Description
There are two recognized subspecies of Rana capito, the Carolina gopher frog, Rana capito capito and the Florida gopher frog, R. c. aesopus. (Rana sevosa used to be a subspecies of R. capito, R. c. sevosa, but has recently been elevated to species status (Young and Crother, 2001) Rana capito capito ranges in size from 2.5 to 3.5 inches and has a variable color pattern ranging from dark gray to brown spotted with black, reddish brown or dark brown (Conant and Collins, 1991). Warts are always prominent, variable in shape and almost pavement-like and the ventral surface is heavily marked with dark flecks (Conant and Collins, 1991). Rana capito aesopus ranges in size from 2.75 to 3.75 inches and has a color pattern that varies from creamy white to brown through yellowish or purplish (Conant and Collins, 1991). The back may be smooth or slightly warty with black or dark brown irregular markings. The chin and throat is usually spotted but the belly is usually unmarked posteriorly (Conant and Collins, 1991). Males may have yellow on the dorsolateral ridges, on the warts, along upper jaw, and in armpits and groins (Conant and Collins, 1991) and females have longer snout vent lengths than males (Greenberg, 2001). Their voice has been characterized by a deep roaring snore. Large choruses produce an effect like that of pounding surf (Conant and Collins, 1991).

Distribution and Habitat

Country distribution from AmphibiaWeb's database: United States

U.S. state distribution from AmphibiaWeb's database: Alabama, Florida, Georgia, North Carolina, South Carolina, Tennessee

	View distribution map in BerkeleyMapper.
	View Bd and Bsal data (6 records).

The geographic range of R. capito, prior to the elevation of the subspecies R. sevosa, extended from extreme southeastern Louisiana along the Gulf coast to throughout the upper three-fourths of the Florida peninsula and north along the Atlantic coast to southeastern North Carolina (Young and Crother, 2001). Now it is assumed that the Louisiana, Mississippi and western Alabama populations were all R. sevosa. The Carolina subspecies, R. c. capito, occurs throughout the coastal plain of the Carolinas and is scattered throughout the southeast part of Georgia (Conant and Collins, 1991). The Florida subspecies, R. c. aesopus, occurs along the coastal plains of South Georgia and three-fourths of the Florida peninsula, with isolated populations in west-central Georgia (Conant and Collins 1991).

This species can be found in a wide range of habitats from dry, upland xeric oak scrub, and oak hammocks to pine flat wood forests. Preferred breeding habitats include seasonally flooded, grassy ponds and cypress heads that lack fish populations. In Florida, this species is closely associated with gopher tortoise burrows, particularly in xeric longleaf pin-wiregrass uplands that are within about 1.7km of suitable breeding ponds (Schmalzer et al., 1999).

Life History, Abundance, Activity, and Special Behaviors
Throughout its range, R. capito is a commensal with the gopher tortoise (Gopherus polyphemus), hence the common name gopher frog, and is almost entirely dependent on G. polyphemus for shelter and to some extent food (Franz, 1988). However, it occasionally occupies a variety of other retreats including the burrows of rodents and crayfish, as well as stump holes and other crevices (Conant and Collins, 1991). Rana capito is the most frequently observed tortoise burrow commensal (Kent et al., 1997). It is generally nocturnal and emerges to sit near the mouth of its burrow to feed on invertebrates and anurans, including toads (Godley, 1992). This regular foraging activity creates a distinct resting area, also called a ‘platform’, outside each frog’s burrow where the soil has been cleared of vegetation and smoothed by the frog’s constant use (Stevenson and Dyer, 2002).

Greenberg (2001) followed the movement patterns of a population of R. c. aesopus in eight isolated ephemeral ponds in longleaf pine-wiregrass sandhills of the Ocala National Forest, Marion and Putnam Counties, Florida, using intermittent drift fences from February 1994 to January 1998. Four of the ponds were located within fire-suppressed sandhills having high densities of laurel oak, other hardwood species and sand pine and the other four ponds were located within regularly burned, savanna-like longleaf pine-wiregrass sandhills. During Greenberg's study, metamorph capture was significantly higher than adult capture. Greenberg observed more juveniles exiting the ponds than entering the ponds (72.8% as apposed to 27.2%). Recruitment was extremely variable between ponds and between years with no obvious correlation with pond hydroperiod. Adult recruitment into ponds did not correspond with upland habitat type; however, juvenile recruitment was consistently higher for ponds within the savanna-like uplands than for ponds within the fire suppressed hardwood-invaded uplands, although it was not obvious why recruitment at these sites was higher.

Trends and Threats
Florida and Alabama have classified R. capito as threatened. Surveys of herpetofauna throughout its range have listed it as uncommon, rare or endangered for at least two decades. Populations are thought to be declining from wetland habitat loss by drainage, filling, or stocking of fish, and upland habitat loss through development, fragmentation and fire suppression (Bailey, 1991). Fire suppression has been linked to population declines because it reduces habitat quality for G. polyphemus. If fire is suppressed, wiregrass and herbaceous vegetative cover (which is preferred by G. polyphemus) declines from increased shading by invading hardwoods. Fire suppression could also affect habitat quality of breeding ponds. For example, changes in hardwood densities could change water chemistry and pond hydrology by increasing transpiration. Greenberg (2001) found that adult recruitment into ponds did not correspond with upland habitat type (fire suppressed or control burned); however, juvenile recruitment was consistently higher for ponds within the savanna-like uplands than for ponds within the fire suppressed hardwood-invaded uplands. Palis (1998) reported high adult usage and egg mass deposition at a pond within a fire-suppressed, longleaf pine-turkey oak sandhill upland but most adults immigrated into the pond from the direction of an early successional hardwood-invaded sandhill, suggesting heavier use of the more open habitat by adult frogs.

Possible reasons for amphibian decline

General habitat alteration and loss
Habitat modification from deforestation, or logging related activities
Urbanization
Disturbance or death from vehicular traffic
Secondary succession
Habitat fragmentation
Predators (natural or introduced)

Comments
Rana capito was first described as a distinct species by LeConte (1855) but then placed with the crayfish frog Rana areolata by Cope (1875). Collins (1990) resurrected R. capito for all populations within the historical range and since then, researchers have either recognized one gopher frog with three subspecies, R. c. capito, R. c. aesopus and R. c. sevosa or they have included all of R. capito as subspecies of Rana areolata, R. a. capito. Young and Crother (2001) have elevated the Mississippi gopher frog, R. c. sevosa, to full species status.

References

Bailey, M. A. (1991). ''The Dusky Gopher Frog in Alabama.'' Journal of the Alabama Academy of Science, 62(1), 28-34.

Conant, R. and Collins, J.T. (1998). A Field Guide to Reptiles and Amphibians of Eastern and Central North America. 3rd Edition. Houghton Mifflin Company, Boston, Massachusetts.

Franz, R. (1988). ''The Florida Gopher Frog and the Florida Pine Snake as burrow associates of the Gopher Tortoise in northern Florida.'' Proceedings of the Annual Meeting of the Gopher and Tortoise Council. D. R. Jackson and R. J. Bryant, eds., Florida State Museum, Gainesville, 16-20.

Godley, S. J. (1992). ''Threatened: Gopher Frog, Rana capito Le Conte.'' Rare and Endangered Biota of Florida. P.E. Moler, eds., University Press of Florida, Gainesville, 15-19.

Greenberg, C. H. (2001). ''Spatio-temporal dynamics of pond use and recruitment in Florida gopher frogs (Rana capito aesopus).'' Journal of Herpetology, 35(1), 74-85.

Kent, D. M., Langston, M. A., and Hanf, D. W. (1997). ''Observations of vertebrates associated with gopher tortoise burrows in Orange County, Florida.'' Florida Scientist, 60(3), 197-201.

Palis, J. G. (1998). ''Breeding biology of the Gopher Frog, Rana capito, in western Florida.'' Journal of Herpetology, 32(2), 217-223.

Schmalzer, P. A., Boyle, S. R., and Swain, H. M. (1999). ''Scrub ecosystems of Brevard County, Florida: a regional characterization.'' Florida Scientist, 62(1), 13-47.

Semlitsch, R. D., Gibbons, J. W., and Tuberville,T. D. (1995). ''Timing of reproduction and metamorphosis in the Carolina Gopher Frog (Rana capito capito) in South Carolina.'' Journal of Herpetology, 29(4), 612-614.

Stevenson, D. J., and Dyer, K. J. (2002). ''Rana capito capito (Carolina Gopher Frog). Refugia.'' Herpetological Review, 33(2), 128-129.

Volpe, E. P. (1957). ''The early development of Rana capito sevosa.'' Tulane Studies in Zoology, 5, 207-225.

Young, J. E. and Crother, B. I. (2001). ''Allozyme evidence for the separation of Rana areolata and Rana capito and for the resurrection of Rana sevosa.'' Copeia, 2001(2), 382-388.

Species Account Citation: AmphibiaWeb 2007 Rana capito: Carolina Gopher Frog <https://amphibiaweb.org/species/6095> University of California, Berkeley, CA, USA. Accessed Dec 31, 2024.

Citation: AmphibiaWeb. 2024. <https://amphibiaweb.org> University of California, Berkeley, CA, USA. Accessed 31 Dec 2024.

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