Lithobates capito

(LeConte, 1855)

Gopher Frog

G2Imperiled (G2G3) Found in 1 roadless area NatureServe Explorer →
G2ImperiledGlobal Rank
VulnerableIUCN
Very highThreat Impact
Identity
Unique IDELEMENT_GLOBAL.2.105963
Element CodeAAABH01270
Record TypeSPECIES
ClassificationSpecies
Classification StatusStandard
Name CategoryVertebrate Animal
IUCNVulnerable
Endemicendemic to a single nation
KingdomAnimalia
PhylumCraniata
ClassAmphibia
OrderAnura
FamilyRanidae
GenusLithobates
Synonyms
Rana areolata capitoRana capitoLeConte, 1855
Other Common Names
Carolina gopher frog (EN) Gopher Frog (EN)
Concept Reference
Collins, J. T. 1990. Standard common and current scientific names for North American amphibians and reptiles. 3rd ed. Society for the Study of Amphibians and Reptiles. Herpetological Circular No. 19. 41 pp.
Taxonomic Comments
Rana capito formerly was included in Rana areolata, and Rana sevosa formerly was included in Rana capito (and Rana areolata).

In a hypothetical phylogenetic diagram based on morphology, Case (1978) placed capito closer to pipiens than to areolata (thereby suggesting the validity of capito as a distinct species) but did not provide any data to justify this hypothesis. Dundee and Rossman (1989) cited evidence of intermediates between areolata and sevosa and regarded them as conspecific (thus not recognizing capito as a distinct species). However, Young and Crother (2001) presented allozyme evidence supporting the separation of Rana capito from R. areolata.

Additionally, Young (1997) and Young and Crother (2001) examined genetic variation in R. capito (sensu lato) from Mississippi, Alabama, Georgia, Florida, and North Carolina and found that populations from Harrison County, Mississippi (the only known population of gopher frogs remaining in the area between Louisiana and the Mobile River delta) were genetically distinct from populations east of the Mobile River drainage (fixed difference at a single locus). Based on this evidence, Young and Crother (2001) resurrected Rana sevosa as a distinct species. "No other specific taxonomic divisions could be determined among the remaining populations of gopher frogs sampled" (USFWS 2000). Hence, recognition of the named subspecies of R. capito (aesopus and capito) appears to be unwarranted.

Devin et al. (2023) identified genetically distinct Florida panhandle and peninsular clusters (lineages) separated by the Aucilla River. Richter et al. (2014) described three lineages, one on the mainland coastal plain and two in peninsular Florida; none of these corresponded to previously named subspecies. Recognizing the three lineages at the species level would influence their conservation status.

Taxonomists continue to debate whether the genus name should be Rana or Lithobates, with a recent argument favoring the retention of Rana (Yuan et al. 2016).
Conservation Status
Rank Method Rank calculation - Biotics v2
Review Date2020-07-06
Change Date2020-07-06
Edition Date2020-07-06
Edition AuthorsYoung, B., W. Dillman, K. Enge, A. Grosse, J. Hall, J. Humphries, B. O'Hanlon, N. Shepard, M. Sisson, A. Smith, J. Smith, D. Sollenberger, C. Threadgill, and T. Wood (2020)
Threat ImpactVery high
Range Extent200,000-2,500,000 square km (about 80,000-1,000,000 square miles)
Number of Occurrences81 - 300
Rank Reasons
This species is restricted to a portion of the southeastern U.S. where its distribution and abundance are greatly reduced from historical levels. The species suffers many ongoing threats, primarily the loss and degradation of habitat caused by silvicultural practices and fire suppression, combined with reduced gopher tortoise populations. Much less than 5% of the range may provide suitable habitat. However, the species appears to have fairly stable populations in protected areas, at least in the Florida portion of the range.
Range Extent Comments
This species occurs on the Coastal Plain of the southeastern US from the southern half of North Carolina (formerly as far north as Beaufort County, currently to Carteret County) to southern Florida (Collier County on the west coast, Broward County on the east coast), west to Alabama; isolated populations in central Alabama (Shelby County), southeastern Alabama, southwestern Georgia, and central Tennessee (Altig and Lohoefener 1983, Bailey 1991, Conant and Collins 1991, Godley 1992, Redmond and Scott 1996, Miller and Campbell 1996). Most of the range is contained within the range of the gopher tortoise (Conant and Collins 1991). This species is likely extirpated from Tennessee where none have been collected since the 1990s (Richter et al. 2014).
Occurrences Comments
Available records with observations of frogs or tadpoles between 2010 and 2020 include 5 occurrences in Alabama, 12 in Georgia, 2 in South Carolina, 7 in North Carolina, and approximately 86 in Florida (Enge et al. 2014). These records are from state natural heritage program and state fish and wildlife agencies, compiled in May 2020. The 2010-2020 time period was chosen to ensure that there was a reasonable chance that the occurrences represent extant populations given the fast pace of habitat conversion and other threats in the region. Additional occurrences without documented records during this time period may also exist.
Threat Impact Comments
Threats are many and include loss of longleaf pine habitat through maximum-yield timber management (e.g., establishment of pine monocultures) and/or decreased frequency of fire; introduction of predatory fishes into breeding ponds; road construction near breeding sites; and declining populations of gopher tortoises whose burrows are used extensively by gopher frogs (Bailey 1991, Godley 1992). Populations are much lower in tree plantations than in naturally regenerated longleaf pine forests with low woody shrub cover (Haggerty et al. 2019). Mechanical site preparation (e.g., roller chopping) destroys burrow openings which may result in entrapment of inhabitants.

Besides direct habitat destruction, housing and urban development restrict land managers' ability to perform prescribed burns to maintain suitable habitat.

Off-road recreational vehicles (ORV) can destroy breeding ponds and kill frogs on sand roads (Roznik and Johnson 2009). Erosion of unpaved roads lying adjacent to breeding sites may result in sedimentation into the ponds. Introduction of sediment is exacerbated by emplacement of wing ditches that divert water from roads into ponds. Road density can also influence the frequency with which prescribed burns can be used as a habitat management tool.

Heavy grazing by cattle in summer in dried pond basins may reduce or eliminate frog oviposition sites and/or alter pond nutrient cycling.

This species is also highly susceptible to ranavirus (Hoverman et al. 2011), although the prevalence of the disease is unknown. In addition, die-off events have been associated with alveolate parasites (Landsberg et al. 2013). Native and nonnative ants (including fire ants) can prey on frogs or drive them from favorable habitat. Rooting behavior by feral hogs can also degrade habitat. Introduction of predatory game fishes (Lepomis spp. and Micropterus spp.) is detrimental as these species feed upon gopher frog tadpoles.

To the extent that it affects precipitation and therefore hydroperiod, climate change may also be a threat (Greenberg et al. 2017). Peninsular Florida has experienced increased droughts, potentially affecting breeding success (Enge et al. 2014). Sea level rise threatens coastal subpopulations in Georgia and North Carolina.
Ecology & Habitat

Description

A stubby frog with short limbs, a large head, and dorsolateral ridges; adults generally are 6-9 cm in snout-vent length (maximum 112 mm) (Conant and Collins 1991). Mass is 47-151 grams (adult male and gravid female, respectively; Palis, unpubl. data). The skin ranges in texture from smooth to warty, and from creamy-white to gray or brown in color. The dorsum and sides are dotted with dark brown or black spots and blotches of various sizes and shapes. The venter is white, cream, or yellowish, and typically is spotted or mottled with dark pigment.

The egg mass is a fist-sized glob, oval to nearly circular in shape. Eggs of gopher frogs and leopard frogs can be distinguished by size (1.44-1.76 mm in diameter in leopard frogs, 1.67-2.7 mm in diameter in gopher frogs; Volpe 1958), and color (leopard frog eggs are dark black, whereas those of gopher frogs are gray to gray-black). Fully expanded gopher frog egg masses are typically larger than those of the leopard frog (pers. obs.).

Gopher frog tadpoles are yellowish-green to olive-green or gray with scattered, relatively large, diffuse black spots on the upper body, tail musculature, and fin. They attain a length of 84 mm before transformation (Wright and Wright 1933). Transforming tadpoles in North Carolina routinely exceed 90 mm in length (A. Braswell, pers. comm.).

The call is a loud snore that lasts up to two seconds and carries nearly 0.4 km (Wright and Wright 1933). However, gopher frogs may also call while submerged beneath the water's surface, which significantly mutes the call (Jensen et al. 1995). The only other species within the range of the gopher frog that have similar calls are the river frog (Rana heckscheri) and pickerel frog (Rana palustris). The river frog, however, breeds in blackwater streams and lakes (K. Dodd, pers. comm.) during the summer months (P. Moler, pers. comm.). Pickerel frogs are not known to breed in the same ponds as gopher frogs (A. Braswell, pers. comm.).

Diagnostic Characteristics

Differs from Rana areolata by the mainly pigmented venter and the absence of light borders around the dark dorsal spots (if present). Differs from Rana heckscheri in lacking white spots on the lips. (Conant and Collins 1991).

The gopher frog is most likely to be confused with the southern leopard frog (Rana utricularia), with which it is sympatric, and the closely related crawfish frog (Rana areolata), which primarily inhabits the Mississippi River drainage from Louisiana northward into Illinois and Indiana. The southern leopard frog has a pointed snout, is more slender with smoother skin, has fewer and more widely scattered spots, and has little or no ventral dark pigment. The dorsal spots of the crawfish frog are encircled by white borders.

Positive identification of larval gopher frogs is difficult because they closely resemble those of the southern leopard frog (pers. obs.). However, as with eggs, fresh hatchlings can be distinguished on the basis of size. In western Florida, hatchling gopher frog tadpoles are 11.9 - 12.7 mm in total length, whereas southern leopard frog hatchlings range from 7.3 - 7.9 mm total length (Palis unpubl. data). Tail depth also differs between hatchlings of the two species. Hatchling gopher frog tadpoles have a deeper tail fin (3.1 - 3.4 mm) than do southern leopard frogs (1.5 - 1.9 mm) (Palis unpubl. data). Larger tadpoles of the two species can be distinguished with limited reliability. Unlike gopher frog tadpoles, southern leopard frog tadpoles often have a light line on each side of the head near the mouth. The skin of gopher frog tadpoles is more transparent than that of southern leopard frogs, which tends to render the internal organs visible through the ventral surface, although this may not always be reliable (A. Braswell pers. comm.). In North Carolina, the origin of the dorsal fin of gopher frogs lies in front of the spiracle, whereas in southern leopard frogs it lies behind the spiracle (A. Braswell, pers. comm.). This character is unreliable in panhandle Florida (J. Palis, pers. obs.).

Habitat

Primary habitat is native xeric upland habitats, particularly longleaf pine-turkey oak sandhill associations; also xeric to mesic longleaf pine flatwoods, sand pine scrub, xeric oak hammocks, and ruderal successional stages of these habitats; generally occurs only where there are gopher tortoises, but rare or absent at most tortoise colonies; absent from most coastal islands and dunes (Godley 1992). Burrows of gopher tortoise or rodents are used for shelter (Gentry and Smith 1968, Lee 1968, Franz 1986); hides also under logs, under or in stumps, and in sewers (Wright and Wright 1949).

Breeding occurs in ephemeral to semi-permanent graminoid-dominated wetlands that lack large predatory fishes (Bailey 1991; Moler and Franz 1987; Palis, unpubl. data). Gopher frogs have also been observed breeding in ditches and borrow pits (Means 1986), and have been heard calling from a recently re-filled, normally permanent wetland following an extreme drought (Franz 1991). The reproductive habitat is best described as a circular or near- circular depression marsh, ranging from 0.02 ha to 33.3 ha (Palis and Jensen 1995, Enge et al. 2014). Dominant emergent graminaceous vegetation typically includes maidencane (Panicum hemitomon), panic grasses (Panicum spp.), bluestem (Andropogon sp.), yellow-eyed grasses (Xyris spp.), pipewort (Eriocaulon compressum), beakrushes (Rhynchospora spp.), and spikerushes (Eleocharis spp.). Most breeding sites have a small component of woody vegetation such as St. John's-wort (Hypericum fasciculatum), myrtle-leaved holly (Ilex myrtifolia), and slash pine that is typically restricted to the edge. Gopher frogs occasionally breed in pond-cypress (Taxodium ascendens)- dominated depressions (Godley 1992, Palis pers. obs.).

Egg masses are laid within an average of 4.5 cm (range 0-20 cm) of the surface, in water 33-78 cm (mean 59 cm) deep, typically attached to vertical stems of graminaceous emergent vegetation (Palis, unpubl. data), as well as to semi-woody weedy vegetation or small (<8 mm) woody stems (e.g., branches of St John's-wort or myrtle-leaved holly). Egg masses may also be deposited on the bottom in shallow water (Bailey, pers. comm.).

Habitat suitability models indicated that number of wetlands, well-drained soil, compatible land cover, and frequent fires were the most important predictors of habitat for the species (Crawford et al. 2020).

Ecology

Egg predators include caddisfly larvae and turtles. Predators on larvae include dragonfly nymphs, diving beetles, introduced game fishes, and undoubtedly many other species. Smaller tadpoles are more susceptible to predation by Notonectids and dragonfly nymphs than are larger tadpoles (Cronin and Travis 1986, Travis et al. 1985).

Among years and ponds, juvenile recruitment is highly variable, but greatest in years with more precipitation (Greenberg 2001, Greenberg et al. 2017).

Predation can be high on recently metamorphosed frogs leaving their breeding ponds, with survivorship only 12.5% during the first month in one study in Florida (Roznik and Johnson 2009).

Reproduction

Breeding generally occurs in winter and early spring in most of the range. In the panhandle of Florida, gopher frogs typically reproduce from January through April (Bailey 1991; Palis, pers. obs.), although Bailey (1991) observed reproduction in southern Alabama in September and October following heavy rains associated with hurricanes. At Eglin Air Force Base, Florida, Palis (1998) observed breeding from October through May, with major breeding events in October, February, and April. Jensen (1994, Herpetol. Rev. 25:161) reported large numbers of egg masses in mid-February in panhandle Florida. Breeding typically occurs between mid-January and April in North and South Carolina (Semlitsch et al. 1995), but calling has been heard in every month of the year in South Carolina (A. Braswell, pers. comm.; S. Bennett, pers. comm.). In South Carolina, breeding did not occur every year at a particular site, breeding events lasted only a short period of less than two weeks, and local breeding populations were very small (Semlitsch et al. 1995). In Florida, breeding ponds ranges in size from 0.02 to 12.22 ha, with a mean of 1.01 ha (Enge et al. 2014).

Individuals spend about 1.5-3.5 weeks in the breeding ponds (Bailey 1991, Palis and Jensen 1995).

Although an egg mass can contain up to 6,000 eggs (Volpe 1958), a typical mass probably contains approximately 1,200-2,500 eggs (M. Bailey, pers. comm.; A. Braswell, pers. comm.). In South Carolina, the larval period was estimated to last 87-113 days (Semlitsch et al. 1995); metamorphosing and emigrating juveniles were found from late May to late July; successful recruitment of metamorphs into the adult population at a particular site occurred only rarely. In north-central Florida, metamorphosis occurs at a snout-vent length of 35-40 mm (Franz 1986). At a mean monthly growth rate of 1.5 mm (Franz 1986), two years are required for gopher frogs to attain the minimum size of sexual maturity (70-75 mm SVL) (Franz, unpubl. data).
Terrestrial Habitats
Forest - MixedWoodland - HardwoodWoodland - ConiferWoodland - MixedShrubland/chaparralSavanna
Palustrine Habitats
TEMPORARY POOLHERBACEOUS WETLANDRiparian
Other Nations (1)
United StatesN2
ProvinceRankNative
AlabamaS2Yes
South CarolinaS1Yes
GeorgiaS1Yes
North CarolinaS2Yes
FloridaS3Yes
TennesseeS1Yes
Threat Assessments
ThreatScopeSeverityTiming
1 - Residential & commercial developmentRestricted (11-30%)Serious or 31-70% pop. declineHigh (continuing)
2 - Agriculture & aquacultureRestricted - smallModerate or 11-30% pop. declineHigh (continuing)
2.2 - Wood & pulp plantationsLarge - restrictedSerious or 31-70% pop. declineInsignificant/negligible or past
2.3 - Livestock farming & ranchingRestricted - smallModerate or 11-30% pop. declineHigh (continuing)
4 - Transportation & service corridorsLarge (31-70%)Serious - moderateHigh (continuing)
4.1 - Roads & railroadsLarge (31-70%)Serious - moderateHigh (continuing)
6 - Human intrusions & disturbanceRestricted - smallSerious - moderateHigh (continuing)
6.1 - Recreational activitiesRestricted - smallSerious - moderateHigh (continuing)
7 - Natural system modificationsPervasive (71-100%)Extreme or 71-100% pop. declineHigh (continuing)
7.1 - Fire & fire suppressionPervasive (71-100%)Extreme or 71-100% pop. declineHigh (continuing)
8 - Invasive & other problematic species, genes & diseasesPervasive - largeUnknownHigh (continuing)
8.1 - Invasive non-native/alien species/diseasesPervasive (71-100%)UnknownHigh (continuing)
8.2 - Problematic native species/diseasesPervasive - largeUnknownHigh (continuing)
11 - Climate change & severe weatherPervasive (71-100%)UnknownHigh (continuing)

Roadless Areas (1)
Florida (1)
AreaForestAcres
Alexander Springs CreekOcala National Forest2,954
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