Rana capito Le Conte, 1855: Gopher Frog. Jensen, J. B & Richter, S. C In Amphibian Declines: The Conservation Status of United States Species, pages 536–538. University of California Press, Berkeley, CA, 2005.
Rana capito Le Conte, 1855: Gopher Frog [link]Paper  abstract   bibtex   
Rana capito Le Conte, 1855 Gopher Frog John B. Jensen1 Stephen C. Richter2 A recent genetic study of gopher frog (Rana capito) populations across the current geographic distribution (Mississippi–North Carolina) by Young and Crother (2001) indicated that the Mississippi population was genetically distinct. Young and Crother (2001) therefore elevated the Mississippi population to specific status by resurrecting Rana sevosa Goin and Netting (1940; dusky gopher frog), because this population is the only one remaining in the historical geographic range of Rana sevosa (Louisiana to Mobile County, Alabama) as described by Goin and Netting (1940). Assigning the population discovered in Baldwin County, Alabama, after the publication of Goin and Netting (1940) to R. sevosa or R. capito is difficult because this population was not included in their study and has since gone extinct. Netting and Goin (1942a) assigned this population to R. sevosa based on the then known distribution of R. sevosa (Louisiana to Mobile County, Alabama) and R. capito (Florida to North Carolina). However, populations later discovered in counties proximate to Baldwin County were included in the genetic study of Young and Crother (2001) and were not genetically distinct from all other populations sampled east of the Mobile Bay (thus remaining R. capito). Based on this evidence, we assign the Baldwin County population to R. capito (as indicated by the geographic range maps) and note the inherent uncertainty. It currently is not possible to determine the location of the contact zone between R. sevosa and R. capito, but the extensive Mobile Basin creates a logical barrier to dispersal and thus probably separates the two species. 1. Historical versus Current Distribution. Historically, gopher frogs (Rana capito) were distributed throughout the Gulf and Atlantic Coastal Plain from southeastern Alabama to North Carolina (Bailey, 1991), with one isolated population known from the Ridge and Valley Province of Alabama (Mount, 1975). Though the species continues to be documented from many new localities within its range (for example, Palis and Jensen, 1995; Stevenson and Davis, 1995), this is undoubtedly due to the increased survey efforts of remaining suitable lands rather than recent colonization. The recent discovery of two Rana cf. capito individuals from a site in the Cumberland Plateau of Tennessee (Miller and Campbell, 1996; B.T. Miller, personal communication) is of great interest and may represent a substantial range extension. Young and Crother (2001) recently provided allozyme data supporting the separation of R. capito from both R. areolata (crawfish frogs) and R. sevosa (dusky gopher frogs). 2. Historical versus Current Abundance. Gopher frogs are considered Endangered, Threatened, or of Special Concern in all of the states within their range (Mount, 1975; Martof et al., 1980; Moler, 1992a; Levell, 1997). Though little data are available concerning changes in the species’ abundance, much information exists on the reduction of their habitat and breeding sites. The longleaf pine (Pinus palustris) community, the ecosystem primarily inhabited by gopher frogs, has been reduced to \textless 5% of its historical range (Frost, 1993; Outcalt and Sheffield, 1996; Means, 2003a), and gopher frogs are just one of many component endemic or nearly endemic species declining as a result of this loss. Means and Means (2000) found that the number of breeding populations of gopher frogs in the Munson Sand Hills of panhandle Florida occur at a much lower percentage on sand pine silviculture lands than in nearby native longleaf pine habitat, and that some historical breeding populations have been extirpated. They hypothesized that the principal reason is intensive soil disturbance resulting in the elimination or severe alteration of the upland habitat. Bailey (M.A., 1994) reported that 8 of 14 known breeding sites in Alabama are considered historical, and of the 6 extant ponds, only 3 supported substantial populations. However, since that study, one “historical” site has been reconfirmed as extant, two additional breeding ponds have been discovered, and one previously known breeding site has been extirpated due to the introduction of predatory fish (M.A. Bailey, personal communication; J.B.J., personal observations). Although a few new North Carolina sites have been documented in recent years (Beane and Hoffman, 1995, 1997), Braswell (1993) found only 11 of 32 previously known breeding sites to be active in North Carolina. Twelve sites are known from South Carolina, though only four remain extant (S. Bennett, personal communication). Of 23 historical Georgia gopher frog breeding sites investigated by Seyle (1994), 12 were judged suitable, 8 were considered degraded but marginally suitable, and 3 were judged unsuitable. Only one site was found to contain gopher frogs during two extensive surveys of Georgia's Coastal Plain in late winter and early spring of 1995; however, heavy rains during the autumn of the previous year may have contributed to early breeding, and therefore, low detectability during the survey period (Moulis, 1995a,b). Extensive surveys are lacking for Florida. 3. Life History Features. A. Breeding. Reproduction is aquatic. i. Breeding migrations. Throughout most of their range, gopher frogs migrate from terrestrial habitats to breeding ponds in winter and early spring (primarily January–April). However, breeding migrations may occur at other times of the year following exceptionally heavy rains, especially those associated with tropical disturbances or hurricanes. Fall breeding (typically October–November) has been documented in Alabama (Bailey, 1991), Florida (Palis, 1998; J.B.J., personal observations) and Georgia (J.B.J., personal observations). Gopher frogs often breed during summer in central and south Florida (Godley, 1992). Julian Harrison (personal communication to S. Bennett) has heard calling males, not necessarily indicative of breeding, in every month of the year in South Carolina (S. Bennett, personal communication). Several migrations may occur throughout the breeding season, with males arriving at reproductive sites prior to females and remaining there longer (Bailey, 1991). Local populations increase during the breeding season as frogs from distant populations take up temporary residence in terrestrial retreats along the way (Franz, 1986). Movements are positively correlated with rainfall and warm temperatures and take place between sunset and sunrise (Bailey, 1991). Individuals enter and exit breeding ponds by approximately the same route (Bailey, 1991; Palis, 1998). Franz et al. (1988) recorded the longest known migration distance (2 km) between breeding site and terrestrial retreat. ii. Breeding habitat. Typically, gopher frogs breed in either temporary or semi‑permanent ponds that are shallow, have an open canopy and emergent herbaceous vegetation, and lack large, predatory fish (Moler and Franz, 1987; Bailey, 1991). Cypress (Taxodium ascendens) ponds are often utilized in Florida and southeast Georgia (Godley, 1992; Stevenson and Davis, 1995; J.B.J., personal observations). Anthropogenic habitats, such as ditches and borrow pits, are occasionally used (Means, 1986b; Jensen and LaClaire, 1995; J.B.J., personal observations). B. Eggs. i. Egg deposition sites. Within breeding wetlands, egg masses are typically attached to vertical stems of upright emergent or submergent herbaceous vegetation (Bailey, 1990; J.B.J., personal observations). The stems of inundated shrubs such as St. John's wort (Hypericum spp.) and myrtle-leaf holly (Ilex myrtifolia) also serve as egg deposition substrates (Bailey and Jensen, 1993; Palis, 1998). ii. Clutch size. Egg masses are fist‑sized clusters (Bailey, 1990), and the mean number of eggs/mass have been reported as 1,244 in Georgia (Phillips, 1995); 2,210 in Florida (Palis, 1998); and 1,500–2,000 in North Carolina (Braswell, 1993). Bailey (1989) reported 1,709 eggs for a single mass in Alabama. Apparently, each female deposits only a single egg mass (Palis, 1998). C. Larvae/Metamorphosis. Studies have shown the length of larval period in the lab ranges from 113–225 days (Phillips, 1995). Palis (1998) documented a larval period of 7 mo at a breeding site in western Florida. Cooler water temperatures may inhibit growth (Phillips, 1995). Larvae transform at 28–37 mm SVL (Franz, 1986; Palis and Jensen, 1995; Semlitsch et al., 1995). Larvae are grazing herbivores and utilize dense emergent and submergent vegetation for cover (J.B.J., personal observations). D. Juvenile Habitat. Recently metamorphosed and immature gopher frogs are thought to require the same terrestrial habitats as adults. E. Adult Habitat. Xeric, fire enhanced habitats, especially longleaf pine–turkey oak (Quercus laevis) sandhill (Palis, 1995a). Other habitats used include pine flatwoods, sand pine (Pinus clausa) scrub, and xeric hammocks (Godley, 1992). Adults seek refuge in the burrows of gopher tortoises (Gopherus polyphemus; Franz, 1986; Jackson and Milstrey, 1989), oldfield mice (Peromyscus polionotus; Gentry and Smith, 1968; Lee, 1968b), and crayfish (Godley, 1992; Phillips, 1995), as well as within stump holes (Wright and Wright, 1949). Nickerson and Celino (2003) report that gopher frogs use the hollow interiors of previously submerged and partially decomposed willow (Salix sp.) tree branches as shelters during droughts. F. Home Range Size. Franz et al. (1988) documented a 2-km movement between an upland retreat and a breeding site in Florida. Phillips (1995) followed two adults for 43 d at a Georgia site and found that both remained within a 10-m radius of the specific burrow they selected. Blihovde (1999) also found strong burrow fidelity, especially among females, at sites monitored in central Florida. G. Territories. In terrestrial habitats, rarely is \textgreater 1 individual in occupancy of a single burrow (Wright and Wright,
@incollection{jensen_rana_2005,
	address = {Berkeley, CA},
	title = {Rana capito {Le} {Conte}, 1855: {Gopher} {Frog}},
	url = {http://amphibiaweb.org/cgi/amphib_query?where-genus=Rana&where-species=capito},
	abstract = {Rana capito Le Conte, 1855 Gopher Frog John B. Jensen1 Stephen C. Richter2 A recent genetic study of gopher frog (Rana capito) populations across the current geographic distribution (Mississippi–North Carolina) by Young and Crother (2001) indicated that the Mississippi population was genetically distinct. Young and Crother (2001) therefore elevated the Mississippi population to specific status by resurrecting Rana sevosa Goin and Netting (1940; dusky gopher frog), because this population is the only one remaining in the historical geographic range of Rana sevosa (Louisiana to Mobile County, Alabama) as described by Goin and Netting (1940). Assigning the population discovered in Baldwin County, Alabama, after the publication of Goin and Netting (1940) to R. sevosa or R. capito is difficult because this population was not included in their study and has since gone extinct. Netting and Goin (1942a) assigned this population to R. sevosa based on the then known distribution of R. sevosa (Louisiana to Mobile County, Alabama) and R. capito (Florida to North Carolina). However, populations later discovered in counties proximate to Baldwin County were included in the genetic study of Young and Crother (2001) and were not genetically distinct from all other populations sampled east of the Mobile Bay (thus remaining R. capito). Based on this evidence, we assign the Baldwin County population to R. capito (as indicated by the geographic range maps) and note the inherent uncertainty. It currently is not possible to determine the location of the contact zone between R. sevosa and R. capito, but the extensive Mobile Basin creates a logical barrier to dispersal and thus probably separates the two species. 1. Historical versus Current Distribution. Historically, gopher frogs (Rana capito) were distributed throughout the Gulf and Atlantic Coastal Plain from southeastern Alabama to North Carolina (Bailey, 1991), with one isolated population known from the Ridge and Valley Province of Alabama (Mount, 1975). Though the species continues to be documented from many new localities within its range (for example, Palis and Jensen, 1995; Stevenson and Davis, 1995), this is undoubtedly due to the increased survey efforts of remaining suitable lands rather than recent colonization. The recent discovery of two Rana cf. capito individuals from a site in the Cumberland Plateau of Tennessee (Miller and Campbell, 1996; B.T. Miller, personal communication) is of great interest and may represent a substantial range extension. Young and Crother (2001) recently provided allozyme data supporting the separation of R. capito from both R. areolata (crawfish frogs) and R. sevosa (dusky gopher frogs). 2. Historical versus Current Abundance. Gopher frogs are considered Endangered, Threatened, or of Special Concern in all of the states within their range (Mount, 1975; Martof et al., 1980; Moler, 1992a; Levell, 1997). Though little data are available concerning changes in the species’ abundance, much information exists on the reduction of their habitat and breeding sites. The longleaf pine (Pinus palustris) community, the ecosystem primarily inhabited by gopher frogs, has been reduced to {\textless} 5\% of its historical range (Frost, 1993; Outcalt and Sheffield, 1996; Means, 2003a), and gopher frogs are just one of many component endemic or nearly endemic species declining as a result of this loss. Means and Means (2000) found that the number of breeding populations of gopher frogs in the Munson Sand Hills of panhandle Florida occur at a much lower percentage on sand pine silviculture lands than in nearby native longleaf pine habitat, and that some historical breeding populations have been extirpated. They hypothesized that the principal reason is intensive soil disturbance resulting in the elimination or severe alteration of the upland habitat. Bailey (M.A., 1994) reported that 8 of 14 known breeding sites in Alabama are considered historical, and of the 6 extant ponds, only 3 supported substantial populations. However, since that study, one “historical” site has been reconfirmed as extant, two additional breeding ponds have been discovered, and one previously known breeding site has been extirpated due to the introduction of predatory fish (M.A. Bailey, personal communication; J.B.J., personal observations). Although a few new North Carolina sites have been documented in recent years (Beane and Hoffman, 1995, 1997), Braswell (1993) found only 11 of 32 previously known breeding sites to be active in North Carolina. Twelve sites are known from South Carolina, though only four remain extant (S. Bennett, personal communication). Of 23 historical Georgia gopher frog breeding sites investigated by Seyle (1994), 12 were judged suitable, 8 were considered degraded but marginally suitable, and 3 were judged unsuitable. Only one site was found to contain gopher frogs during two extensive surveys of Georgia's Coastal Plain in late winter and early spring of 1995; however, heavy rains during the autumn of the previous year may have contributed to early breeding, and therefore, low detectability during the survey period (Moulis, 1995a,b). Extensive surveys are lacking for Florida. 3. Life History Features. A. Breeding. Reproduction is aquatic. i. Breeding migrations. Throughout most of their range, gopher frogs migrate from terrestrial habitats to breeding ponds in winter and early spring (primarily January–April). However, breeding migrations may occur at other times of the year following exceptionally heavy rains, especially those associated with tropical disturbances or hurricanes. Fall breeding (typically October–November) has been documented in Alabama (Bailey, 1991), Florida (Palis, 1998; J.B.J., personal observations) and Georgia (J.B.J., personal observations). Gopher frogs often breed during summer in central and south Florida (Godley, 1992). Julian Harrison (personal communication to S. Bennett) has heard calling males, not necessarily indicative of breeding, in every month of the year in South Carolina (S. Bennett, personal communication). Several migrations may occur throughout the breeding season, with males arriving at reproductive sites prior to females and remaining there longer (Bailey, 1991). Local populations increase during the breeding season as frogs from distant populations take up temporary residence in terrestrial retreats along the way (Franz, 1986). Movements are positively correlated with rainfall and warm temperatures and take place between sunset and sunrise (Bailey, 1991). Individuals enter and exit breeding ponds by approximately the same route (Bailey, 1991; Palis, 1998). Franz et al. (1988) recorded the longest known migration distance (2 km) between breeding site and terrestrial retreat. ii. Breeding habitat. Typically, gopher frogs breed in either temporary or semi‑permanent ponds that are shallow, have an open canopy and emergent herbaceous vegetation, and lack large, predatory fish (Moler and Franz, 1987; Bailey, 1991). Cypress (Taxodium ascendens) ponds are often utilized in Florida and southeast Georgia (Godley, 1992; Stevenson and Davis, 1995; J.B.J., personal observations). Anthropogenic habitats, such as ditches and borrow pits, are occasionally used (Means, 1986b; Jensen and LaClaire, 1995; J.B.J., personal observations). B. Eggs. i. Egg deposition sites. Within breeding wetlands, egg masses are typically attached to vertical stems of upright emergent or submergent herbaceous vegetation (Bailey, 1990; J.B.J., personal observations). The stems of inundated shrubs such as St. John's wort (Hypericum spp.) and myrtle-leaf holly (Ilex myrtifolia) also serve as egg deposition substrates (Bailey and Jensen, 1993; Palis, 1998). ii. Clutch size. Egg masses are fist‑sized clusters (Bailey, 1990), and the mean number of eggs/mass have been reported as 1,244 in Georgia (Phillips, 1995); 2,210 in Florida (Palis, 1998); and 1,500–2,000 in North Carolina (Braswell, 1993). Bailey (1989) reported 1,709 eggs for a single mass in Alabama. Apparently, each female deposits only a single egg mass (Palis, 1998). C. Larvae/Metamorphosis. Studies have shown the length of larval period in the lab ranges from 113–225 days (Phillips, 1995). Palis (1998) documented a larval period of 7 mo at a breeding site in western Florida. Cooler water temperatures may inhibit growth (Phillips, 1995). Larvae transform at 28–37 mm SVL (Franz, 1986; Palis and Jensen, 1995; Semlitsch et al., 1995). Larvae are grazing herbivores and utilize dense emergent and submergent vegetation for cover (J.B.J., personal observations). D. Juvenile Habitat. Recently metamorphosed and immature gopher frogs are thought to require the same terrestrial habitats as adults. E. Adult Habitat. Xeric, fire enhanced habitats, especially longleaf pine–turkey oak (Quercus laevis) sandhill (Palis, 1995a). Other habitats used include pine flatwoods, sand pine (Pinus clausa) scrub, and xeric hammocks (Godley, 1992). Adults seek refuge in the burrows of gopher tortoises (Gopherus polyphemus; Franz, 1986; Jackson and Milstrey, 1989), oldfield mice (Peromyscus polionotus; Gentry and Smith, 1968; Lee, 1968b), and crayfish (Godley, 1992; Phillips, 1995), as well as within stump holes (Wright and Wright, 1949). Nickerson and Celino (2003) report that gopher frogs use the hollow interiors of previously submerged and partially decomposed willow (Salix sp.) tree branches as shelters during droughts. F. Home Range Size. Franz et al. (1988) documented a 2-km movement between an upland retreat and a breeding site in Florida. Phillips (1995) followed two adults for 43 d at a Georgia site and found that both remained within a 10-m radius of the specific burrow they selected. Blihovde (1999) also found strong burrow fidelity, especially among females, at sites monitored in central Florida. G. Territories. In terrestrial habitats, rarely is {\textgreater} 1 individual in occupancy of a single burrow (Wright and Wright,},
	booktitle = {Amphibian {Declines}: {The} {Conservation} {Status} of {United} {States} {Species}},
	publisher = {University of California Press},
	author = {Jensen, John B and Richter, Stephen C},
	editor = {Lannoo, M J},
	year = {2005},
	pages = {536--538},
}
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