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).