Postbreeding Movements of the Dark Gopher Frog, Rana Sevosa Goin and Netting: Implications for Conservation and Management

Home , Gopher frog, Rana (genus)

Journal o fieryefology, Vol. 35, No. 2, pp. 336-321, 2001 CopyriJt 2001 Society for the Study of Amphibians and Reptiles

Postbreeding Movements of the Dark Gopher Frog, Rana sevosa Goin and Netting: Implications for Conservation and Management

'Departmmzt of Biological Sciences, Southeastern Louisiana Uniwsity, SLU 10736, Hammod, Louisiamla 70403-0736, USA 4USDA Forest Service, Southern Research Station, Southern Institute of Forest Genetics, 23332 Highmy 67, Saucier, Mississippi 39574, USA

ABSTRACT.-Conservation plans for amphibians often focus on activities at the breeding site, but for species that use temstrial habitats for much of the year, an understanding of nonbreeding habitat use is also essential. We used radio telemetry to study the postbreeding movements of individuals of the only known population of dark gopher frogs, Rana sevosa, during two breeding seasons (1994 and 1996). Move- ments away from the pond were relatively short (< 300 m) and usually occurred within a two-day period after frogs initially exited the breeding pond. However, dispersal distances for some individuals may have been constrained by a recent clearcut on adjacent private property. Final recorded locations for all individuals were underground retreats associated with stump holes, root mounds of fallen trees, or mammal burrows in surrounding upland areas. When implementing a conservation plan for Rana sevosa and other amphibians with similar habitat utilization patterns, we recommend that a temstrial buffer zone of pro- tection include the aquatic breeding site and adjacent nonbreeding season habitat. When the habitat is fragmented, the buffer zone should include additional habitat to lessen edge effects and provide connec- tivity between critical habitats. For our study site, we recommend a 1000-m buffer zone around the primary breeding site and each of two other potential breeding ponds.

For amphibians that breed in temporary habitat selection. Although" these data are critical ponds, the hibernation sites, breeding sites, and in understanding habitat use by frogs, few te- foraging areas may be temporally and spatially lemetry data are yet published. One study of separated, and individuals must migrate to and ranid frogs (Rana clamitans) using terrestrial from these sites in seasonal cycles (Semlitsch, habitats showed a maximal dispersal distance of 1981; Sinsch, 1990). Because individuals are gen- 560 m, although there was considerable inter- erally concentrated only during the breeding individual variation (Lamoureux and Madison, season, many studies of amphibian biology take 1999). place in and adjacent to breeding sites and not Gopher frogs are rare and poorly studied in the nonbreeding habitats. However, design- frogs whose geographic range once extended ing a comprehensive management plan for any throughout the southeastern coastal plain from amphibian that uses terrestrial habitats for North Carolina to Louisiana. Although once mu& of the year requires an understanding of common to abundant in coastal Mississippi and habitat use during both breeding and nonbreed- Louisiana (Allen, 1932; Dundee and Rossman, ing seasons (Dodd, 1996; Dodd and Cade, 1998). 1989), breeding populations of gopher frogs Postbreeding movements of amphibians into west of Alabama have been severely reduced in adjacent terrestrial habitats are poorly under- numbers. They are thought to be extirpated in stood, and distances that most species normally Louisiana and are now known to occu; only at disperse are unknown (Dodd, 1996; Dodd and a single location in the De Soto National Forest Cade, 1998). Although mark-recapture studies in Harrison County, Mississippi. A recent study have provided valuable data on dispersal dis- by Young and Crother (2001) indicated that this tances, recent advances in radio-telemetric methods make this a more suitable method for population is genetically distinct from other studying postbreeding movement pattems and populations of gopher frogs and that it should be recognized as Rana smsa Goin and Netting. Thus, &ere 1s a v~talneed for ~nformat~on0% Author Present both breeding and postbreedlng actlvltles ment of Zoology, University of Oklahoma, Norman, Oklahoma 73019, USA, E-mail richter@ou edu Because gopher frogs are secretwe and d~ffi- Present Address Applied Ecology Rewarch to locate of the breeding Group, Un~vers~tyof Canberra, AustralIan Capit,,l knowledge of their ecology 1s generally llmlted Territory 2601, Australia to stud~esof reproductwe ecology at breed~ng

MOVEMENTS AND CONSEIiVATlON OF GOPIiER FliOGS 31 7 s~tes(eg, Ra~ley,1991; Sernl~tschct a1 , 1995, harness plus transm~tterfor all ~ndivtdualswas Young, 1997, I'al~s, 1998, Rtcliter, 1998) During 3-5'%, of the total mass of the frog Th~sis well the nonbreed~ngseason, gopher frogs have been below the general rule of 1O1%as tlie maximum reported to take shelter in the burrows of go- we~ghtratio of traiism~tterpackages to body pher torto~scs, Gop\lcrlri po/t/p/rc7~iilli(Franz, Illass (R~cliardset al., 1994) 1986) D~staiicesmoved from the breed~ngs~tc To determ~nepotcnt~al ~iegat~ve effects on go- after reproduct~onare unknown, except for two pher trogs, thc harness design n7as tested on ~nd~vidualsin Flor~dathat were' tound 1 6 and southern leopard frogs, Roll(? sf~l~~~izo~~~~~l~oio,111 2 0 l

I Iistance M,iss SVI. N.1 irom pitn(3 l'r

der leaf l~tterFlowevcr, ftnal recordcd locat~otis the north, three tct thcl south, and two to tlie for all ~iid~vtdualswere underground retreats west (Fig. 1). assoctatecl w~thstump holes, root moutidi of The skeletoti of one frog (#12) was found 44 fallen trees, or small mammal burrows. days following initla1 release Ca~~seof death is Two ~ndlvtduals(#4 and #5 111 Table 1) in 1994 unknown, but its burrow was submerged by remained within the area of the drled pond (49 heavy rain tliree times following the frog's re- m from the pond center), aiici In 1996, one frog treat undergrouncl. After the water receded for (#8) remalned wrthtn the area of the complete the final time, tlie dead frog was found outside drift fence (82 ni from the pond center) but not of the burrow wtthin the actual pond area (Ftg 1) The hnal rfc>cfs qf Ck.nrc~if\find Cor~fro\k~?dRurl7i.-Bc- postt~onsof all other ~ndtv~dualswere beyond tween 30 March and 2 Apr~l1994, the prlvate the drift-fenced area Of these, four lnoved to land 200 m north of Glen4 Pond (then owned by 11') was clearcut, roller-chopped, arid bedded for ptne treei w~thlieavy machinery No frogs with tranimitteri vvcrc obier\.ed entering the

I ic, I I'rnal locallt1c~5ot racilo-lrackcd gitplrc,r trogb (m1nibelc.d dot\) ~clatlveto the 'rre't enclosed by 1 I(. 2 Amount of r'lrn (cnr) 'rnd rno\enicnt~ot thc drltt fence (lrorr/o~it~~llylratchcd alc,a), pond center tc~lt,mcte~tdgoplici frog5 tlirougliot~tthe stn~lypci I- (X), acijacurt prnate land (.relt~callyhatchti1 area), od N~linbcrot flogs rno\lng on '1 glven day reprc- ,ind USIIA l orest \el\ rcc lctndi (~~tihatclii~cla1 ca) ~tntcciby numbers along the r,trnlall Illre MOVEMEN'TS AND CONSEIZVATION OF GOPHER FROGS 31 9

clearcut, but of the frogs leavlng tlie ~mmediate Our niaxtnitrni d~stanct.is mucl? less than has victil~tyof the pond, four moved toward thls been observcd for incltvldual gopher frogs 111 general area, three of whtch moved qulte close Florrda (1 6 and 2 0 km; Carr, 1940, Frana ct al , to the clearcut (Fig 1) In 1994, two ~iidiv~duals1988) Tliese differences havc at least two expla- (#1 and #6) were tracked to final positions 10 m nat~ons (I) ol?scr\7atlons on gopher frogs in and 30 m from this 'trea In 1996, one frog (#11) Flonda rc.presented unusually long movements, movcd twitce (dur~ngtwo separate mrgratlon and (2) gopher frogs 111 Florida generally have per~ods)to the L7o~indaryseparating tlic US De- longer dtsperial d~stanccsthan gopht-r frogs In 1.xrtnient of Agrlculturc Forest Serv~ce(TS) and Mss5issippi We lack data to sufflcrently teit I1"roperty and took res~denccIn a11abandoned these hypotheses, although recent stud~esby B mamma\ burrow literally under the bou~id~lryBlthovde (pers comm ) suggest that long-d~s- fence (Fig 1) Thts suggests that some frogs tancc niovements (: 1 km) are rare In gopher ~mglitIi'tve moved farther north had the 11' land frogs in central Flortda not been clearcut at thc time We achleved our orig~nalgoals by sampl~ng Becauic suitable hab~tdtfor gopher frogs and postbrecdlng inlgratlons, but caut~onmust be other spccles In th~slongleaf psne ecoiyste~i~1s taken In applysng thew data to nlaxlmum maliita~iiedby period~cfire, a growlng season movement distances or total habitat used prescribed burn of the entire study area was througl~outthe year Sttiscl? (1990) expla~nedtlie conducted by the FS on 26 Aprll 1994, clurlng potential use of three "spatial units" (breeding which tiine three frogs had funct~onaltransmlt- s~te,nutr~tlon site, anct hibernation site) 111 frcsgs ters attached All three frogs wcre act~vcafter througliout a year Br~efly,after breeding, maiiy the burn, and two postburii movemetits were anurans mtgrate from the pond to an area to recorded for each of two frogs The thlrd frog feed and may subsequently m~grateto a d~ffer- was not recaptured but was seen at a burrow cnt area for hlbernatlon Gopher frogs breed eiitrance tollowtng the burn. Thus, as mtght be during the winter, thus, they do not liibertiate expected for frogs that occur In a fire-dependent but do have periods of inactlvsty. We are uncer- habitat, there was no evldence that mortality re- tain to what extent R stvosa uses different spa- sulted froin the burn. tsal units durlng periods beyond our study. A Bchm~oral Obsc.r.iat1o17.:-After movemeiit to year-round examination of migratory behavior their upland refuges, most gopher frogs did not would be required for sucli data Some tsine af- remain underground but were often seen out- ter removal of transmitters (146 days), frogs s~deof thew burrows throughout the study Dls- were no longer found perched outslde of thesr tinctwe resting areas were observcd outstde of burrows nor were tlie charactertstic worii areas each frog's burrow, conststtng of so11 cleared of near the burrows seen. This llidicatcs erther that vegetatlc>nand smoothed by the frog's constant the frogs remained at these bu~rowsunder- use In 1994, tlie activ~tyof frogs outirde therr gr~undor that they moved to another site, as IS burrows was not closely monitored, although known for c011i11ioii toads, NclfO 171lfO (Sinsch, ~nchvrctual frogs were often observed In tlie 1990) open In 1996, a11 frogs (except #12) wert, corn- Ejcccts of Hahrtnt Altrratlol~-In areas such as monly scen bask~iigIn direct sunl~ghtoutstdc nat~onal forests, where timber inanagemcnt their respective burrows practices arc impletiiented, ,111 understartdlng of Althotigh no negative effects of the transniit- habitat use dur~ngboth brecdtng and noiibreed- ters were found In 1994, in 1996 three frogs ing per~ods1s essential Forestry pract~ces,such werc found wtth skin abrastons 21-26 days fol- as tlie removal of trees and controlled bur~i~~ig, lowi~igreleasc When abrasions were found, tlie are known to have dctri~nerttaleffects on arn- transin~ttcrs werc rclnoved, 'n~d frogs were phlblans but 111 some cases may be necessary for mon~tctred 111 the field All frogs had healccl tlwr burv1v~11it1 human-altered liab~tats(Wcll- when recaptured 8-14 days follow~ngtransmtt- dlck, 1997, and refcre~iccstherein) Knowledge ter rerno\.al and were seen outsrde of their br~r- of the beliav~orof the spec1c.s L~llowsInanage- rows 8-46 clays 1atc.r No turther <~tteniptsat tc- ment pract~cesto be pcrfornied wtieii tlie frogs Icmetry were made are least hkely to be affected. Two major liabltat dtsturbanccs occurred durlng tlic. st~ldythat may have affectcd gopher frogs clearcuttlng 111 Mtrric~trri,irt llllrl Ac t~iilty l'attc7r 115 -All post- an adjacent habrtat and controlled b~~rn~iigin brecctlng movements of gopher frogs occur rcd tlic rnlined~atc\icin~ty ot the pond We found wrtliln 700 111 of Glen's Pond, and dlst'n~ces 1-10 short-term nipa act of prcscrlbed burmrlg on inoved were slm~larbetween years (Tablc 1) 1si gopher frogi All tliree indi\r~dualsrnonitorcd both ycari, most movements were c~ssociatcd during tlie bur11 snrv~vedwith 110 apparelit with rzltntall evcmts, an ~~ssoci;ttsoiiseen 111 other harm arict resunlcd normal inovcrneiits wttli~n ;Inurans (e g , I'cclimann and Sernlltsch, 1986) 1-2 days Ho\vcver, we crn?~ictt cletcrnilne 320 S. C. RICHTER ET AL whether tlie burn may have ~mpactedforag~ng success (el tlier p)s~t~vely or ncgat~vely) or whether there werc any long-term effects Also, gtve~ithe low samplcx s~/cof frogs at tlie burn stte, we cannot generalr/e from our results Dr~scnlland lioberts (1997) tound more exten- s~vernortal~ty from burns in GL~OCIIIIIIIllriln m Australla Our data suggest that at least some frog movements were affected by the clearc~~tNo telenietcred frogs entered tlie clearcut, although sonic moved to arcas along thc bounclary between tlie FS land and tlic clearcut (see Ftg 1) We stro~iglysuspect tliat goplier frogs lnay have ~tscdthe clearcut area pr~orto habttat alteration in 1994, grvcti tliat (1) thc clearcut 15 'rvcll w~thtn the rC~diusof maxllnuni d~spersal d~stances troiii tlic pond (I e, < 299 m), (2) one frog tw~ce moved to' tlie very edge of the clearcut, 'lnd (3) therc, were gopher tortotse burrows in thrs area bc,fore the liabitat was clearcut and bedded (go- Flc, '3 I'rol.7used 1000-in terrestr~albuffer /oncrs pher frogs are reported to use gophe~torto~se for C;lcn's I'oncl (I), 5econdary Pond (2), and I'ony burrows extens~velyIn Flortda, Fran~,1986, I3 lianch I'ond (7) Tlie inset box around Glen's I'ond (I) B~lhovdt: per5 comm ) Clearcuts arc known to indrcates tlie area clcplcted In I'lgure 1 strongly tnflucncc the abundance of salamanders (e.g., Petranka et dl, 1993); however, addl- d~stanceswe found In our study niay have been tlonal data on ~mpactsof clearcuts on anurans truncated by the presence of the clearcut to the IS a niajor management need nortli of our study site. Second, the privde land Conscr-ontto~~lrrtplrcntton~ -Protectton of the 200 ni north of Glen's Pond IS be~ngcurrently ~nimed~atev~crntty of wetlands w~tlioutconsid- dt.veloped as a ret~rementcommun~ty, wlitch cratton of t1ie acljacent terrestr~alhab~tat (as IS means that thcse trogs may be exposed to hu- often the case wlicn federal statutes regulate, nia~i-~nducedstresses s~ichas fert~lt/errunoff thts protectron) 1s msuft~c~etitto ma~nt't~nmany and recreat~onalacltvittes Thus, an addit~onal wetland ipectes (Burke and G~bbons, 1995, buffer is requtred to lessen edge effects (Murcia, Dodd, 1996, Dodd and Cade, 1998) Because an- 1995) I hrrd, two other poncls in the v~c~nityof urans that breed In temporary w7ctla1ids spend Glen's I'ond appear to be su~tablebreecl~ng s~tcs much of the ycar In terrestr~alIiabttats, a buffer but arc not currently being used by gopher /one (as recommended by Burke and G~bbons, frogs These ponds (lieserve Pond and Pony 199i, IDodd and Cade, 1998, Scnil~tsch,1998) at lianch Pond, 10 ancl 1 6 kni fro111 Glen's I'ond, least completely eiicornpasstng tlie nonbreed~ng rcspectively) iieecl to bc ~ncluded111 a stnglc season hab~tat1s cssentlal tor thetr protectton bufter /one tliat would Increase the l~kel~liood Tlie sl/e of sucli bufter /ones depends on sev- of (re)coIoiit/at~on of these s~tesOur recom- eral factors, notably the degree of fragnientat~on mendcltlon 1s for a terrestr~albuffer /one for and the riiob~lttyof tlie spccles under cons~der- each pond of 1000 111, wli~clienconipasses the at~onThe d~recteffects of d~sturbanceto r~sed nonbreedtng season hab~tatdnd an add~tron~il liab~tat(e g , loss of m~croliab~tat,prey av;t~lab~l-buffcr to reduce edge. effects (I'lg 3) These buff- ~ty,change In liyc~ropc.r~od)m'ly be obv~ous,but er /ones would also crc,dtc corrrdors between alteratton of tlie adjacent habitat nus st also he the three ponds tli'it ,ire 'it least 800 m froni tlie cons~clered If thc liab~tdtacljncent lo tlic buffer edge of altered h,ibttats /one 1s altc,rccl (c g , cleCircutor dcvcloped), or- gdnlsms w~thinthe now frag~iientedliab~tat arc Rtki~o~c~B~iigiii(~iti~-We want to thank tlic potentrally exposed to clianges III nitcrctcl~ni~~temany peoplc who helpcd in tlie t~eld,espec~ally s~~clias w~de~ fluctuat~ons i11id changes In mean j S Il)oody, M I'tlgr~ni,'2nd J 13cmuth We also Iiu~ii~d~ty,litter mo~sture,sol~r rad~ation, and woulcl like to thank li Uoudr~ckand others at teniperaturc (I c,, edge etft>cts, Sciunclers et al , tlie Harr~sonFxp~r~tiic~ntaI Forest for tlie vcry 1991, Mntl'lck, 19'13, MLI~~I;~,IC)95) gr~~ctot~s~~seof tlie~r c'ib~n and ICiboratorjifac~l- Moi7o;{c~ii1(,iiiof tl~~,MI~~I\~I~I;JI l'op~rlni~oii -A)- ~t~esT1i'~nks to J S Doody, J C;lldwcll, ancl C thougli goplier trogs at C;leni I'ond mo\d fatr- l.edry tor Iielpful comments on 'In early \ers~on ly short J~stclnces(all < 299 ni), tlircc factors of tli~stnanuscrrpt We arc grateful to tlic US argue. tor a much largci- buftcr /one F~rst,the Ftsh ;~ndW~ldltte Scrv~ce, USDA 12c)restScrvrte, MOVEMENTS AND CONSEIIVATION OF GOPHER FROGS and the Mississippi Department of Wildlife, MATLACK,G. 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