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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).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.

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).

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.

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). From a few studies scattered throughout its range, it appears that the timing of breeding and larval development varies from shorter breeding and larval periods farther north to multiple breeding episodes and longer larval periods farther south (Bailey, 1991; Semlitsch et al., 1995; Palis, 1998; Greenberg, 2001). In the Atlantic coastal plains of the Savannah River Site in South Carolina at the very edge of their range, Semlitsch et al. (1995), found that the breeding season in some years lasted only a few days and varied in timing between January and April. Palis (1998), who studied their breeding biology in western Florida, reported an eight-month breeding season (October through May) that encompassed three major breeding events (one each in October, February, and April). Reports of larval developmental periods range from 87-113 days in South Carolina (Semlitsch et al., 1995), 141-155 days in laboratory-reared tadpoles (Volpe, 1958) and 210 days in the Florida panhandle (Palis, 1998). Most studies report May through July as the peak emergence time for metamorphic juveniles. Semlitsch et al. (1995) rarely found metamorphs in pitfall traps and the few metamorphs that were seen were caught in drift fences between 27 May and 12 July. Greenberg (2001) found that, in Marion and Putnam Counties, Florida, most emigration of metamorphic juveniles occurred within a 14-86 day period between May and October. Rainfall did not appear to trigger emigration and had a negligible influence on daily emigration rates. 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.

Gopher frogs can live for up to 6 years in the wild and 7 years in captivity.

Range lifespan
Status: wild: 0 to 6 years.

Range lifespan
Status: captivity: 0 to 7 years.

Caddisfly larvae are significant predator on gopher frog egg masses, however, dragonfly nymphs, diving beetles and turtles also prey on egg masses. Occasionally, snakes have been documented at breeding sites. Gopher frogs breed in seasonally flooded ponds absent of predatory fish. Transitioning from aquatic habitat as tadpoles to the terrestrial habitat as juveniles results in high mortality. Juveniles are unfamiliar with their new habitat and the location of burrows, making them easy targets for predation. Approximately 5% of fertilized eggs develop into juveniles. Gopher frogs use burrows to escape potential predators and their camouflaged coloration likely helps reduce predation risk.

Known Predators:

• caddisfly larvae (Trichoptera)
• dragonfly nymphs (Odonata)
• diving beetles (Dytiscidae)
• turtles (Testudines)
• snakes (Serpentes)

Anti-predator Adaptations: cryptic

Gopher frogs have robust, stocky bodies with relatively short forelimbs. They have tapered snouts and a single lateral ridge down each side of the back. Their light-colored body is marked with dark brown or black blotches of various sizes and shapes. Adult snout-vent length ranges from 6 to 9 cm and weight ranges from 47 to 151 g. Adult males are smaller than adult females. Skin texture can ranges from rough to smooth and the color ranges from yellow-white to brown or gray. The venter is often white, cream, or yellow and usually mottled with dark spots. Tadpoles range from yellow-green to olive-green or gray with large black spots on the upper body, tail and fin. Tadpoles typically reach a length of 84 mm; however, in North Carolina tadpoles can exceed 90 mm in length.

Range mass: 47 to 151 g.

Range length: 6 to 9 cm.

Other Physical Features: ectothermic ; heterothermic ; bilateral symmetry

Sexual Dimorphism: female larger

The primary habitat of gopher frogs is native xeric upland habitats, comprised mainly of longleaf pine and sandy substrates. The habitat also contains xeric to mesic hardwoods such as sand pine scrub, longleaf pine flatwoods, and xeric hammocks. These habitats in early successional stages are ideal. Gopher frogs seek shelter in underground refuges, such as the burrows of gopher tortoises, after which they were names, and several species of small mammals, such as rodents, as well as under logs and in stump holes. Gopher frogs also use clumps of grass and leaf litter as refuge during its migration. Refuges protect against adverse weather and predation. Newly metamorphosed gopher frogs are at a high risk of predation and desiccation due to their unfamiliarity with refuge habitat. Fire-maintained habitats with open canopy contain a higher density of gopher tortoise and small mammal burrows. Thus, juvenile gopher frogs avoid closed-canopy habitat and select open-canopy habitat that has been maintained by fire. As a result of fire suppression, habitat loss and degradation, many wildlife species including the gopher frog associated with longleaf pine forests have declined. Gopher frogs breed in temporary or semipermanent (seasonally flooded) ponds but spend the majority of their lives in the burrows of surrounding terrestrial habitat.

Habitat Regions: temperate ; terrestrial

Terrestrial Biomes: forest ; scrub forest

Aquatic Biomes: temporary pools

Other Habitat Features: suburban ; agricultural

Lithobates capito, also known as Lithobates capito, is found mainly in the Coastal Plain of the southeastern United States. Its range extends from central North Carolina to the east and west coasts of southern Florida. There are isolated populations in central and southeastern Alabama, central Tennessee and southwestern Georgia.

Biogeographic Regions: nearctic (Native )

Gopher frogs are carnivorous and are known to consume a variety of invertebrates, including earthworms, cockroaches, spiders, grasshoppers, beetles as well as other toads and frogs. They travel significant distance at night to forage. Tadpoles eat microscopic algae, organic debris, bacteria and protozoans found on underwater vegetation or along the pond bottom. Water quality and lack of canopy cover have a significant impact on prey abundance.

Animal Foods: amphibians; insects; terrestrial non-insect arthropods; other marine invertebrates

Plant Foods: algae; phytoplankton

Primary Diet: carnivore (Insectivore ); herbivore (Algivore)

Both as larvae and adults, gopher frogs are preyed upon by a number of vertebrate and invertebrate predators. In general, amphibians are often used as biological indicators of habitat quality.

There no known adverse effects of gopher frogs on humans.

Little is known of development of gopher frogs. Egg masses are laid in semiperminant ponds just below the surface of the water. Tadpoles metamorphose after 87 to 225 days and disperse into drier upland habitat.

Development - Life Cycle: neotenic/paedomorphic; metamorphosis

Because of declining populations, Lithobates capito is listed as “near threatened” on the IUCN's Red List of Threatened Species. Mississippi gopher frogs, a subspecies of Lithobates capito, is listed as endangered on the U.S. Federal List. Major threats include fire suppression, habitat loss and fragmentation from roads and construction, agriculture and off-road vehicles. Gopher tortoise populations are also declining, leaving fewer burrows for L. capito to use. In general, the area occupied by this species is rather small and is very specific, leaving L. capito highly vulnerable to habitat change. Although precise numbers are difficult to estimate, recent population estimates suggest that less 10,000 individuals remain in the wild, which is significantly reduced from historical numbers.

US Federal List: endangered

CITES: no special status

State of Michigan List: no special status

IUCN Red List of Threatened Species: near threatened

Gopher frogs, and amphibians in general, are often used as biological indicators of habitat quality. Biological indicators are species that are sensitive to environmental change such as pollution or climate change. Because of their permeable skin they are susceptible to environmental stress in both aquatic and terrestrial environments and serve as an early warning to conservationists of poor aquatic habitat conditions.

The call of gopher frogs is often described as a deep, throaty “snore”, which can last up to two seconds and can be heard nearly 0.4 km away. They have also been documented calling while submerged. Their calls can be heard throughout the year, especially after heavy rains, but are much more abundant during breeding season.

Communication Channels: acoustic

Perception Channels: visual ; tactile ; acoustic ; chemical

In recent years, there has been a taxonomic reassignment of genus Rana to Lithobates. There is still ongoing research to determine if the reassignment is correct. As a result, Lithobates capito is also known as Lithobates capito.

Gopher frogs are polygynous and breed from January to April, immediately after a heavy rain. Males actively call to potential mates during this time. Males occupy breeding ponds for about a month while females stay less than a week.

Mating System: polygynous

Breeding generally occurs during winter and early spring from January through April. Reproduction can occur as early as September and October following heavy rains. Breeding sites are seasonally flooded isolated ponds free of predatory fish. Males usually occupy breeding ponds for about a month while the females stay less than a week. Females lay cluster containing thousands of eggs. Eggs are gray to gray-black and range in size from 1.67 to 2.7 mm in diameter. Evidence suggests that each female lays one egg mass per breeding season. She deposits the egg mass near the surface of the water on a rigid support (e.g., semi-submerged shrub) in order to keep the eggs at a desired depth. As surface waters warm, development of embryos progresses. Tadpoles metamorphose after a larval period ranging from 87 to 225 days, and disperse into the drier uplands.

Breeding interval: Gopher frogs breed once yearly.

Breeding season: Gopher frogs breed from January to April.

Range time to hatching: 4 to 5 days.

Range time to independence: 87 to 225 days.

Average age at sexual or reproductive maturity (female): 2 years.

Average age at sexual or reproductive maturity (male): 1.5 years.

Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; fertilization (External ); oviparous

Similar to most amphibians, once eggs are laid and fertilized, adult gopher frogs abandon the eggs. Young are completely independent upon hatching.

Parental Investment: no parental involvement

The gopher frog, Lithobates capito, is a ranid frog native to the southeastern United States coastal plain from southern North Carolina through central Alabama. It inhabits a variety of dry sandy upland environments including turkey-oak sandhills, sand pine scrub, longleaf pine flat woods and oak hammocks.

A large, plump frog, Lithobates capito is 2-4.5 inches (5.1 - 11.4 cm) long.Its back is covered with prominent bumps and varies in color from brown to dark gray, with black, brown or red markings.Male Florida gopher frogs sometimes have various yellow markings.

Two gopher frog subspecies are currently recognized. The Carolina gopher frog (L. c. capito), lives on the Atlantic coast of the Carolinas and is slightly smaller than the Florida gopher frog (subspecies L. c. aesopus), which occurs in Florida and southern Georgia. The critically endangered dusky gopher frog Rana sevosa, found in Mississippi, was considered a third subspecies (L. c. sevosa) until 2001, when genetic evidence elevated it to species level (Young and Crother 2001).

The gopher frog is a secretive animal, and difficult to find. Nocturnal, it hides much of the day near by but away from water sources.Especially in Florida, it lives mostly in the burrows of gopher tortoises (Gopherus polyphemus).Indeed, its common name derives from this commensal relationship with gopher tortoises. Gopher frogs will also sometimes seek shelter in rodent warrens, crayfish burrows, in crevices and under logs.At night, gopher frogs sit at the edge of their gopher tortoise burrow to forage on insects, other invertebrates and even small frogs and toads that come within range.This edge of their burrow becomes worn into a characteristic smoothed “pad” where they consistently hunt at the same spot each night, wearing away any vegetation.

During the winter rainy season, rains trigger congregations of adults around temporary water holes and seasonally flooded grassy ponds to breed.After migrating to breeding sites, males set up territories and start calling, a sound described as a “deep roaring snore.”After mating, females deposit large egg masses on submerged vegetation.Especially in northern locations (L. c. capito), breeding seasons are explosive, lasting just a few days.Larvae hatch and develop into adults quickly, within 3.5 months.Further south along the Florida panhandle (L. c. aesopus), breeding season can last for several months and larvae develop over the course of up to seven months.

Listed as “Near Threatened” by the IUCN, this frog is vulnerable to a diversity of environmental effects due to human activity.The main issue is habitat degradation.It does not do well in degraded habitats, and is threatened by timber industry management of forests promoting monocultures and control of seasonal wildfires. Research shows larger numbers in breeding pools where fires were not suppressed and savannah-like conditions prevailed, than in uplands where fire suppression allowed hardwood trees to invade.Introduced predatory fish in their breeding areas has also taken a toll on gopher frog eggs and tadpoles.Construction, increased sedimentation, agricultural grazing, and off road vehicles have caused destruction of wetlands and necessary wetland vegetation required for gopher frog breeding. Furthermore, because gopher turtle populations are also in decline, gopher frogs face a crisis in finding shelter.

Eglin Air Force Base (Florida), Conecuh National Forest (Alabama), and Fort Benning (Georgia) provide protected habitat for the gopher frog.Gopher frogs are listed as a species of special concern in Florida, North Carolina, and South Carolina, and are protected in Alabama. The United States Fish and Wildlife Service is currently evaluating recommendations for possible federal listing of Lithobates capito as a threatened species.

Note: some consider Lithobates a subgenus name, and use the generic name Rana for this species.

(Doubledee 2003; Richter and Jensen 2005; Franz 1988; Georgia Wildlife Web 2000; Hammerson and Jensen 2004; Palis 1998; Wikipedia 2016; Young and Crother 2001)

Rana capito (Rana capito en clasificación scientifica (LeConte, 1855), tamién claseficato como Lithobates capito) ye una especie d'amfibios d'a orden Anura, d'a familia Ranidae y d'o chenero Rana, que se distribuye por o sudeste d'Estatos Unitos (Carolina d'o Norte, Carolina d'o Sud, Cheorchia, Florida, Alabama y Tennessee).

The gopher frog (Lithobates capito) is a species of frog in the family Ranidae, endemic to the south-eastern United States. It primarily inhabits the threatened sandhill communities, flatwoods, and scrub in the Atlantic coastal plain, where it is usually found near ponds.

Lithobates capito​​​ es una especie de anfibio anuro de la familia Ranidae.

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Lithobates capito Lithobates generoko animalia da. Anfibioen barruko Ranidae familian sailkatuta dago, Anura ordenan.

Lithobates capito est une espèce d'amphibiens de la famille des Ranidae.

Lithobates capito é uma espécie de anfíbio da família Ranidae.

É endémica dos Estados Unidos da América.

Os seus habitats naturais são: florestas temperadas, pântanos, lagos de água doce intermitentes e marismas intermitentes de água doce.

Está ameaçada por perda de habitat.