they are developmentally mature. After substantial rainfall the area begins to flood. Due to this inundation flatwoods salamanders will almost immediately hatch and move into the water. They can hatch as early as 9 days and as late as 60 days (average = 39 days), depending on rainfall. The parents will remain in the area burrowed underground until they emigrate back to their yearly habitats in late December to January. Hatchlings are independent upon hatching. (Anderson and Williamson, 1976; Haas, et al., 2009; Lannoo, 2005)

  • Breeding interval
    Flatwoods salamanders breed once yearly.
  • Breeding season
    Mating for the flatwoods salamander begins in late October to early December.
  • Range number of offspring
    1 to 34
  • Average number of offspring
    7.2
  • Range time to hatching
    9 to 60 days
  • Average time to hatching
    39 days
  • Range time to independence
    0 to 0 minutes
  • Range age at sexual or reproductive maturity (female)
    1 to 2 years
  • Average age at sexual or reproductive maturity (male)
    1 years

Flatwoods salamanders are independent at hatching. After the female lays her eggs, she slowly burrows into the surrounding area and goes dormant for up to 38 days. The male will also burrow and remain dormant for a period of time. After the rainfall has saturated the area, the adult flatwoods salamanders will emigrate back to their non-breeding upland habitats. (Anderson and Williamson, 1976; Haas, et al., 2009; Lannoo, 2005)

  • Parental Investment
  • no parental involvement

Lifespan/Longevity

Flatwoods salamanders live in captivity up to 4 years. It is unknown how long they can survive in the wild, but considering a female does not reach maturity until 2 years of age, and is expected to lay multiple clutches in her lifetime, 4 to 5 years is assumed. Batrachochytrium dendrobatidis or chytrid fungus may decrease lifespans of local populations. A recent study found Hedruris siredonis (a parasitic nematode) in the stomachs of flatwoods salamanders, and with the introduction of non-native predatory fish species, freshwater crustacean populations have plummeted. Flatwoods salamander main diet includes these freshwater crustaceans and starvation has begun to affect multiple populations. (Anderson and Williamson, 1976; Bishop, et al., 2006; Lannoo, 2005)

  • Range lifespan
    Status: captivity
    2 to 4 years

Behavior

Flatwoods salamanders are a solitary species. They spend almost entire year alone but will migrate to reproduce in the fall. Flatwoods salamander individuals may travel up to 1700 meters breed in the winter, but otherwise will never stray more than a couple meters from their underground burrows. They travel to breed in water bodies. They are known to burrow underground for most of the year but year-round activity suggests that they do not hibernate.

They are believed to be nocturnal; during a rain event in Florida, flatwoods salamanders were equally active from 1800 h until midnight and from midnight until 0700 h. During the heat of summer, they are believed to spend the entirety of their time in their underground burrows.

They are known to communicate with chemicals and use glandular secretions an an anti-predator defense. (Jones, et al., 2012; Lannoo, 2005)

Home Range

According to Lannoo (2005), only one study examined the home range of a single individual. Ashton (1992; cited in Lannoo) reported the home range of this individual flatwoods salamander to be 1500 square meters. These salamanders are not believed to be territorial, although this question has not been addressed scientifically. (Lannoo, 2005)

Communication and Perception

Little is currently known about how flatwoods salamanders communicate. Genetic research speculates the use of specialized organelles for tactile communication. The salamanders will a secrete distasteful mucus on the underside of their tail if threatened, they will then coil their heads under their tail to deter the predator. Occasionally they will lash their tails as warning but this is a rare occurrence.

Although there are no studies on the visual importance of this species, one can conclude that because of their ability to lash their tail, they must be able to have some sort of vision. Like most salamanders, flatwoods salamanders have glands on the underside of their chin (mental gland) that secrete pheromones. Males rub these glands on the head of the female, stimulating her to mate. (Jones, et al., 2012; Lannoo, 2005)

Food Habits

The flatwoods salamander diet consists of small freshwater invertebrates, with isopods and amphipods. Mature flatwoods salamanders seem to coexist with Aristida stricta (wiregrass) and although never proven, their climbing behavior around the grass implies that they may eat it in times of need. They also eat other salamander larva and were primarily raised in research labs on earthworm diets (which may not be a stable part of their diet elsewhere). With the introduction of predatory fish with the same diet, flatwoods salamanders may also eat small insects and other worms to avoid starvation. (Jones, et al., 2012; Lannoo, 2005; Whiles, et al., 2004)

  • Animal Foods
  • insects
  • terrestrial non-insect arthropods
  • terrestrial worms
  • other marine invertebrates

Predation

Predators are only a real threat to flatwoods salamanders during the larval stage. Many small pools and vegetation are available during the wet season and numerous species of fish will eat or out-compete the larvae for food. One adaptation flatwoods salamanders use to deter predators is their camouflage. As a larvae, their coloring is similar to the pond's surroundings helping to hide from predatory fish. As mature adults, the bright white-ish spots and dark background will help them almost completely disappear into the brush and sandy terrain. A gland located under the tail and will secrete a distasteful and bad smelling aroma that will turn away most predatory animals. Adults may also lash their tail as a sign of warning. (Anderson and Williamson, 1976; Lannoo, 2005)

  • Anti-predator Adaptations
  • cryptic

Ecosystem Roles

Parasites of flatwoods salamanders include a nematode Hedrus siredonis and chytrid fungus Batrachochytrium dendrobatidis. Chytrid fungus can be transmitted through direct contact between amphibians in areas of low water. Flatwoods salamanders have been known to carry this disease and transmit it to other organisms. The nematode uses flatwoods salamanders as a host and primarily feeds on their stomach contents. (Green, et al., 2002)

Commensal/Parasitic Species
  • nematode Hedruris siredonis
  • Chytrid fungus Batrachochytrium dendrobatidis

Economic Importance for Humans: Positive

Flatwoods salamanders do not have any known positive economic importance for humans.

Economic Importance for Humans: Negative

Flatwoods salamanders do not have any known negative economic impact on humans.

Conservation Status

Frosted flatwoods salamanders are listed as "Vulnerable" on the IUCN Red List. They are also listed as a federally threatened species. They have no special status through CITES or the State of Michigan List. They are state-endangered in South Carolina and are believed to be extinct in Alabama. Georgia lists them as state-threatened, and Florida considers them a species of special concern, although theses statuses were published before flatwoods salamanders were federally listed. None of these state listings afforded them much protections. Heavy development in areas of Florida, and eradication of natural longleaf pine Pinus palustris forests (reduced 97% in land area) all negatively impacted these salamanders. Longleaf pine ecosystem restorations efforts are in place and natural fire regimes partly restored. These controlled fires allow for reduction of woody vegetation, increase in herbaceous ground cover, increases in water temperatures and oxygen levels, and increased invertebrate prey.

At one time, it was estimated that less than 25 populations were left in the wild. Recent studies show that 27 have been observed: 6 in Georgia, 3 in South Carolina and 18 in Florida. However, most populations are isolated and metapopulation analyses show that 40% of known salamander populations depend entirely on a single, unconnected pond (different ponds for each population) for breeding.

Connecting isolated breeding pools to permanent waterbodies has introduced non-native predatory fish that eat the same diet as flatwood salamanders. Continued fire suppression and habit conversion on private lands (12% of known salamander populations), combined with threats from chytrid fungus and iridovirus, only make the threats worse.

Efforts to restore fire-maintained longleaf pine ecosystems are in place for public lands. Stable longleaf pine systems, coupled with access to wet, isolated vernal pools appears to the goal of conservation efforts. Avoiding further human disturbance (dredging, connecting pools to permanent waterbodies) also is a goal. ("Federal Register: Endangered and threatened wildlife and plants; Determination of endangered status for reticulated flatwoods salamander; Designation of critical habitat for frosted flatwoods salamander and reticulated flatwoods salamander; Final rule", 2009; Haas, et al., 2013; Palis and Hammerson, 2008; Pauly, et al., 2011)

Contributors

Diana Nickle (author), Radford University, Alex Atwood (editor), Radford University, Karen Powers (editor), Radford University, Joshua Turner (editor), Radford University, Tanya Dewey (editor), University of Michigan-Ann Arbor.

Glossary

Nearctic

living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.

World Map

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

bog

a wetland area rich in accumulated plant material and with acidic soils surrounding a body of open water. Bogs have a flora dominated by sedges, heaths, and sphagnum.

carnivore

an animal that mainly eats meat

chemical

uses smells or other chemicals to communicate

cryptic

having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.

ectothermic

animals which must use heat acquired from the environment and behavioral adaptations to regulate body temperature

fertilization

union of egg and spermatozoan

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

fossorial

Referring to a burrowing life-style or behavior, specialized for digging or burrowing.

freshwater

mainly lives in water that is not salty.

insectivore

An animal that eats mainly insects or spiders.

internal fertilization

fertilization takes place within the female's body

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

marsh

marshes are wetland areas often dominated by grasses and reeds.

metamorphosis

A large change in the shape or structure of an animal that happens as the animal grows. In insects, "incomplete metamorphosis" is when young animals are similar to adults and change gradually into the adult form, and "complete metamorphosis" is when there is a profound change between larval and adult forms. Butterflies have complete metamorphosis, grasshoppers have incomplete metamorphosis.

migratory

makes seasonal movements between breeding and wintering grounds

motile

having the capacity to move from one place to another.

native range

the area in which the animal is naturally found, the region in which it is endemic.

nocturnal

active during the night

oviparous

reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.

pheromones

chemicals released into air or water that are detected by and responded to by other animals of the same species

polygynandrous

the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.

seasonal breeding

breeding is confined to a particular season

sedentary

remains in the same area

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

solitary

lives alone

swamp

a wetland area that may be permanently or intermittently covered in water, often dominated by woody vegetation.

tactile

uses touch to communicate

temperate

that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).

threatened

The term is used in the 1994 IUCN Red List of Threatened Animals to refer collectively to species categorized as Endangered (E), Vulnerable (V), Rare (R), Indeterminate (I), or Insufficiently Known (K) and in the 1996 IUCN Red List of Threatened Animals to refer collectively to species categorized as Critically Endangered (CR), Endangered (EN), or Vulnerable (VU).

visual

uses sight to communicate

References

U.S. Fish and Wildlife Service. Federal Register: Endangered and threatened wildlife and plants; Determination of endangered status for reticulated flatwoods salamander; Designation of critical habitat for frosted flatwoods salamander and reticulated flatwoods salamander; Final rule. 50 CFR Part 17; Volume 74, Number 26. Washington, DC: U.S. Fish and Wildlife Service. 2009. Accessed November 19, 2017 at https://www.gpo.gov/fdsys/pkg/FR-2009-02-10/pdf/E9-2403.pdf.

Anderson, J., G. Williamson. 1976. Terrestrial mode of reproduction in Ambystoma cingulatum. Herpetologica, 32/2: 214-221.

Bevelhimer, M., D. Stevenson, N. Giffen, K. Ravenscroft. 2008. Annual surveys of larval Ambystoma cingulatum reveal large differences in pond residency. Southeastern Naturalist, 7/2: 311-322.

Bishop, D., J. Palis, K. Enge, P. David, D. Stevenson. 2006. Capture rate, body size, and survey recommendations for larval Ambystoma cingulatum (flatwoods salamander). Southeastern Naturalist, 5/1: 9-16.

Green, D., K. Converse, A. Schrader. 2002. Epizootiology of sixty-four amphibian morbidity and mortality events in the USA, 1996-2001. Annals of the New Academy of Sciences, 969: 323-339.

Haas, C. 2005. Burning trends and potential negative effects of suppressing wetland fires on flatwoods salamanders. Natural Areas Journal, 25/3: 290-294.

Haas, C., T. Gorman, D. Bishop. 2009. Factors related to occupancy of breeding wetlands by flatwoods salamander larvae. Wetlands, 29/1: 323-329.

Haas, C., T. Gorman, J. Himes. 2013. Evaluating methods to restore amphibian habitat in fire-suppressed pine flatwoods wetlands. Fire Ecology, 9/1: 96-109.

Jones, K., P. Hill, T. Gorman, C. Haas. 2012. Climbing behavior of flatwoods salamander (Ambystoma bishopi/ A. cingulatum). Southeastern Naturalist, 11/3: 537-542.

Lannoo, M. 2005. Amphibian Declines: The Conservation Status of United States Species. Los Angeles, California: University of California Press.

Means, B., J. Palis, M. Baggett. 1996. Effects of slash pine silviculture on a Florida population of flatwoods salamander. Conservation Biology, 10/2: 426-437.

Palis, J., G. Hammerson. 2008. "Ambystoma cingulatum" (On-line). The IUCN Red List of Threatened Species 2008: e.T1099A3236962. Accessed November 19, 2017 at http://dx.doi.org/10.2305/IUCN.UK.2008.RLTS.T1099A3236962.en.

Palis, J., M. Aresco. 2007. Immigration orientation and migration distance of four-pond breeding amphibians in northwestern Florida. Florida Scientist, 70/3: 251-263.

Palis, J., M. Aresco, S. Kilpatrick. 2006. Breeding biology of a Florida population of Ambystoma cingulatum (flatwoods salamander) during a drought. Southeastern Naturalist, 5/1: 1-8.

Pauly, G., S. Bennett, J. Palis, H. Bradley Shaffer. 2011. Conservation and genetics of the frosted flatwoods salamander (Ambystoma cingulatum) on the Atlantic coastal plain. Conservation Genetics, 13/1: 1-7.

Semlitsch, R., S. Walls, W. Barichivich, K. O'Donnell. 2017. Extinction debt as a driver of the amphibian declines: An example with imperiled flatwoods salamander. Journal of Herpetology, 51/1: 12-18.

Westervelt, J., J. Sperry, J. Burton, J. Palis. 2013. Modeling response of frosted flatwoods salamander populations to historic and predicted climate variables. Ecological Modelling, 1/1: 18-24.

Whiles, M., J. Jensen, J. Palis, W. Dyer. 2004. Diets of larval flatwoods salamanders, Ambystoma cinguatum, from Florida and South Carolina. Journal of Herpetology, 38/2: 208-214.

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