Amphibians of North Carolina
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NC Records

Lithobates capito - Gopher Frog

Class: Amphibia Order: Anura Family: Ranidae Synonym: Rana capito
Taxonomic Comments: Frost et al. (2006) placed species in North American that were formerly in the very large genus Rana into a separate genus, Lithobates, to distinguish them from a large and predominantly Eurasian genus Rana (sensu stricto). There have been numerous arguments put forth for well over a decade about whether these species should be placed back into Rana or retained as Lithobates, with some supporting placing Lithobates as a subgroup within Rana and others supporting the recognition of both genera. There has been no clear resolution of the issue, and both Lithobates and Rana continue to be widely used in recent published literature on North American species. Here, we follow the recommendations of the Society for the Study of Amphibians and Reptiles' Standard English Names Committee and use Lithobates for North American representatives of this group.

Three subspecies of Lithobates capito were previously recognized based on external morphology that included the Carolina Gopher Frog (L. c. capito ), the Florida Gopher Frog (L. c. aesopus ), and the Dusky Gopher Frog (L. c. sevosus ). Authorities today recognize L. sevosus as a separate species based on genetic analyses and treat L. capito as a monotypic taxon with no recognized subspecies (Young and Crother 2001). More recent studies by Richter et al. (2014) that examined mtDNA variation supported previous interpretations and showed L. capito to consists of three highly supported allopatric clades. One occurs in the Coastal Plain of Mississippi, Alabama, Georgia, South Carolina, North Carolina, and the panhandle of Florida. A second clade is restricted to northeastern Florida, while the third clade occurs in southern peninsular Florida. Divergence time between the Coastal Plain and the two Florida peninsular lineages was estimated to be 1.9–2.3 mya, and between the northern and southern peninsular clades to be 1.1–1.3 mya. Enge et al. (2017) examined microsatellite variation in Florida populations and found two genetic clusters that correspond to the Panhandle and combined peninsular clades of Richter et al. (2014). However, they were not able to distinguish between a northern and southern peninsular group.
Species Comments:
Description: The Gopher Frog is a medium-sized frog that has a large head and short hind limbs relative to other ranids in the eastern US. The ground color of the dorsum of the back, sides, and legs varies from off-white to gray or light brown. The ground is overlain with numerous dark brown to blackish spots or blotches of varying shapes and sizes. Barring is usually evident on the hind limbs, and the skin is slightly rugose with small, scattered tubercles. A prominent dorsolateral fold extends on each side from behind the eye posteriorly to near the hind limb insertion. The folds can vary from being gray to orangish yellow or brown (Dodd 2013, Beane et al. 2010, Jensen et al. 2008). The venter varies from off-whitish to somewhat yellowish and is mottled or spotted with darker pigment. Many specimens have yellowish coloration in the groin region. Young metamorphs resemble the adults but lack mottling on the belly.

Breeding males have enlarged thumbs and a pair of very large lateral vocal sacs (Wright 1932). Males in local populations typically average slightly smaller than the females. Males that were trapped by Palis (1998) in Florida averaged 80 mm SUL (range = 61–93 mm SUL) versus 94 mm SUL (range = 78–112 mm SUL) for females. Alabama females that Bailey (1990) collected varied from 70–94 mm SUL compared with 61–87 mm SUL for males. Adult males in museum specimens that Goldberg (2021) examined from several sites in the Carolinas averaged 82 mm (range = 73–103 mm SUL) versus 85 mm (range = 67–98 mm) for females.

The larvae vary from yellowish to olive green and have scattered large dark blotches or spots on the body, tail musculature, and tail fin (Dodd 2013). Some individuals lack fine spotting and the tail fins are sometimes clear (Gregoire 2005). The throat is unpigmented and the gut may or may not be visible through the cream to yellowish colored venter. The maximum size of mature larvae is 90 mm TL. Dodd (2013) noted that the larvae are difficult to distinguish from those of L. sphenocephalus that frequently share breeding ponds.
Vocalizations: The advertisement call of L. capito has been described as a deep snore or prolonged groan that last about two seconds. Individuals may pause for a second or two before issuing another call.
Technical Reference: Dodd (2013)
Online Photos:    Google   iNaturalist
Observation Methods: Individuals are best found by searching for calling males.
AmphibiaWeb Account
Distribution in North Carolina
Distribution Comments: Lithobates capito has been documented in Coastal Plain habitats from central North Carolina southward throughout South Carolina, Georgia, southern Alabama, and most of Florida southward to Broward County (Dodd 2013). Disjunct populations were previously documented in central Alabama (Shelby Co.) and Tennessee (Coffee Co.). The breeding site in Alabama was destroyed by a subdivision project and the Tennessee population may have been extirpated (Niemiller and Reynolds 2011). This species had undergone significant declines in North Carolina and other areas of its range, although populations are still widespread and relatively stable in Florida (Krysko et al. 2019). Twenty-three populations were known from North Carolina historically from the coastal pine forests and the Sandhills, but only seven are known today. The historical range extended northward to Beaufort County, but the current northernmost locality records are from Carteret County.
Distribution Reference: Beane et al. (2013), Dodd (2013)
County Map: Clicking on a county returns the records for the species in that county.
GBIF Global Distribution
Key Habitat Requirements
Habitat: The Gopher Frog thrives in fire-maintained sandhills or flatwoods that are dominated by Longleaf Pine or Turkey Oak-pine associations. In Florida they can also be found in other upland habitats such as scrub, xeric hammocks, mesic and scrubby flatwoods, dry prairies, mixed hardwood-pine communities, and a variety of disturbed habitats (Krysko et al. 2019). This species is fossorial and relies heavily on animal burrows for shelter, particularly those of the Gopher Tortoise (Gopheris polyphemus) and small mammals. Where Gopher Tortoise burrows or other burrows are not present, they seek cover in hollow logs, stump holes, root mounds, dead vegetation and clumps of grass.

The adults breed in seasonal or semipermanent ponds that usually lack predatory fish. Ponds that are in the open are preferred, and heavily shaded ponds with closed-canopies are rarely used. The breeding ponds usually have substantial growths of grasses or sedges in and around the margins. Habitats that were listed by Dodd (2013) include depression wetlands, sinkhole ponds, cypress ponds, cypress/gum ponds, pine savanna wetlands and Carolina bays. Artificial habitats such as ditches and borrow pits are also occasionally used. In North Carolina populations are found in fire-maintained sandhills or flatwoods that are dominated by Longleaf Pine or Turkey Oak-pine associations. The historical breeding sites include open, depressional wetlands, Carolina bays, and in rare instances borrow pits.
Biotic Relationships: Both the eggs and larvae are vulnerable to attacks from numerous aquatic predators. Bailey (1990) noted that the eggs are eaten by caddisfly larvae and newts (Notophthalmus viridescens). The larvae are usually found in seasonal or semipermanent ponds that lack fish, although fish can sometimes invade sites if connected to permanent sites during flood events. The larvae appear to lack chemical defenses against fish and invertebrate predators, and are highly vulnerable to predation from both. They also do not respond to chemical cues from predators (Dodd 2013, Phillips 1995).

Gregoire and Gunzburger (2008) found that several species of fish would readily consume the tadpoles, with the Warmouth Sunfish (Lepomis gulosus) being the most effective predator. The Eastern Mosquitofish (Gambusia affinis) was an ineffective predator due to gape limitations, but caused more injuries to tadpoles than any other test species. Injuries of this sort often lead to infections and death. The tadpoles increased their use of cover relative to controls when attacked by small fish such as the Eastern Mosquitofish.

Travis et al. (1985a) manipulated the sizes and densities of tadpoles and found that predation rates from overwintering dragonfly larvae decreased as the tadpole's body size increased. In addition, the number of tadpoles consumed by a dragonfly larva increased with increasing tadpole density, even though the proportion of the total number that were eaten decreased. Cronin and Travis (1986) documented similar outcomes when using two species of backswimmers as predators, with predation rates being lower for the larger size classes of tadpoles.

The juveniles and adults are preyed upon by numerous vertebrates, including snakes (Coluber constrictor, Nerodia fasciata, Thamnophis sirtalis), turtles (Apalone ferox, Kinosternon subrubrum), owls and raccoons (Dodd 2013, Jensen et al. 2008, Roznik and Johnson 2009b). Dodd (2013) noted that L. capito juveniles and adults have a peculiar odor when handled that may function in predator deterrence, and that individuals in burrows are wary and will rapidly retreat down into burrows when approached. Individuals are capable of changing their dorsal coloration to better match their substrates (Wright 1932).
See also Habitat Account for Longleaf Pine Woodlands with Isolated Pools
Life History and Autecology
Breeding and Courtship: Calling and breeding can occur during most times of the year but is most concentrated from October through April depending on the location. The adults migrate from the surrounding uplands when conditions are damp, and typically on rainy nights (Bailey 1990, Palis 1998). Breeding effort is positively correlated with the amount of precipitation just prior to and during the main breeding season, and individuals may skip breeding in drought years (Dodd 2013, Jensen et al. 2003, Palis 1998).

Some reported breeding dates include February and March in Alabama (Mount 1975), January-April in South Carolina (Semlitsch et al. 1995), October through May in the Florida Panhandle (Palis 1998), and mid-February to mid-April in North Carolina, with a peak in March (Beane et al. 2010, Dorcas et al. 2007). Populations in the central and southern Florida can breed year-round (Carr 1940, Krysko et al. 2019, Meshaka and Layne 2015), while in Georgia most occurs from January through the spring months, with occasional breeding during the summer. Heavy rains from tropical storms can trigger breeding outside of the peak breeding season. Semlitsch et al. (1995) noted that breeding populations on the Savannah River Site in South Carolina are very small, with no more than 10 adults breeding at a given site during a single year. Populations at this site regularly skipped breeding for one or more years, and the breeding season lasted less than a week or two during most years at most sites.

Palis (1998) found that the adults bred over an extended period at his study site in the Florida Panhandle, with three major pulses of breeding in October, February, and April. Breeding was first observed in early October following a tropical storm and continued through May, with a peak in February. Both sexes migrated on a given night, but males arrived in greater numbers early in the breeding season. Operational sex ratios were highly biased towards males from October through January, but less so for the remainder of the breeding season. Males spent an average of 18 days at the site as opposed to 9.5 days for females. Some males breed twice during the study and the collective time spent at the site for these averaged 27 days. Bailey (1990, 1991) found that males at a breeding site in Alabama remained for an average of 25 days versus 9 days for females. Fewer males (N = 93) were collected in traps at this site relative to females (N = 176).

The males normally call at night while floating in open water or positioned in or around the shallow margins of ponds, but occasionally call during the day (Meshaka and Layne 2015). Individuals have been observed calling from thick grass tussocks and sedges, piles of debris, and the bases of stumps and trees both in and out of the water (Dodd 2013, Wright 1932). Calling mostly occurs at night, but occasionally in late afternoon, with peaks in chorusing occurring after dark and before dawn. Jensen et al. (1995) reported numerous instances of observers witnessing calling from underwater at several sites. Two males were observed calling while sitting on the pond bottom and produced muffled calls that could be heard from as far as 10 m away. Calling while underwater appears to be commonplace and may hamper the ability to detect local populations.

Information on male-male and male-female social interactions are poorly documented. It is uncertain is males establish and defend calling territories or if females discriminate between calling males when selecting a mate. Females presumably begin laying eggs within several hours after being amplexed, but details are lacking.
Reproductive Mode: Each female produces a ovoid to spherical mass that is about 8–12 cm in diameter and 2.5–7 cm in depth when fully swollen (Bailey 1990, Dodd 2013, Phillips 1995). The egg masses are attached to support structures, including twigs, brush, and the stems of aquatic plants such as grasses, sedges and rushes. The masses are typically laid within 10-20 cm or less of the water surface (Bailey 1990, Dodd 2013, Palis 1998, Wright 1932). The freshly laid eggs are gray to blackish above and cream-colored to whitish below and are surrounded by two gelatinous envelopes. The eggs are large for a ranid of this size and vary from 1.8–2.4 mm in diameter. The outer jelly envelop varies from around 4.5-7 mm in diameter (Wright 1932).

A very large egg mass may contain as many as 5,000 eggs (Wright 1932), but most masses have far fewer eggs. Palis (1998) estimated an average of 2,210 eggs per mass (range = 540–4,825) for 67 masses from Florida and Phillips (1995) reported clutches of 988, 1,281 and 1,463 eggs in Georgia. Bailey (1990) counted a single clutch of 1,709 eggs from Alabama. Hatching occurs in four to seven days and the hatchlings are 12–13 mm TL (Dodd 2013).
Aquatic Life History: The larval ecology of this species is poorly documented. The hatchlings may remain in the vicinity of the jelly masses for a few days and continue development before dispersing widely in the breeding sites. There are no reports of social aggregates or social feeding for this species, but few observations have been made of microhabitat use and feeding behavior. Greenberg (2001a) reported one instance of a negative correlation between the number of juveniles exiting her study ponds and average body size, which suggests that intraspecific competition for food resources may occasionally occur when larvae are crowded. Juvenile recruitment was highly variable among years and ponds, with no more than 195 metamorphs exiting a pond in any one year during the study.

The larval period is thought to last for 3-6 months (Dodd 2013), but detailed studies of local populations are largely lacking. Jensen et al. (2008) reported a larval period of 145–169 days for Georgia populations, while Semlitsch et al. (1995) estimated a larval period of 87–113 days for a South Carolina population across years. Palis (1998) collected metamorphs in May that were likely from eggs that hatched in early October, which implies a larval period of around 7 months for fall-breeding cohorts. The larval stage in North Carolina lasts 3-4 months, with transformation usually occurring from May to July when tadpoles grow larger than 85 mm in total length (NC Wildlife Resources Commission 2018). Newly transformed individuals vary from 26–43 mm SUL (Dodd 2013, Jensen et al. 2008, Palis 1998, Phillips 1995, Semlitsch et al. 1995, Wright 1932).
Terrestrial Life History: The juveniles and adults that leave the ponds often move considerable distances into upland habitats. Humphries and Sisson (2012) tracked adults in the Sandhills of North Carolina as they dispersed from a breeding pond. Nine were successfully tracked to their final summer homes that were 0.5-3.5 km (mean = 1.3 km) from the breeding pond. Frogs that either were killed or had equipment failure had moved an average of 1.0 km before tracking was terminated. Major migrations occurred only on rainy nights, with single night movements varying from 263- 1,200 m (mean = 743 m). Five of the nine individuals made the entire journey to their summer homes in a single night, while others moved over many days (mean = 12 days). They either dug shallow burrows or rested on the surface beneath grass clumps or in leaf litter while moving. The summer refugia that were used were mostly holes associated with tree stumps. A single frog that travelled 3.5 km one year returned to the same stump hole the following year after breeding - a remarkable instance of homing! This result is consistent with other studies showing that the adults tend to emigrate and immigrate to and from ponds in the same direction (Palis 1998), and can return to specific refuges they have used in the past (Blihovde 2006, Roznik 2007).

Roznik et al. (2009) tracked animals in Florida and found that juveniles moved an average of 215 m from ponds (range = 31-665 m) versus 279 m (63-730 m) for adults. When dispersing, the adults used refuges more often than juveniles, and began using them more quickly after leaving a pond. This suggests that the adults may be familiar with the locations of specific refuges surrounding breeding ponds, while recently metamorphosed frogs must learn the spatial locations of refuges by exploring their habitat. In a survey of Gopher tortoise burrows in Florida, Smith et al. (2021) found that the distance from frog-occupied tortoise burrows to the nearest breeding wetland at four sites ranged from 141 to 3,402 m, with the mean distance varying from 1,000 m to 2,200 m among sites.

The movement of the young metamorphs to upland sites presents several challenges, including the risk of predation and desiccation. Dispersal away from the ponds occurs during the summer months and often in the absence of rain (Greenberg 2001a). Roznik and Johnson (2009b) tracked young metamorphs that were moving to upland habitats and documented very high mortality rates, with only 12.5% surviving their first month on land. Most were killed by predators, particularly the Eastern Racer (Coluber constrictor) and Common Gartersnake (Thamnophis sirtalis). Frogs that used Gopher Tortoise burrows, mammal burrows, root holes and other underground refuges while migrating had markedly higher survival than those that did not. Individuals in this study were tracked an average of 157 m from the ponds (maximum = 671 m). Roznik and Johnson (2009a) found that juveniles emigrated in nonrandom directions from ponds and selected fire-maintained habitats that had open canopies, few hardwood trees, small amounts of leaf litter, and large amounts of wiregrass. These habitats also contained higher densities of Gopher Tortoise burrows.

After the juveniles and adults reach their summer homes they are largely fossorial and spend much of the daytime in underground retreats that offer shelter from cold, drought, and periodic fires (Dodd 2013). Populations in Florida and surrounding areas rely heavily on Gopher Tortoise burrows (Franz 1986, Meshaka and Layne 2015, Smith et al. 2021), but will also use the burrows of crayfishes and small mammals --particularly the Southeastern Pocket Gopher (Geomys pinetis), the Oldfield Mouse (Peromyscus polionotus), and the Florida Mouse (Podomys floridanus). Populations in North Carolina and elsewhere outside of the range of the Gopher Tortoise use small mammal burrows and other protective retreats such as hollow logs, stump holes, and root mounds where conditions are relatively cool and moist.

Blihovde (2006) studied burrow use over a 14-month period in central Florida and radio-tracked nine adults over a nine-month period. The adults relied on Gopher Tortoise and Southern Pocket Gopher burrows for retreats. Individuals showed strong site fidelity to the burrows and in some cases used the same burrow over a 14-month period. Others used from 2-4 burrows during the study period and had home ranges that averaged 45 m2. About one-third of the burrows that were monitored were used by the frogs. Blihovde (2006) only found singles, but Kent et al. (1997) and Smith et al. (2021) found as many as four or five frogs using a single Gopher Tortoise burrow in Florida.

Individuals usually remain in burrows or other retreats during the day and forage at night when conditions permit. They can remain active year-round in Florida and other southern localities except during cold bouts (Branch and Hokit 2000, Dodd 2013). Wright (1932) observed frogs sitting at or near the mouth of burrows in Georgia during the day and foraging on the ground surface at night. The juveniles and adults appear to be gape-limited, generalist predators that will eat just about any palatable prey that is small enough to swallow. Dietary studies are largely lacking, but some of the known prey include earthworms, beetles, hemipterans, grasshoppers and other species of frogs (e.g., Anaxyrus quercicus; Carr 1940a, Dodd 2013). They will readily eat toads in captivity (Wright 1932).
General Ecology
Population Ecology: The number of adults that appear annually at local breeding sites is often very small and many populations appear to be maintained by using clusters of local ponds. At the Savannah River Site, no more than 10 adults were collected at a single site in a given year by Semlitsch et al. (1995). Greenberg (2001b) also reported very small annual breeding populations at her study ponds in central Florida. Dodd (1992) captured 44 adults at a small pond in Florida in a single year, and Cash (1994) captured 154 adults from a pond in southeastern Georgia. Palis (1998) collected 301 adults in one year at a study site in the Florida Panhandle. Dodd (2013) noted that relatively large populations sometimes occur in optimum terrestrial upland sandhill habitats.

Smith et al. (2021) recorded observations of Gopher Frogs during gopher tortoise surveys at four conservation sites in Florida and found 274 frogs in 1,097 tortoise burrows that were inspected, with from 17-25% of the burrows having frogs. Population estimates for the sites ranged from 134-742 frogs, and density estimates ranged from 0.50-1.42 frogs per hectare. These were presumably conservative estimates since mammal burrows are also used for cover and frogs that were hiding behind tortoises may not have been detected. The authors used camera scopes to inspect burrows and did not report the percentage of individuals that were juveniles versus adults.

Given that the juveniles and adults move long distances from breeding ponds, connectivity between neighboring ponds that are separated by 1-3 km is likely high if there are no barriers to dispersal. Enge et al. (2017) examined microsatellite variation in Florida populations and found that genetic diversity within populations of gopher frogs in Florida was high. Their analyses revealed only slight to moderate population structure. However, little migration existed between two ponds that were situated only 1.8 km apart but separated by a small stream. Their data suggests that very limited migration may be occurring in some areas where streams, unsuitable wetland or developed habitats appear to be significant barriers to migration. Migration rates appear to be highest in ponds that were connected by contiguous xeric upland habitats.
Community Ecology: Larvae of the Gopher Frog shares breeding ponds with numerous other amphibians but community interactions have not been carefully examined. Predator-prey interactions are discussion above under biotic interactions.
Adverse Environmental Impacts
Habitat Loss: Populations of Gopher Frogs have declined markedly throughout the range of this species except in Florida where populations appear to be relatively stable (Krysko et al. 2019). The losses reflect a combination of factors, including the loss and degradation of terrestrial habitats from agriculture, silviculture, and urbanization. The failure to maintain fire-dependent Longleaf Pine communities has resulted in many pine communities being invaded and replaced by hardwoods that provide low-quality habitat for this species. Numerous seasonal wetlands have been destroyed and fish have been introduced to many breeding sites. The decline of the Gopher Tortoise has eliminated critical microhabitats for the juveniles and adults. Populations outside of Florida tend to be small and geographically isolated from one another and are the most threatened (Dodd 2013).
Habitat Fragmentation: Populations were historically organized as clusters of small breeding populations or sub-populations that had high levels of connectivity between breeding sites. These were embedded in an expansive forested landscape that was dominated by Longleaf Pine-wiregrass communities. Many regional populations have now been severely fragmented by the loss of seasonal wetlands and degradation or destruction of fire-maintained, xeric communities.
Status in North Carolina
NHP State Rank: S2
Global Rank: G2G3
Status in North Carolina: E
Environmental Threats: Populations of the Gopher Frog have declined in most areas of the range, primarily due to the loss, degradation, and fragmentation of aquatic and terrestrial habitats. Most of the historical populations in North Carolina have suffered similar losses, with the remaining populations being small, isolated, and vulnerable to local extinction.
Status Comments: Populations of the Coastal Plain clade that was recognized Richter et al. (2014) have suffered major declines with very few populations remaining in Alabama, Georgia, and South Carolina. As of 2018, Only 7 of 23 historical populations in North Carolina remain, and only 14 of the original 53 known breeding sites remain. Egg mass counts suggest that the total number of adult L. capito in North Carolina is in the order of 200-300 animals. The remaining populations are fragmented and face numerous threats from metapopulation collapse, disease, severe weather, prolonged droughts, development, and lack of proper forest management (NC Wildlife Resources Commission 2018). Only one of the seven recognized populations appears to be somewhat secure. The Gopher Frog is currently under review for listing as threatened or endangered under the U.S. Endangered Species Act.
Stewardship: Local breeding populations that inhabit ephemeral wetlands in xeric sandhills and pines savannas are often very small (e.g., Greenberg 2001a, Semlitsch et al. 1995). The adults sometimes skip breeding for one or more years and juvenile recruitment is erratic. This strongly suggest that viable populations require clusters of local ponds with varying hydroperiods that are embedded in a large fire-maintained upland matrix.

Photo Gallery for Lithobates capito - Gopher Frog

2 photos are shown.

Recorded by: Jeff Beane et al.
Scotland Co.
Recorded by: Jeff Beane et al.
Scotland Co.