Ambystoma cingulatum Cope, 1867 (1868)
John G. Palis
D. Bruce Means
1. Historical versus Current Distribution. Flatwoods salamanders (Ambystoma cingulatum) are restricted to the Coastal Plain of the southeastern United States; ranging from southern South Carolina south to Marion County, Florida, and west through southern Georgia and the Florida Panhandle to southwestern Alabama (Conant and Collins, 1991). Formerly, Louisiana, Mississippi, and North Carolina were included in their range (Martof, 1968); however, these records have been discounted as misidentifications (Goin, 1950; Hardy and Olmon, 1974; P. Moler, personal communication).
Flatwoods salamanders may no longer occur at many historical locations. The last observation of flatwoods salamanders in Alabama was 1981 (Jones et al., 1982). Palis (1997b) was unable to confirm the continued existence of flatwoods salamanders at/near 2/3 of historical breeding sites sampled in Florida. Recent (post-1989) surveys failed to detect flatwoods salamanders at 33 historical locations in Georgia and at 26 of 29 (90%) historical sites in South Carolina (U.S.F.W.S., 1997c). However, recent surveys documented 102 previously unknown breeding sites in Florida (n = 75) and Georgia (n = 27; U.S.F.W.S., 1997c). Despite intense efforts, no new breeding sites have been located in Alabama or South Carolina (U.S.F.W.S., 1997c).
2. Historical versus Current Abundance. Historical (Goin, 1950) and recent (Palis, 1997a) observations indicate that flatwoods salamanders can be common locally. However, due to habitat alteration, they may be less common now than they were historically. For example, Means et al. (1996) observed a 99% decline in the breeding migration of adults at a Florida site from the 1970s–'90s.
3. Life History Features.
A. Breeding. Reproduction is aquatic.
i. Breeding migrations. Flatwoods salamanders are autumn breeders; most individuals migrate to breeding sites in October–November and emigrate in December–January (Means, 1972; Anderson and Williamson, 1976; Means et al., 1996; Palis, 1997a). Adults typically move during rains associated with passing cold fronts, but will also migrate on rainless nights when soils are near saturation (Palis, 1997a). Males and females migrate simultaneously (Palis, 1997a). Post-breeding salamanders tend to emigrate in the direction of immigration, suggesting the ability to return to a particular terrestrial retreat (Palis, 1997a). In an all-night (29–30 October 1993) investigation of a flatwoods salamander breeding migration at a site in Okaloosa County, Florida (see Palis, 1997a), nearly equal numbers of salamanders were observed between 1800–2400 (n = 27) and 2401–0700 (n = 26; J.G.P., unpublished data). Rain fell continuously and the temperatures remained nearly constant across both observation periods. Adults may move as far as 1,700 m from breeding sites (Ashton, 1992). However, this distance may be exceptional inasmuch as other Ambystoma are reported to travel shorter distances from breeding sites (Semlitsch, 1998; P.K. Williams, 1973).
ii. Breeding habitat. Breeding sites include isolated swamps where pond cypress or blackgum predominate, marshy pasture ponds, roadside ditches, or small, shallow borrow pits (Anderson and Williamson, 1976; Palis, 1995b, 1997b; D. Stevenson, personal communication). Where woody regeneration is poor, breeding sites that were logged may now be dominated by shrubs or graminaceous vegetation.
i. Egg deposition sites. Eggs are deposited singly in small groups (1–34 eggs) in dry pond basins either in the open on bare soil or under cover beneath logs, leaf litter, dead grass, Sphagnum mats or within crayfish burrows (Anderson and Williamson, 1976). Anderson and Williamson's (1976) report that most eggs are deposited in the lowest elevations of depressions may be an artifact of sampling. Eggs are more easily located in the open, lower elevations of breeding basins than in the more densely vegetated grassy edges (G. Williamson, personal communication). Captures of recently hatched larvae suggest that many eggs are deposited in these upper grassy zones (J.G.P., personal observations). Cryptic oviposition sites offer protection from fire (J. Jensen, personal communication). In Florida, flatwoods salamanders oviposit on vegetation in water (R. Ashton, personal communication).
ii. Clutch size. Based on ovarian counts, from 97–222 (mean 163) in South Carolina and Georgia (Anderson and Williamson, 1976) and about 225 in Florida (Ashton, 1992). Egg number correlates with female size (see also Petranka, 1998).
i. Length of larval stage. Palis (1995b) determined the larval period at two Florida breeding sites. At a site containing two distinct size classes of larvae, larvae grew 1.7–1.9 mm/wk, and the larval period encompassed 15–18 wk. At the second site, larvae grew 2.5 mm/wk and transformed in 11–13 wk. Sekerak et al. (1996) observed metamorphosis after an approximate 12-wk larval period.
ii. Larval requirements.
a. Food. Food habits have not been studied, but larvae likely prey on a variety of aquatic invertebrates and perhaps small vertebrates (e.g., other amphibian larvae, smaller conspecifics). Larvae have been raised in the laboratory on earthworms (Sparganophilus spp., Diplocardia eiseni), fairy shrimp (Eubranchiopoda), aquatic amphipods (Hyallela spp.), and mosquito (dipteran) larvae.
b. Cover. Larvae hide in submerged herbaceous vegetation during the day. In a quantitative study of ten larval sites in Florida, Sekerak et al. (1996) found larvae most often in sedge- and pipewort-dominated plots. In a daytime survey of 82 breeding sites in Florida, Palis (1997b) captured larvae principally in graminaceous vegetation. At night, however, larvae enter the water column (Palis, 1995b).
iii. Larval polymorphisms. None known.
iv. Features of metamorphosis. Larvae typically metamorphose within a 5–10 d span in March–April (Mecham and Hellman, 1952; Palis, 1995b). Metamorphic animals range in size from 35.5–46.0 mm SVL (Goin, 1950; Mecham and Hellman, 1952; Palis, 1995b; J.G.P., unpublished data).
v. Post-metamorphic migrations. Post-metamorphic migrations have not been studied. However, at a Florida site, 28% of the yearlings first observed at a drift fence in the fall were captured emigrating from the pond, suggesting that they summered at the breeding site (Palis, 1997a). Metamorphic animals have been found within 15 m of a breeding site in Liberty County, Florida, in April (J.G.P., personal observations) and collected while emigrating from a breeding site in Jasper County, South Carolina, in mid April (data with specimens in collection of Savannah Science Museum).
vi. Neoteny. Not known to occur.
D. Juvenile Habitat. Unknown, but presumably similar to that of adults. Because small salamanders are susceptible to desiccation (Spotila, 1972), juveniles may be restricted to the vicinity of the breeding site in dry years (Palis, 1997a).
E. Adult Habitat. Often described as slash pine flatwoods (Goin, 1950; Martof, 1968; Conant and Collins, 1991), but Palis (1996a) and Means et al. (1996) argued that the primary habitats of post-metamorphic animals are longleaf pine flatwoods and savannas. The distinction is important because longleaf pine-dominated uplands typically have a more open canopy and a greater herbaceous component to the ground cover than slash pine-dominated uplands. Neill (1951b) and Ashton (1992) have observed adults in crawfish burrows.
F. Home Range Size. Observations of home range size are limited to Ashton (1992) who determined that the activity range of one individual encompassed 1,500 m2.
G. Territories. The potential for territorial behavior has not been studied.
H. Aestivation/Avoiding Dessication. Unknown. However, we are not aware of any specimens collected in July–August, suggesting that either flatwoods salamanders are active only underground or aestivate during these months.
I. Seasonal Migrations. Documented migrations involve breeding (adults to/from breeding sites in the fall) and migrations of newly metamorphosed animals in the spring.
J. Torpor (Hibernation). Unknown and unlikely, given that the species is active during the winter (Goin, 1950; Palis, 1997a).
K. Interspecific Associations/Exclusions. Larvae co-occur with a wide variety of invertebrates and vertebrates in breeding sites, including other salamander species (Anderson and Williamson, 1976; Palis, 1997b). Competitive interactions with other salamander species, either in larval or post metamorphic stages, is unknown.
L. Age/Size at Reproductive Maturity. In a western Florida population, males attained sexual maturity in 1 yr, but most females apparently did not mature for at least 2 yr (Palis, 1997a). In this population, the smallest mature male was 44 mm SVL, the smallest gravid female 53 mm. In the laboratory, Means (1972) raised larvae to adult sizes in 1 yr.
M. Longevity. Flatwoods salamanders were not included among the species for which captive longevity records were available to Bowler (1977). One of us (D.B.M.) and Clive Longden (personal communication) kept adults raised from larvae in captivity for ≤ 4 yr. Longevity of individuals in the wild is unknown.
N. Feeding Behavior. Although feeding behavior was not observed, Goin (1950) noted that stomachs of adults contained earthworms (Diplocardia sp.). Prey capture kinematics were investigated by Beneski et al. (1995). In the laboratory, newly metamorphosed animals were raised to adulthood on earthworms (Diplocardia eiseni, Pheretima spp.) and small slugs (D.B.M., personal observations).
O. Predators. Unknown, but some associated species are potential predators (Anderson and Williamson, 1976; Palis, 1997b).
P. Anti-Predator Mechanisms. As with other members of the genus Ambystoma, flatwoods salamanders have concentrations of granular glands along the length of their tail that produce distasteful secretions (Brodie, 1977). When threatened, adults coil with their head positioned under the base of their tail (Brodie, 1977). Occasionally individuals will weakly lash their tail (Brodie, 1977). Juveniles will raise their tail and slowly undulate it from side to side (J.G.P., personal observations).
Q. Diseases. Unknown; however, one of us observed a dying larva at a breeding site in Liberty County, Florida, that appeared to have red-leg disease (J.G.P., personal observations).
R. Parasites. Unknown.
4. Conservation. Flatwoods salamanders may no longer occur at many historical locations and may be extirpated in Alabama (Jones et al., 1982). Flatwoods salamanders are listed as Threatened by the Federal Government (U.S.F.W.S., 1997c), Protected in Georgia (http://georgiawildlife.dnr.state.ga.us), and State Endangered in South Carolina (www.dnr.state.sc.us). The main threat to this species appears to be habitat alteration (Means et al., 1996).
Acknowledgments. We thank Ray Ashton, John Jensen, Clive Longden, Dirk Stevenson, and Gerald Williamson for sharing their observations on flatwoods salamanders with us.
Literature references for Amphibian Declines: The Conservation Status of United States Species, edited by Michael Lannoo, are here.
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