The Flatwoods salamander (Ambystoma cingulatum) was first described by Cope in 1868 but has remained rare in museum collections to this day. This species is a small, slender species of Ambystoma with a light gray reticulate or frosted pattern on a dark gray or black dorsum (Petranka 1998), though gulf coast populations often have a brownish gray reticulate pattern on the dorsum. The venter is typically dark gray with a light gray flecking or spotting pattern present. Adults may reach lengths of 13.5cm in total length (TL) (Palis 1996) with an average of 15 costal grooves in a range of 13-16. Larvae develop a yellow - gold vertebral stripe along the length of the body shortly after hatching. Hatchlings are identified by their uniformly dark brown dorsal patterning and their pale brown venter. Hatchlings typically measure 7.5-11.5 mm SVL and 10-19 mm TL (Anderson and Williamson 1976; Palis 1995).

Ambystoma cingulatum has a very narrow distribution occupying seasonally wet, pine flatwoods and savannas from southern South Carolina, southern Georgia, and northern Florida west to southern Alabama (Petranka 1998). Prior to European settlement Ambystoma cingulatum was found primarily in seasonally wet, longleaf pine (Pinus palustris)-wiregrass (Aristida stricta) communities. Populations are discontinuous and rapidly becoming fragmented throughout the range as longleaf pine habitat is destroyed and replaced with slash pine. There may have been extirpations occurring in Alabama, and recent work suggests the extirpation of a sizable breeding population in a period of approximately 22 years (Means et al. 1996).

South Carolina, Southern Georgia, and Northern Florida, west to southern Alabama.

Biogeographic Regions: nearctic (Native )

endemic to a single nation

United States

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

Type of Residency: Year-round

Global Range: (5000-20,000 square km (about 2000-8000 square miles)) Range includes the lower southeastern Coastal Plain of the United States from southern South Carolina southward to Marion County (north-central Florida) and disjunctly westward through southern Georgia (Jensen et al. 2008) and northern Florida to the Apalachicola and Flint rivers (mid-Panhandle of Florida and northward) (Pauly et al. 2007).

Length: 3 1/2- 5 1/16 in.

The Flatwoods salamander is a small highly variable ambystomid, with coloration ranging from specks, to grayish lines that resemble a frosted or lichenlike reticulated pattern. About 5% of A. cingulatum lack the frosted appearance altogether, with 2% having light annuli on their dorsum (Petranka 1998). Costal grooves average at about fifteen. The head is small compared to the shoulder and neck. Sexual dimorphism is slight to non-existent, with sexually active males being slightly shorter in length, and having a swollen cloacal region. Hatchling A. cingulatum are 7.5-11.5 mm long, and are pale brown underneath, and dark brown above (Petranka 1998). The young develop a yellow stripe that runs the length of the body. Older larvae have a paler stripe that may be retained up to one year after metamorphosis. While variations in coloration are often geographic, subspecies are not recognized (Conant & Collins 1998).

Length: 13 cm

Adults may be confused with the slimy salamander (Plethodon grobmani), small-mouthed salamander (Ambystoma texanum), or Mabee's salamander (Ambystoma mabeei). Slimy salamanders are readily distinguished by the presence of a small groove (nasolabial groove) from the nostril to upper lip (absent in all Ambystoma). Ambystoma texanum (smallmouth salamander) overlaps the range of Ambystoma cingulatum in extreme southwestern Alabama. Small-mouthed salamanders have a very short, rounded snout and, in Alabama, are brown or dark gray with lichen-like light blotches (Mount 1975). In South Carolina, Ambystoma cingulatum has been observed breeding in the same wetland as Ambystoma mabeei (Anderson and Williamson 1976). The body of Mabee's salamander is dark brown or black with pale specks that are concentrated along the sides.

Although the flatwoods salamander larval pattern is distinctive, two other Ambystoma larvae may appear similar to the untrained eye. Like Ambystoma cingulatum, Ambystoma mabeei larvae have a light mid-lateral stripe between two dark lateral stripes. However, unlike the continuous lateral stripes of Ambystoma cingulatum, those of Ambystoma mabeei are broken into blotches (Hardy and Olmon 1974). In addition, the stripe extending from the snout to the gills in Ambystoma mabeei is diffuse and indistinct, and the upper lip stripe is replaced by a series of spots (Hardy and Olmon 1974). Larval mole salamanders (Ambystoma talpoideum) may have an indistinct, light mid-lateral stripe, but are readily distinguished from Ambystoma cingulatum larvae by the presence of a dark mid-ventral stripe and dark dorsal crossbands (pers. obs.). The light mid-lateral stripe of larval Ambystoma cingulatum is retained by metamorphs through their first year (pers. obs.). It is best observed by shining a bright light through the body.

• Terrestrial
• Freshwater

Ambystoma cingulatum occupies seasonally wet, pine flatwoods, and pine savannas in the southern United States. The Flatwoods salamander is typically found under logs near small cypress ponds. Traditionally, the Flatwoods salamander was found in sandy, seasonally wet, longleaf pine communities. But, due to European settlement these areas have been replaced by slash pine, or destroyed altogether (Petranka 1998). Adult A. cingulatum are subterranean, living mainly underground in root channels or crayfish burrows (Conant & Collins 1998).

Terrestrial Biomes: forest

Comments: The following information pertains to the Ambystoma cingulatum/bishopi complex as a whole.

Post-larval individuals inhabit mesic longleaf pine (Pinus palustris)-wiregrass (Aristida stricta) flatwoods and savannas. The terrestrial habitat is best described as a topographically flat or slightly rolling wiregrass-dominated grassland having little to no midstory and an open overstory of widely scattered longleaf pine. Low-growing shrubs, such as saw palmetto (Serenoa repens), gallberry (Ilex glabra) and blueberries (Vaccinium spp.), co-exist with grasses and forbs in the groundcover. Groundcover plant diversity is usually very high. The underlying soil is typically poorly drained sand that becomes seasonally inundated.

Slash pine flatwoods is often cited as the preferred terrestrial habitat of the flatwoods salamander (e.g., Conant and Collins 1991). This may be the result of an error made by Martof (1968) in which he referred to longleaf pine as slash pine (Pinus elliottii). In addition, slash pine now dominates or co-occurs with longleaf pine in many pine flatwoods communities as a result of fire suppression and preferential harvest of longleaf pine (Avers and Bracy 1975). Historically, however, fire-tolerant longleaf pine dominated the flatwoods, whereas slash pine was confined principally to wetlands (Harper 1914, Avers and Bracy 1975). Post-larval individuals are fossorial (live underground) and occupy burrows (Goin 1950, Neill 1951, Mount 1975, Ashton 1992). Presumably, they remain underground during the lightning-season (May through September). Adults are rarely encountered under cover objects at or near breeding sites (J. Palis, pers. obs.).

Breeding occurs in acidic (pH 3.6-5.6 (Palis, unpubl. data)), tannin-stained ephemeral wetlands (swamps or graminoid-dominated depressions) that range in size from 0.02 to 9.5 ha, and are usually not more than 0.5 m deep (Palis, unpubl. data). The overstory is typically dominated by pond cypress (Taxodium ascendens), blackgum (Nyssa sylvatica var. biflora) and slash pine, but can also include red maple (Acer rubrum), sweetgum (Liquidambar styraciflua), sweetbay (Magnolia virginiana), and loblolly bay (Gordonia lasianthus). Canopy coverage ranges from near zero to almost 100% (Palis, unpubl. data). The midstory, which is often very dense, is most often composed of young of the aforementioned species, myrtle-leaved holly (Ilex myrtifolia), Chapman's St. John's-wort (Hypericum chapmanii), sandweed (Hypericum fasciculatum), titi (Cyrilla racemiflora), storax (Styrax americana), popash (Fraxinus caroliniana), sweet pepperbush (Clethra alnifolia), fetterbush (Lyonia lucida), vine-wicky (Pieris phillyreifolia), and bamboo-vine (Smilax laurifolia). Depending on closure of the canopy and midstory, the herbaceous groundcover of breeding sites can range from about 5% to nearly 100% (Palis, unpubl. data). The groundcover is dominated by graminaceous species, including beakrushes (Rhynchospora spp.), sedges (Carex spp.), panic grasses (Panicum spp.), bluestems (Andropogon spp.), jointtails (Manisurus spp.), three-awned grass (Aristida affinis), plumegrass (Erianthus giganteus), nutrush (Sclera baldwinii) and yellow-eyed grasses (Xyris spp.). The floor of breeding sites is riddled with the burrows of crayfish (genus Procambarus). Breeding sites are typically encircled by a wiregrass-dominated graminaceous ecotone. Breeding sites can include roadside ditches (Anderson and Williamson 1976; Palis, pers. obs.) and borrow pits (D. Stevenson, pers. comm.). Breeding sites often harbor fishes, the most typical species include pygmy sunfishes (Elassoma spp.), mosquitofish (Gambusia holbrookii), and banded sunfish (Enneacanthus obesus) (Palis, unpubl. data). Favorable breeding habitat lacks large predatory fishes.

Before breeding sites fill with water, eggs are deposited singly or in small groups on the ground beneath leaf litter, under logs and Sphagnum mats, at the base of grasses, shrubs or trees, or at the entrance to crayfish burrows (Anderson and Williamson 1976). In wetlands that fill incrementally, eggs are deposited amid graminaceous vegetation at the edge (J. Palis, pers. obs.). Egg deposition in shallow water also has been reported (Ashton 1992). Larvae hide amid inundated graminaceous vegetation by day, but will enter the water column at night (J. Palis, pers. obs.).

Non-Migrant: No. All populations of this species make significant seasonal migrations.

Locally Migrant: Yes. At least some populations of this species make local extended movements (generally less than 200 km) at particular times of the year (e.g., to breeding or wintering grounds, to hibernation sites).

Locally Migrant: No. No populations of this species make annual migrations of over 200 km.

Salamanders of the Ambystoma cingulatum/bishopi complex migrate up to hundreds of meters between breeding and nonbreeding habitats; Ashton (1992) mentioned movements of over 1,700 meters. Migrations to breeding sites occur at night in conjunction with rains and passing cold fronts from mid-fall through early winter (Means 1972, Anderson and Williamson 1976; Palis, unpubl. data).

Hatchlings begin feeding immediately on invertebrates and zooplankton, which aids them in growing at a rapid pace. As A. cingulatum becomes juveniles and adults, they remain carnivorous, feeding primarily on earthworms and other insects (Petranka 1998).

Comments: Goin (1950) found earthworm remains in the stomachs of adults. Larvae feed primarily on small crustaceans (Whiles et al. 2004).

Note: For many non-migratory species, occurrences are roughly equivalent to populations.

Estimated Number of Occurrences: 21 - 80

Comments: Surveys completed since 1990 indicate that 37 populations are known from across the historical range, with 6 in Georgia, 3 in South Carolina, and the remainder in Florida (USFWS 2005).

2500 - 100,000 individuals

Comments: Secretive habits of adults make population estimates difficult. Total adult population size likely is at least several thousand, but actual number is unknown.

Comments: Seldom seen except during the breeding season. Small numbers of post-larval salamanders continue to be active on the surface during the winter months (Palis, unpubl. data).

The male Flatwoods salamander reaches sexual maturity at one year, but often does not breed until the following season. The females are mature and breed during their second year (Petranka 1998). Breeding populations range from 200-400 adults. Migration to breeding sites such as ephemeral habitats like ditches, burrow pits, marshy ponds, and swamps is triggered by rainy weather from mid October to early February. The adults move during the heavy rain and cease moving within five hours of the end of rainfall (Petranka 1998). The salamanders travel to the breeding sites located at or near pine flatwoods that support long leaf pine, slash pine, and wiregrass. Males and females migrate together and emigrate in December and January, after spending approximately thirty eight days at the breeding site. They return to their home range, females weighing 37% less after ovipositing. A. cingulatum is one of the only Ambystoma species, other than A. opacum, that courts terrestrially (Petranka 1998). Females lay 1-34 eggs in linear or clumped fashion beneath logs, leaf litter, sphagnum mats, bare soil, bases of bushes, and at the entrances of crayfish burrows (Conant & Collins 1998). The female abandons the eggs and leaves them where they will hatch in about two weeks, triggered by heavy rains that raise the water level. Hatching becomes staggered when it requires multiple rains to fill the pond. The larval period lasts from three to five months (Petranka 1998).

The following information pertains to the Ambystoma cingulatum/bishopi complex as a whole.

Movements to breeding ponds occur usually between early October and January during rainy evenings when the barometric pressure is falling (Ashton 1992). In Florida, salamanders that entered and exited the breeding site only once remained in the basin an average of 38 days (range 3-117 days) (Palis 1997). Individual females lay up to 225 eggs (Ashton 1992) singly or in small clusters, with larger individuals producing more eggs than smaller ones (Anderson and Williamson 1976). Eggs are laid terrestrially before depressions fill with water; The eggs develop to hatching size within three weeks, but do not hatch until inundated (Anderson and Williamson 1976). The larval period lasts three to four months (11-18 weeks) (Means 1986, Palis and Jensen 1995). Metamorphs emigrate from their natal ponds during the months of March and April (J. Palis, pers. obs.). In captivity, adult size can be reached within one year (Means 1972). Preliminary field data, however, suggest that full size is not attained until the third or fourth year in the wild (Palis, unpubl. data). Although not much bigger than metamorphs, males attain sexual maturity in their first year (Palis 1997). Females, however, do not sexually mature until at least two years old (Palis and Jensen 1995, Palis 1997). Generation length is presumed to be about 8 years.

The Flatwoods salamander is a small, rare ambystomid with widely distributed populations. Clearing of land, creating ditches, filling wetlands, and conservation of native longleaf pine forests to create managed tree farms, have destroyed and reduced populations. A. cingulatum is a candidate for federal concern because populations seem to be declining throughout the range (Petranka 1998). It is endangered in South Carolina, rare in Alabama, Georgia, and Florida. Management is complicated, involving both aquatic and terrestrial preservation (Conant 1998).

IUCN Red List of Threatened Species: vulnerable

United States

Rounded National Status Rank: N2 - Imperiled

Rounded Global Status Rank: G2 - Imperiled

Reasons: Endemic to a small portion of the southeastern U.S. Coastal Plain; typically collected in low numbers; few recent collections; trend data indicate a loss of a large majority of historical local breeding populations; adult and larval habitats continue to be threatened by conversion to other uses.

Intrinsic Vulnerability: Moderately vulnerable

Environmental Specificity: Narrow. Specialist or community with key requirements common.

This species breeds in ephemeral ditches, including roadside ditches and pasture ponds, in addition to pond cypress and black gum habitat. The flatwoods salamander is a fall breeder. Adults typically migrate to breeding sites during rains from mid-October through early February (Means et al.1996). Courtship has not been observed, but it is believed that individuals court on land. Ambystoma cinglatum is one of only two species (see A. opacum) of mole salamanders that are known to court and deposit eggs in terrestrial habitat. Individuals leave the breeding site near their entry point after a period of approximately 38 days. Females deposit 1-34 clumped eggs at or near the water's edge in depressions. As in A. opacum, hatching is triggered as rains that fill the pools inundate the eggs. Females from populations in southern South Carolina and into Georgia have been shown to deposit eggs singly under logs, leaf litter, and sphagnum mats (Anderson and Williamson 1976).

Global Short Term Trend: Relatively stable to decline of 30%

Comments: Currently, the species presumably is declining in concert with continued loss of remaining intact pine flatwoods community (particularly degradation of groundcover). The rate of decline is unknown.

Global Long Term Trend: Decline of 30-70%

Comments: During extensive surveys of historical (pre-1990) breeding ponds, researchers recorded the species at only a small minority of formerly inhabited sites. The salamander has lost much of its former habitat in Georgia and South Carolina.

Habitat fragmentation and destruction are rapidly affecting the flatwoods salamander. The longleaf pine-wiregrass communities that once dominated the Coastal Plain from North Carolina to east Texas are rapidly being destroyed for slash pine silviculture practices. It is estimated that the longleaf pine forests once comprised 60.6% of the upland landscape prior to early settlement, yet by 1990 it had dropped to 1.4% of the landscape (Ware et al. 1993). Despite earlier controversy about whether the flatwoods salamander preferred the slash pine habitat, Means et al. (1996) demonstrated that adult members of the species are primarily inhabitants of the longleaf pine-wiregrass flatwoods, and that the slash pine flatwoods only form a small portion of the adult habitat.

Degree of Threat: B : Moderately threatened throughout its range, communities provide natural resources that when exploited alter the composition and structure of the community over the long-term, but are apparently recoverable

Comments: Potential threats include conversion of pine flatwoods habitat for agriculture, silviculture, or commercial/residential development; drainage or enlargement (with subsequent introduction of predatory fishes) of breeding ponds; habitat alteration resulting from suppression of fire; mortality and collecting losses associated with crayfish harvest; and highway mortality during migration.

The principal threat is habitat destruction as a result of agriculture, silviculture, and residential and commercial development. Modern silvicultural methods rely on altering soil hydrology, suppressing fire, shortening timber rotations, and replacing widely-spaced longleaf pine with dense plantations of slash pine. Loss of groundcover vegetation due to mechanical soil preparation, fire suppression, and shading by overstories of slash pine have been implicated in the decline in north Florida (Means et al. 1994, 1996).

Larvae are threatened in some wetlands by the harvest of crayfish as bait. Bait harvesters drag large hardware cloth buckets through inundated vegetation, dump the contents of the bucket on the ground, and then sort out the crayfish. Flatwoods salamander larvae taken in this manner are left to die or are collected as bait (J. Palis, pers. obs.).

The effect of herbicide or fertilization application on flatwoods salamanders is unknown. However, fertilization of plantations often results in eutrophication of wetlands, promoting algal blooms. Larval flatwoods salamanders have not been observed in algal-choked wetlands (J. Palis, pers. obs.).

Ditching or berming of small, isolated pond-cypress wetlands, a common practice when establishing slash pine plantations on mesic sites, results in lowered water levels and shortened hydroperiods (Marois and Ewel 1983). These hydrologic perturbations could prevent successful flatwoods salamander reproduction by preventing egg inundation or stranding larvae before they are capable of metamorphosis. Altered hydrology, in association with fire exclusion, results in a shift in dominance from pond-cypress to broad-leaved hardwoods that reduce herbaceous groundcover vegetation through shading (Marois and Ewel 1983). This may be detrimental since A. cingulatum larvae take shelter in herbaceous vegetation during the day.

Ephemeral pond-cypress depressions are sometimes converted into permanent water bodies, rendering them unsuitable for flatwoods salamander reproduction (J. Palis, pers. obs.).

A constant winter-burn fire plan could be detrimental (Ashton 1992).

See USFWS (1999) for additional information.

Restoration Potential: Recovery is directly linked with the ability to preserve existing habitat and restore degraded habitat. Given the drastic decline in the extent of longleaf pine-dominated communities (Ware et al. 1993), elevation of flatwoods salamander populations above present levels is unlikely. Restoration of degraded mesic, seasonally inundated longleaf pine flatwoods and savannas has not been attempted, and may only be feasible in cases where soil disturbance is minimal. The effectiveness of reintroduction into areas where extirpated is unknown.

Preserve Selection and Design Considerations: High quality occurrences include several wetlands within a matrix of pine flatwoods and savanna. Based on the maximum distance adults are known to travel between reproductive and nonreproductive habitat (1.7 km), each breeding site should be surrounded by at least 10 sq km of terrestrial habitat. Longterm perpetuation of a viable population of flatwoods salamanders will presumably require protection of a larger area of terrestrial habitat encompassing a suite of alternative breeding sites (Travis 1994). A suite of wetlands guards against extirpation at any one breeding site, since animals can immigrate from nearby wetlands. The minimum viable population size needed to sustain a population longterm is not known. Preliminary drift fence data at Eglin Air Force Base, Florida, suggests that breeding population sizes are low relative to other Ambystoma (Palis, unpubl. data). However, this may be a site specific observation as larger breeding migrations have been observed elsewhere in the range (R. Moulis, pers. comm.). Presently, there is no method of assessing an occurrence based on the number of animals captured at a drift fence or the number of larvae inhabiting a breeding site.

Management Requirements: Maintenance of intact mesic longleaf pine-wiregrass flatwoods and ephemeral wetlands by mimicking natural forces, such as lightning-season fire, is the most appropriate form of management. On sites where timber extraction is practiced, several precautions should be taken to limit the impact to flatwoods salamanders. Tree harvest should be restricted to dry periods to prevent soil compaction and rutting. Clearcutting should be replaced with selective timber harvest and natural regeneration enhanced by fire, particularly lightning-season fire. If off-site species such as slash pine have been planted, they should be removed and replaced with longleaf pine at densities found in nature. Mechanical preparation of the soil should be avoided. If a site supports mature, closed-canopy pine plantations, they should be thinned with as little disturbance to the soil and remaining groundcover as possible. The natural hydrology and fire regime of terrestrial and aquatic habitats should be restored on sites where altered.

The wetland/upland ecotone appears to be critical to successful flatwoods salamander reproduction. Some areas are in need of periodic burning to clear encroaching shrubby vegetation that shades out herbaceous ground cover (Palis and Jensen 1995). Maintenance of a graminaceous ecotone and breeding site will require burning in the lightning-season when wetlands are dry or nearly dry (Huffman and Blanchard 1990) . Bury et al. (1980) recommended that wiregrass not be burned in winter (destructive to wiregrass [used for egg attachment] and possibly to salamanders directly). Palis and Jensen (1995) stated that winter burns may be needed to avoid catastrophic fires when warm-season burning is initiated.

Mechanical disturbance of the wetland-upland ecotone should be avoided. The practice of "protecting" wetlands by encircling them with plow line should be abandoned. Where present, berms should be removed and drainage ditches filled.

Breeding ponds should not be dredged or stocked with fishes.

Management Research Needs: Development of a quantitative means of using surveys of larvae to indicate the size of the adult breeding population is needed.

Demographic data are needed to better understand the natural history and, in particular, factors that limit population size (e.g., egg, larval, and metamorph survivorship; competition with other species).

Longterm drift-fence studies are needed at several nearby sites to examine inter-pond salamander movement and to delineate the range of natural population fluctuations.

More information is needed on the extent of upland habitat required to support a population breeding in a particular pond. Radiotelemetry or radioactive tagging of adults could be used to address this need.

Effects on salamander populations of different forms of resource management and of anthropogenic habitat disturbance need to be examined (Palis and Jensen 1995).

Global Protection: Several (4-12) occurrences appropriately protected and managed

Comments: This species occurs on several managed areas (see USFWS 1997). A substantial number of populations are on public land (USFWS 1997). Sites include Apalachicola and Osceola national forests and St. Marks National Wildlife Refuge in Florida, Fort Stewart in Georgia, and Francis Marion National Forest in South Carolina. Outlook on private lands is poor.

Needs: Maintain ecological integrity of managed areas from which species is known. Limit collecting if exploitation is extensive. Protect populations on private lands by conservation agreements or land purchases.

Other than preservation of biodiversity and treasuring a native species to maintain ecosystem equilibrium, there is little economic importance to humans. It could however be hypothesized that salamanders keep insect populations under control

The flatwoods salamander is a species with a relatively restricted range and specific habitat requirements. The longleaf pine communities, which were once the dominant upland habitat type in the coastal plain from North Carolina to east Texas, have declined by more than 50% since presettlement times. Silvicultural practices have driven this habitat destruction resulting in the fragmentation of the remaining flatwoods salamander populations and potentially other species dependent on these communities. Thus, the species provides an example where reevaluating the impact economic forestry practices may have on species may be warranted.

Stewardship Overview: Activities aimed at restoring/maintaining the ecological integrity of mesic longleaf pine-wiregrass flatwoods and associated ephemeral wetlands are needed to preserve extant populations of flatwoods salamanders.

Comments: Based on patterns of genetic and morphological variation, Pauly et al. (2007) concluded that Ambystoma cingulatum should be split into two species, A. bishopi west of the Apalachicola-Flint rivers and A. cingulatum east of those rivers.

Based on mtDNA evidence, A. cingulatum (sensu stricto) inhabiting the eastern Gulf Coastal Plain and the Atlantic Coastal Plain in northern Florida are distinct lineages; the Suwannee River and a large distributional gap separate them (Pauly et al. 2007). Further data are needed to determine whether these two lineages warrant recognition as different species (Pauly et al. 2007).