Lithobates capito - (LeConte, 1855)
Carolina Gopher Frog
Other English Common Names: Carolina gopher frog, Gopher Frog
Synonym(s): Rana areolata capito ;Rana capito LeConte, 1855
Taxonomic Status: Accepted
Related ITIS Name(s): Lithobates capito (LeConte, 1855) (TSN 775083)
Unique Identifier: ELEMENT_GLOBAL.2.105963
Element Code: AAABH01270
Informal Taxonomy: Animals, Vertebrates - Amphibians - Frogs and Toads
Kingdom Phylum Class Order Family Genus
Animalia Craniata Amphibia Anura Ranidae Lithobates
Genus Size: D - Medium to large genus (21+ species)
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Concept Reference
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.
Concept Reference Code: B90COL01NAUS
Name Used in Concept Reference: Rana capito
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.
Conservation Status

NatureServe Status

Global Status: G3
Global Status Last Reviewed: 18Feb2014
Global Status Last Changed: 15Nov2001
Rounded Global Status: G3 - Vulnerable
Reasons: Restricted to a portion of the southeastern U.S., where distribution and abundance are reduced from historical levels, probably due mainly to loss and degradation of habitat caused by silvicultural practices and fire suppression, combined with reduced gopher tortoise populations; population estimates are not precise but may be fewer than 10,000 individuals remaining.
Nation: United States
National Status: N3 (15Nov2001)

U.S. & Canada State/Province Status
Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
United States Alabama (S2), Florida (S3), Georgia (S2S3), North Carolina (S2), South Carolina (S1), Tennessee (S1)

Other Statuses

IUCN Red List Category: NT - Near threatened

NatureServe Global Conservation Status Factors

Range Extent: 20,000-2,500,000 square km (about 8000-1,000,000 square miles)
Range Extent Comments: Coastal Plain from the southern half of North Carolina (Beaufort 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 (Atlig 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).

Number of Occurrences: 21 - 80
Number of Occurrences Comments: In Alabama, occurs at six breeding sites (M. Bailey pers. comm.). In Florida, extant at 79 sites east of the Apalachicola River (Franz, unpubl. data) and breeds in at least 25 sites west of the Apalachicola River (Palis, pers. comm.). Known from one specimen in Tennessee (Redmond and Scott 1996). Known to occur at five sites in Georgia, including Fort Benning (60-110 breeding adults in a two acre pond; L. Andrews, pers. comm.), Fort Stewart (26-60 adults in three ponds of up to 4 acres; D. Stevenson, pers. comm.), McIntosh County, Baker County, and Charton County (Seyle unpubl. data). Known from two sites (Savannah River Ecology Lab and Santee Coastal Reserve) in South Carolina. In North Carolina, breeds at 11 sites (A. Braswell, unpubl. data).

Population Size: 10,000 - 1,000,000 individuals
Population Size Comments: Total population size is not known but based on data throughout the range is estimated to be 10,000 or more individuals. In Florida, usually uncommon but locally fairly common (Bartlett and Bartlett 1999).

Overall Threat Impact: High
Overall 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). Mechanical site preparation (e.g., roller chopping) destroys burrow openings which may result in entrapment of inhabitants. Routine pine straw harvest and associated removal of herbaceous vegetation degrades habitat quality of the gopher frog (A. Braswell, pers. comm.). Introduction of predatory game fishes (LEPOMIS spp. and MICROPTERUS spp.) is detrimental as these species feed upon gopher frog tadpoles. In some instances, introduction of GAMBUSIA can be detrimental to gopher frogs and other larval amphibians (A. Braswell, pers. comm.). Gopher frog breeding sites are often degraded by off-road recreational vehicle (ORV) use or by sand roads that pass through or adjacent to the ponds (J. Palis, pers. comm.). Vehicular traffic disrupts pond floor micro-topography and eliminates herbaceous vegetation (J. Palis, pers. comm.). Large tires of ORVs may break the organic hardpan that lies below the pond floor. This hardpan prevents water from draining into the sand below the wetland (LaClaire and Franz 1991). Breaking the hardpan could result in a shorter hydroperiod and thus make some wetlands unsuitable for gopher frog reproduction. Loss of herbaceous vegetation from ORV use could also discourage gopher frog reproduction since egg masses are attached to stems of herbaceous vegetation (Bailey 1990; J. Palis, pers. comm.). 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. Heavy grazing by cattle in summer in dried pond basins may reduce or eliminate frog oviposition sites and/or alter pond nutrient cycling.

Short-term Trend: Decline of 10-30%
Short-term Trend Comments: Apparently declining in many areas. Has become less common in recent years in Florida (Bartlett and Bartlett 1999).

Intrinsic Vulnerability: Moderately vulnerable

Environmental Specificity: Very narrow to narrow.

Other NatureServe Conservation Status Information

Inventory Needs: Survey potential breeding sites and remnant upland pine communities to better define present distribution. Use drift fence surveys in order to obtain more accurate population estimates.

Protection Needs: Protect breeding sites and surrounding uplands. Encourage actions that enhance gopher tortoise populations.

Global Range: (20,000-2,500,000 square km (about 8000-1,000,000 square miles)) Coastal Plain from the southern half of North Carolina (Beaufort 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 (Atlig 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).

U.S. States and Canadian Provinces

Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
Color legend for Distribution Map
Endemism: endemic to a single nation

U.S. & Canada State/Province Distribution
United States AL, FL, GA, NC, SC, TN

Range Map
Note: Range depicted for New World only. The scale of the maps may cause narrow coastal ranges or ranges on small islands not to appear. Not all vagrant or small disjunct occurrences are depicted. For migratory birds, some individuals occur outside of the passage migrant range depicted. For information on how to obtain shapefiles of species ranges see our Species Mapping pages at

Range Map Compilers: IUCN, Conservation International, NatureServe, and collaborators, 2004

U.S. Distribution by County Help
State County Name (FIPS Code)
AL Barbour (01005)*, Coffee (01031)*, Covington (01039), Escambia (01053), Shelby (01117)*
FL Alachua (12001), Baker (12003), Bradford (12007), Brevard (12009), Broward (12011), Calhoun (12013), Charlotte (12015), Citrus (12017), Clay (12019), Collier (12021)*, Columbia (12023), DeSoto (12027), Flagler (12035), Gilchrist (12041), Glades (12043), Gulf (12045)*, Hamilton (12047), Hernando (12053), Highlands (12055), Hillsborough (12057), Lake (12069), Lee (12071), Leon (12073), Levy (12075), Liberty (12077), Madison (12079), Manatee (12081), Marion (12083), Martin (12085), Nassau (12089), Okaloosa (12091), Okeechobee (12093), Orange (12095), Osceola (12097), Palm Beach (12099), Pasco (12101), Polk (12105), Putnam (12107), Santa Rosa (12113), Sarasota (12115), Seminole (12117), St. Lucie (12111), Sumter (12119), Suwannee (12121), Taylor (12123), Volusia (12127), Walton (12131), Washington (12133)
GA Baker (13007), Ben Hill (13017)*, Berrien (13019)*, Bleckley (13023)*, Brantley (13025)*, Bryan (13029)*, Burke (13033)*, Camden (13039), Charlton (13049), Chatham (13051)*, Chattahoochee (13053), Early (13099), Emanuel (13107)*, Evans (13109)*, Irwin (13155), Jenkins (13165)*, Laurens (13175)*, Liberty (13179), Long (13183), Marion (13197), Mcintosh (13191), Pierce (13229)*, Richmond (13245)*, Screven (13251)*, Talbot (13263)*, Taylor (13269), Telfair (13271)*, Ware (13299)*, Wheeler (13309), Wilcox (13315)*
NC Beaufort (37013)*, Bladen (37017)*, Brunswick (37019), Carteret (37031), Cumberland (37051), Hoke (37093), Jones (37103)*, New Hanover (37129)*, Onslow (37133), Pender (37141), Robeson (37155), Sampson (37163), Scotland (37165)
SC Aiken (45003), Barnwell (45011)*, Berkeley (45015), Charleston (45019)*, Dorchester (45035)*, Hampton (45049)*, Orangeburg (45075)*
TN Coffee (47031)
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
03 Pamlico (03020104)+*, Lower Neuse (03020204)+*, White Oak River (03020301)+, New River (03020302)+, Upper Cape Fear (03030004)+, Lower Cape Fear (03030005)+, Black (03030006)+, Northeast Cape Fear (03030007)+, Lumber (03040203)+, Little Pee Dee (03040204)+, Coastal Carolina (03040208)+, Santee (03050112)+, Cooper (03050201)+, South Fork Edisto (03050204)+*, Edisto (03050205)+*, Four Hole Swamp (03050206)+*, Salkehatchie (03050207)+*, Middle Savannah (03060106)+, Brier (03060108)+*, Lower Savannah (03060109)+*, Upper Ogeechee (03060201)+*, Lower Ogeechee (03060202)+*, Canoochee (03060203)+, Ogeechee Coastal (03060204)+, Lower Oconee (03070102)+*, Lower Ocmulgee (03070104)+*, Little Ocmulgee (03070105)+, Altamaha (03070106)+, Satilla (03070201)+, Cumberland-St. Simons (03070203)+, St. Marys (03070204)+, Nassau (03070205)+, Upper St. Johns (03080101)+, Oklawaha (03080102)+, Lower St. Johns (03080103)+, Daytona - St. Augustine (03080201)+, Cape Canaveral (03080202)+, Kissimmee (03090101)+, Northern Okeechobee Inflow (03090102)+, Western Okeechobee Inflow (03090103)+, Big Cypress Swamp (03090204)+*, Caloosahatchee (03090205)+, Florida Southeast Coast (03090206)+, Peace (03100101)+, Myakka (03100102)+, Sarasota Bay (03100201)+, Manatee (03100202)+, Little Manatee (03100203)+, Hillsborough (03100205)+, Crystal-Pithlachascotee (03100207)+, Withlacoochee (03100208)+, Waccasassa (03110101)+, Econfina-Steinhatchee (03110102)+, Upper Suwannee (03110201)+, Alapaha (03110202)+, withlacoochee (03110203)+*, Lower Suwannee (03110205)+, Santa Fe (03110206)+, Apalachee Bay-St. Marks (03120001)+, Middle Chattahoochee-Walter F. George Reservoir (03130003)+, Lower Chattahoochee (03130004)+, Upper Flint (03130005)+, Ichawaynochaway (03130009)+, Apalachicola (03130011)+, Chipola (03130012)+, St. Andrew-St. Joseph Bays (03140101)+, Choctawhatchee Bay (03140102)+, Yellow (03140103)+, Blackwater (03140104)+, Pensacola Bay (03140105)+, Pea (03140202)+*, Lower Conecuh (03140304)+, Lower Coosa (03150107)+*, Cahaba (03150202)+*
06 Upper Duck (06040002)+
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
Ecology & Life History
Basic Description: A stocky frog with a usual adult snout-vent length of about 6-9 cm.
General 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.).

Reproduction Comments: Breeding generally occurs in winter and early spring in most of the range. In southern Alabama and 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. In Alabama, males commonly occupied breeding ponds for about a month, most females less than a week (Bailey 1991).

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

Subspecies AESOPUS: In peninsular Florida, breeding most often occurs in the summer months (Godley 1992).

Subspecies CAPITO: 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).

Although an egg mass can contain up to 6000 eggs (Volpe 1958), a typical mass probably contains approximately 1200-2500 eggs (M. Bailey, pers. comm.; A. Braswell, pers. comm.). A larval period of 4 months was observed in Salt Pond on Conecuh National Forest, Alabama (subspecies SEVOSA) (M. Bailey, pers. comm.). In South Carolina (subspecies CAPITO), 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 (subspecies AESOPUS), 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).

Ecology Comments: 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 (Greenberg 2001).

Habitat Type: Freshwater
Non-Migrant: N
Locally Migrant: Y
Long Distance Migrant: N
Mobility and Migration Comments: In Florida, a gopher frog (subspecies AESOPUS) was found at a tortoise burrow 2 km from the pond in which it was marked 4 months earlier (Franz et al. 1988). In Alabama, migrations to and from breeding ponds were positively correlated with rainfall and warming air temperatures (Bailey 1991). In Florida, migrations occurred mainly on warm, rainy nights (Palis and Jensen 1995). Apparently maintains small home range in vicinity of burrow when not breeding.
Palustrine Habitat(s): HERBACEOUS WETLAND, Riparian, TEMPORARY POOL
Terrestrial Habitat(s): Forest - Mixed, Savanna, Shrubland/chaparral, Woodland - Conifer, Woodland - Hardwood, Woodland - Mixed
Special Habitat Factors: Benthic, Burrowing in or using soil, Fallen log/debris
Habitat Comments: 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 0.12 ha to 33.3 ha (Palis and Jensen 1995). 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.).

Adult Food Habits: Invertivore
Immature Food Habits: Herbivore
Food Comments: Metamorphosed frogs eat various invertebrates and occasionally small anurans (Godley 1992). According to Wright and Wright (1949), gopher frogs range "some distance from their burrows in foraging at night." Larvae eat suspended matter, organic debris, algae, minute organisms, and plant tissue. Tadpoles can be raised on boiled spinach (Volpe 1958).
Adult Phenology: Hibernates/aestivates, Nocturnal
Immature Phenology: Hibernates/aestivates, Nocturnal
Phenology Comments: Inactive during cold weather. Mainly nocturnal but sometimes active on the surface in daylight (Einem and Ober 1956).
Colonial Breeder: Y
Length: 9 centimeters
Economic Attributes Not yet assessed
Management Summary
Stewardship Overview: Stewardship activities aimed at restoring/maintaining the ecological integrity of mesic longleaf pine-turkey oak sandhill and associated ephemeral and semi-permanent wetlands will preserve extant populations of gopher frogs. Because gopher frogs are sensitive to anthropogenic alteration of their habitat (Bailey 1991, Godley 1992), perpetuation of existing populations will require preservation of relatively undisturbed xeric longleaf pine-dominated uplands, and graminaceous, fish-free ephemeral to semi-permanent wetlands. Based on the maximum distance gopher frogs are known to move between reproductive and nonreproductive habitat (2 km), at least 12 sq km of terrestrial habitat surrounding each breeding site is probably needed to sustain each breeding population. Long term perpetuation of a viable population of gopher frogs will presumably require protection of a larger area of terrestrial habitat encompassing a suite of alternative breeding sites (Travis 1994). Protection of terrestrial and aquatic habitats will require implementation of the steps outlined in this stewardship report.
Restoration Potential: Gopher frog recovery is directly linked with the ability to preserve existing habitat and restore degraded habitat. Given the drastic decline in the extent of longleaf pine-dominated communities (Ware et al. 1993), elevation of gopher frog populations above present levels is unlikely. However, experimental restoration of a degraded xeric upland longleaf pine sandhill is being conducted by The Nature Conservancy in Florida (G. Seamon, pers. comm.). In addition, Alvin Braswell (pers. comm.) is experimenting with construction of breeding sites in North Carolina, and the U.S. Forest Service is attempting to eliminate introduced predatory fishes from two historic gopher frog breeding sites in Alabama (Bailey, pers. comm.). Should these efforts prove successful, then reintroduction of gopher frogs onto historic sites where extirpated may be feasible on a larger scale.
Preserve Selection & Design Considerations: High quality occurrences include several wetlands within xeric upland pine-dominated communities. In the range of the gopher tortoise, a high quality occurrence would also include a large, viable population of tortoises. Based on the maximum distance adults are known to travel between reproductive and nonreproductive habitat (2 km), each breeding site should be surrounded by at least 12 sq km of terrestrial habitat. A suite of wetlands guards against extirpation at any one breeding site, since animals can immigrate from nearby wetlands or use different sites as conditions vary from year to year. The minimum viable population size needed to sustain a population of gopher frogs longterm is not known. A total of 269 gopher frogs was captured during one breeding season at a drift fence in southern Alabama (Bailey 1990). At a site in South Carolina, however, fewer than 10 adults per year were captured at breeding sites during a 25-year period (Semlitsch et al., in press). Presently, there are no standards for assessing an occurrence based on the number of animals captured at a drift fence, number of egg masses observed, or the number of larvae captured at a breeding site.
Management Requirements: Maintenance of intact xeric longleaf pine-dominated uplands and ephemeral and semi-permanent wetlands by mimicking natural forces, such as lightning-season fire, is the most appropriate form of management. Clearcutting should be replaced with selective timber harvest and natural regeneration enhanced by fire, particularly lightning-season fire. If off-site species such as slash or sand pine have been planted, they should be removed and the site replanted with longleaf pine at densities found in nature. Complete removal of off-site pine in one operation may not be warranted, unless sand pine is the unwanted species. Precommercial thinning of slash pine, coupled with planting longleaf pine, may be less disruptive to the habitat, while providing a canopy that mimics an open longleaf pine stand (M. Bailey, pers. comm.). Mechanical preparation of the soil should be avoided. In addition, silvicultural salvage of naturally fallen trees should be minimized within 12 sq km of breeding ponds, as rotting logs and other debris may be used as temporary refugia for emigrating metamorphs (M. Bailey, pers. comm.).

Seasonality of fire is an important consideration in a prescribed fire plan. Historically, fire was ignited by lightning, most often between May and September (Robbins and Myers 1992). Many components of the groundcover flora, such as wiregrass, flower profusely only after lightning-season fire (Robbins and Myers 1992). Furthermore, lightning-season fires are outside the reproductive period of gopher frogs and thus would not interfere with movements to and from breeding sites. Caution must be exercised, however, when reintroducing fire at fire-suppressed sites. Unnaturally high fuel loads, a consequence of fire exclusion, could endanger longleaf pine if a lightning-season fire is ignited without first reducing the fuel load with cool-season fire(s).

Lightning-season fire may also be important to the maintenance of larval gopher frog habitat. Wetlands used by gopher frogs for reproduction are typically dry or partially dry during the lightning season. A fire passing through these sites when dry would consume herbaceous vegetation and possibly kill woody vegetation. Elimination or suppression of woody vegetation is important in maintaining the open, graminaceous character typical of most gopher frog breeding ponds. In addition, fire releases nutrients bound in plant material. This release of nutrients results in a flush of primary productivity that would be available to herbivorous gopher frog tadpoles the following winter. However, elimination of excessive dead vegetation build-up, with associated release of tannic acid, may be important in sites with marginally low pH (A. Braswell, pers. comm.).

This frog benefits from management that enhances gopher torpoise populations.

Introduction of predatory fishes (e.g., green sunfish) may render a pond unsuitable for breeding (Bailey 1991).

Monitoring Requirements: Adult populations can be censused annually during their arrival at breeding sites by encircling the ponds with plastic drift fences combined with pitfall traps. This method allows capture-mark-release studies to be conducted. Due to annual variation in hydrological conditions, multiple-year surveys are needed to assess the presence or absence of gopher frogs; absence of breeding in one year does not preclude breeding at the same site in subsequent years (Palis and Jensen 1995).

Gopher frog breeding activity can be monitored by surveying for egg masses. Egg mass surveys should be conducted several days after heavy rains during the breeding season. Surveying consists of slowly wading through the wetland and visually investigating vegetation to which eggs are commonly attached. Egg masses are most readily located on overcast, windless days (J. Palis, pers. obs.). Tadpole surveys can be effective when diagnostic characters are known or when tadpoles are reared to transformation to confirm identification (A. Braswell, pers. comm.). Nocturnal vocalization surveys can also be performed during conditions conducive to frog calling activity (usually during or following rains, with air temperatures above 7 C). Caution must be exercised when conducting aural surveys, however, as gopher frogs can produce muted calls from under water that are audible at less than 10 m (J. Palis, pers. obs.; Jensen et al. 1995, Hepetological Review 26:98). To avoid missing submerged calling males, vocalization surveys should be conducted at the pond edge at various points around the pond. On rainy nights when frogs are moving to and from breeding sites, they can often be encountered crossing roads. Gopher frogs can also be monitored at tortoise burrows. Gopher frogs are readily trapped at night at the mouth of tortoise burrows with screen funnel traps, Sherman traps, or bucket traps. Burrow trapping is most successful outside the breeding season since animals abandon upland habitats during this period (Franz 1986).

Management Research Needs: 1. Response to anthropogenic habitat disturbance and altered fire regimes needs study. Drift fence monitoring programs should be implemented at various locations exhibiting varying degrees of anthropogenic disturbance. Tracking of individuals at different places in the range is needed to better assess home range sizes, population densities (A. Braswell, pers. comm.), and the extent of upland habitat needed to support a viable breeding population.

2. Determination of the number of egg masses deposited by each female is needed to provide an indication of the size of the adult breeding population based on egg mass counts.

3. A positive means of identifying live gopher frog tadpoles is needed rangewide to facilitate surveying and monitoring.

4. Additional demographic data are needed to improve our knowledge of the life history, particularly factors that limit population size (e.g., egg, larval, and metamorph survivorship; competition with other species; burrow availability).

5. Additional research efforts should be directed toward understanding water quality needs, larval competitive interactions, density dependent development rates, and factors that may stimulate breeding (A. Braswell, pers. comm.).

Biological Research Needs: Conduct research on habitat requirements (Godley 1992). Investigate the ecology of young frogs.
Population/Occurrence Delineation
Group Name: Ranid Frogs

Use Class: Not applicable
Subtype(s): Breeding Location
Minimum Criteria for an Occurrence: Occurrences are based on evidence of historical presence, or current and likely recurring presence, at a given location. Such evidence minimally includes collection or reliable observation and documentation of one or more individuals (including larvae or eggs) in or near appropriate habitat where the species is presumed to be established and breeding.
Separation Barriers: Busy major highway, especially at night, such that frogs rarely if ever cross successfully; urban development dominated by buildings and pavement; habitat in which site-specific data indicate the frogs virtually never occur.
Separation Distance for Unsuitable Habitat: 1 km
Separation Distance for Suitable Habitat: 5 km
Separation Justification: BARRIERS/UNSUITABLE HABITAT: Rivers may or may not be effective barriers, depending on stream width and flow dynamics; identification of streams as barriers is a subjective determination. Ranid frog species vary in habitat use, but even the most aquatic species may traverse upland habitat when conditions are suitable (Pope and Matthews 2001); natural and seminatural upland habitat generally does not constitute a barrier. Here, unsuitable habitat refers to upland habitat devoid or nearly devoid of wetlands, streams, ponds, or lakes. Bodies of water dominated by predatory fishes may be barriers to some species but suitable habitat for others; in most cases, such waters probably should be regarded as unsuitable habitat.

SUITABLE HABITAT: Suitable habitat includes riparian/riverine corridors, wetlands, and wetland/upland mosaics in which wetland patches are separated by less than 1 km of upland habitat; it also includes any upland habitat regularly used for feeding or wintering (e.g., mesic forest for wood frogs).

MOVEMENTS: Available information indicates that individual ranids occasionally move distances of several km (R. luteiventris: Reaser 1996, cited by Koch et al. 1997; R. blairi: Gillis 1975) but most individuals stay within a few kilometers of their breeding sites (R. aurora draytonii: USFWS, Federal Register, 11 September 2000; R. capito: Franz et al. 1988; R. clamitans: Lamoureux and Madison 1999; R. luteiventris: Turner 1960, Hollenbeck 1974, Bull and Hayes 2001). Similarly, maximum distance between capture points generally is a few kilometers or less (R. aurora: Hayes et al. 2001; USFWS, Federal Register, 11 September 2000; R. catesbeiana: Willis et al. 1956; R. luteiventris: Reaser and Pilliod, in press; Engle 2000; R. muscosa: Pope and Matthews 2001). Dispersal data for juveniles are lacking for most species.

Adult and juvenile R. sylvatica readily traveled in excess of 300 m from their pools of origin (Vasconcelos and Calhoun 2004). Bellis (1965) determined that adult and juvenile R. sylvatica in a peat bog had traveled at least 410 m from the nearest breeding pool. Berven and Grudzien (1990) found that dispersing R. sylvatica juveniles traveled an average of 1,208 m from their natal pools. In the Shenandoah Mountains, data for R. sylvatica indicated that ponds separated by a distance greater than 1,000 m should experience little gene flow (Berven and Grudzien 1991). In contrast, populations in Minnesota were very similar in allelic frequencies, even at distances greater than several kilometers (Squire and Newman 2002). However, sample sizes and number of loci examined were small, and genetic patterns do not necessarily reflect movement distances.

The preponderance of data for ranids indicate that a separation distance of several kilometers may be appropriate for suitable habitat and practical for occurrence delineation, despite occasional movements that are longer and that may allow some genetic interchange between distant populations. The movement data for ranids are here regarded as consistent enough to allow the same separation distance to be used for different species; much of the apparent variation in movements doubtless reflects differences in study methods and in the ability to detect long-distance movements.

Date: 01Apr2005
Author: Hammerson, G.
Population/Occurrence Viability
U.S. Invasive Species Impact Rank (I-Rank) Not yet assessed
NatureServe Conservation Status Factors Edition Date: 17Apr2002
NatureServe Conservation Status Factors Author: Palis, J. G., L. Glass-Godwin, and G. Hammerson
Management Information Edition Date: 11Sep1995
Management Information Edition Author: Palis, J. G.
Element Ecology & Life History Edition Date: 06Feb2003
Element Ecology & Life History Author(s): Hammerson, G.

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