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EFFECTS OF PRESCRIBED FIRE ONHERPETOFAUNA IN BOTTOMLANDHARDWOOD FORESTSAuthor(s) Kurtis R Moseley Steven B Castleberry Sara HSchweitzerSource Southeastern Naturalist 2(4)475-486 2003Published By Eagle Hill InstituteDOI httpdxdoiorg1016561528-7092(2003)002[0475EOPFOH]20CO2URL httpwwwbiooneorgdoifull1016561528-7092282003290025B04753AEOPFOH5D20CO3B2

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SOUTHEASTERN NATURALIST2003 2(4)475486

EFFECTS OF PRESCRIBED FIRE ON HERPETOFAUNAIN BOTTOMLAND HARDWOOD FORESTS

KURTIS R MOSELEY1 STEVEN B CASTLEBERRY

1AND Sara H Schweitzer1

ABSTRACT - Abundance diversity and richness of herpetofaunal species werecompared between burned and unburned bottomland hardwood stands at Di-Lane Plantation Wildlife Management Area (WMA) in Burke County Georgiafrom July to October 2001 Two trap clusters each consisting of a drift fencepitfall array four coverboards and three polyvinyl chloride (PVC) pipe refugiawere randomly placed within each of three burned and three unburned standsHabitat variables were measured within a 004-ha circular plot centered on eachtrap cluster A total of 346 individuals representing 21 species was captured in348 array nights Amphibian abundance diversity and richness were similarbetween burned and unburned stands Reptile abundance and diversity weregreater in burned stands whereas reptile species richness was similar betweenburned and unburned stands Winter burns did not significantly affect coarsewoody debris volumes between burned and unburned stands and may haveaccounted for similar amphibian abundance diversity and richness Greaterreptile abundance and diversity in burned stands likely was a result of decreasedground cover providing greater thermoregulatory opportunities

INTRODUCTION

Prescribed burning is used by land managers to control hardwoodinvasion reduce litter buildup recycle nutrients destroy on-site slashprepare seed beds and improve wildlife habitat (Nyland 1996 Smith etal 1997 Stoddard 1962 Van Lear and Waldrop 1989 Walker 1999)Prescribed burning also provides a means of maintaining habitat diver-sity where natural fire cycles have been excluded A variety of south-eastern forest ecosystems such as the Florida sandhill longleaf pine(Pinus palustris Mill)-wiregrass (Aristida beyrichiana Trin andRupr) and slash pine (Pinus elliotii Engelm) flatwood communitiesdepend on frequent fire regimes to eliminate hardwood competitors andencourage regeneration Fire maintained communities are important tomany herpetofaunal species (Means and Campbell 1981 Mushinsky1985) While the effects of fire on herpetofaunal communities in thesesystems have been examined effects of fire on herpetofauna in hard-wood stands of the southeastern Coastal Plain are relatively unknown(deMaynadier and Hunter 1995 Russell et al 1999)

1DB Warnell School of Forest Resources University of Georgia Athens GA30602-2152 Corresponding author - krm9575owlforestryugaedu

Southeastern Naturalist Vol 2 No 4476

When fire is limited or eliminated altogether hardwood speciesbecome established in pine (Pinus spp) dominated stands Shade toler-ant hardwoods outcompete and eventually eliminate the more shadeintolerant pine species If left undisturbed this eventually leads tosuccession of a hardwood dominated forest that may not provide ad-equate habitat for herpetofauna adapted to fire-maintained pine commu-nities (Means and Campbell 1981) While some species may declinehardwood stands have been shown to exhibit a greater abundance ofherpetofauna than pine stands Miller (1997 unpub data) found thathardwood sites sampled at Di-Lane Plantation WMA in the GeorgiaCoastal Plain contained greater herpetofauna diversity than planted pinestands An oak (Quercus spp)-hickory (Carya spp) forest at the Savan-nah River Site South Carolina had 50 more amphibians than plantedpine areas (Bennett et al 1980) The moist cool microenvironmentprovided by hardwood stands is important to a variety of herpetofaunalspecies (Bennett et al 1980)

Microhabitat structure such as snags decaying logs and leaf littermay be drastically altered by fire (Greenberg et al 1994 Smith et al1997) Reduction of microhabitat structure leads to a decrease in on-sitemoisture important to amphibians The epidermis of amphibians mustremain moist to respire making them highly susceptible to dry condi-tions (Duellman and Trueb 1994 Spotila 1972) Reptiles howeverpossess a relatively impermeable skin that increases their tolerance todry conditions (Pough et al 2001)

The objective of our study was to compare herpetofaunal abundancediversity and richness in bottomland hardwood stands subjected toprescribed burning to those in unburned stands Because of the potentialnegative effect of fire on vital microhabitat features such as leaf litterand coarse woody debris (CWD) we hypothesized that the stands sub-jected to prescribed burns would have a lower abundance ofherpetofauna than the unburned stands We also hypothesized that thediversity of species would not decline in burned stands because previ-ously excluded fire-adapted species would be present in burned stands

FIELD-SITE DESCRIPTION

Our study was conducted on Di-Lane Plantation WMA (hereafterDi-Lane) a 3282-ha tract in Burke County Georgia located in theupper Coastal Plain Topography generally is flat with elevations rang-ing from 58 to 70 m Experimental stands were poorly drained bottom-land hardwoods interspersed in a matrix of closed canopy pine planta-tions and open fields The canopy of hardwood stands consisted of 74 mixed hardwood species and 24 loblolly pine (Pinus taeda L)Dominant hardwood canopy tree species included water oak (Quercus

KR Moseley SB Castleberry and SH Schweitzer2003 477

nigra L) laurel oak (Q laurifolia Michx) and sweetgum (Liquidam-bar styraciflua L) Midstory tree species included water oak laurel oaksweetgum and black gum (Nyssa sylvatica Marsh) The understoryconsisted of blueberry (Vaccinium spp) greenbrier (Smilax spp)grapevine (Vitis spp) American beautyberry (Callicarpa americana L)and other seedling hardwood species

In an effort to improve northern bobwhite (Colinus virginianus L)habitat on Di-Lane pine and hardwood stands were burned to increasecanopy openings and stimulate herbaceous growth Stands were burnedduring winter using aerial ignition Burns in bottomland hardwood standswere light due to moist soil and litter conditions and numerous ephemeralponds present during winter Treatment stands considered for the studywere those burned every two to three years for the past nine years withthe most recent burn occurring in January 2001 (J Bearden GeorgiaDepartment of Natural Resources pers comm June 2001) Three standswere randomly chosen from all stands meeting the criteria (Fig 1)Treatment stand sizes were 364 ha 445 ha and 931 ha Three standswith similar plant species composition hydrology and elevation to treat-ment stands but which had not been burned in at least 9 years wererandomly selected as reference stands All stands contained numerouspools of standing water from late fall 2000 through early spring 2001

Figure 1 Di-Lane Wildlife Management Area Burke County Georgia andthe locations of trap clusters in burned and unburned bottomland hardwoodstands

Southeastern Naturalist Vol 2 No 4478

METHODS

Two trap clusters each consisting of a drift fence pitfall array ply-wood coverboards and polyvinyl chloride (PVC) pipe refugia (Fig 2)were installed in each burned and unburned stand Y-shaped drift fencearrays consisting of four pitfall traps per array were constructed of 30-cm aluminum flashing Each leg of the array was five m with 189 Lbucket pitfall traps located at the end of each leg and one bucket in thecenter (Corn 1994) Four 39 x 122 cm plywood coverboards were ran-domly placed around each array (Fellers and Drost 1994) Three PVCpipes were attached to overstory hardwood trees around each array at 213m from the ground to sample tree frogs (Hylidae) (Boughton et al 2000)Each trap cluster was within a 004-ha area centered around the pitfallarray (Fig 2) Trap clusters were monitored for 29 days between 11 Julyand seven October 2001 for a total of 348 array nights where one arraynight was considered one open trap cluster for one night Traps werechecked every 2448 hrs Each animal captured was identified to speciestoe clipped weighed (g) measured (snout-vent length cm) and released

Habitat variables were measured within a 004-ha plot centered oneach array (n = 12) Species and diameter at breast height (DBH) were

Figure 2 Arrangement of trap clusters used to sample herpetofauna in burned inunburned bottomland hardwood stands at Di-Lane Plantation Wildlife Manage-ment Area Burke County Georgia JulyOctober 2001

KR Moseley SB Castleberry and SH Schweitzer2003 479

recorded for all trees ge 72 cm DBH Basal area was calculated for pinehardwood and snag components Midstory stems (lt 72 cm DBH andge155 m tall) were tallied by species Diameter and length of CWDgreater than 72 cm mean diameter were recorded to determine volumeof CWD All CWD measured was categorized based on degree ofdecomposition class one included logs with a hard outer covering andno soft parts class two included logs with hard or soft outer parts andclass three included logs with completely soft outer parts (Maidens et al1998) Canopy and midstory height (m) were determined using a cli-nometer One canopy and one midstory tree representative of the aver-age height in each plot were measured (m)

Within each 004-ha plot three transect lines 120o apart wereestablished beginning at the center of each array and extending out113 m (Fig 2) Two 1-m2 plots were randomly located along eachtransect line Within each plot understory stems (0155 m) werecounted and percentage of ground cover was estimated Understorystems were placed into one of five categories grass shrub vinewoody and other A stem representing average understory height (cm)within each 1-m2 plot was measured Percent ground cover of pinelitter hardwood litter bare ground and woody debris was estimatedAt three random points along each transect litter depth was measuredto the nearest mm by inserting a ruler into the litter layer Soil pH wasdetermined by taking a mixture of six random soil samples from withinthe 004-ha plot Soil samples were analyzed by the Georgia Coopera-tive Extension Service Athens Distance of array to habitat edge andnearest body of water was determined using ARCVIEW (Environmen-tal Systems Research Institute 1999)

Captured species were pooled into the following taxonomic catego-ries for analysis total herpetofauna amphibians salamanders anuransand reptiles Reptilian taxa were not examined separately because oflow sample sizes Abundance was expressed as the total number ofcaptures per trap cluster and diversity was calculated using the Shan-non-Weiner index (Pielou 1977) Recaptures were not included in abun-dance or diversity calculations A one-way analysis of variance(ANOVA) was used to test taxonomic categories for differences inabundance diversity and richness between treatments Because a par-ticular stand received only a single treatment (burn or no burn) theeffect of stand was nested within treatments and used as the error termwhen testing for treatment effects Habitat variables and distance ofarray to water and habitat edge also were analyzed as a one-wayANOVA with nested effects as described above Habitat variables thatwere not normally distributed were log10 transformed

Spearmans rank correlation was used to examine relationships be-tween habitat variables and abundance of individual species with gt 20

Southeastern Naturalist Vol 2 No 4480

total captures as well as abundance richness and diversity of taxo-nomic categories Only habitat variables that differed between burnedand unburned stands using ANOVA were examined in correlationanalysis All statistical analyses were conducted using SAS statisticalsoftware (SAS Institute 1997)

RESULTS

A total of 346 individuals representing 21 species was captured in348 array nights (Table 1) Of those captured 33 (eight species) werereptiles and 313 (13 species) were amphibians Seven species wererepresented by a single capture Over the trapping period 33 individualswere recaptured

Overall herpetofaunal abundance diversity and richness were notdifferent (P gt 005) between burned and unburned stands (Table 2) Nodifferences (P gt 005) were observed in amphibian salamander oranuran abundance diversity or richness between burned and unburnedstands (Table 2) Reptile abundance (P = 00080) and diversity (P =00462) were greater in burned than unburned stands However reptilespecies richness was not different (P = 04216) between burned andunburned stands (Table 2)

Of habitat variables examined litter depth (P = 00242) and canopyheight (P = 00247) were greater in unburned than in burned stands

Table 1 Numbers of herpetofauna species captured in burned and unburned bottomlandhardwood stands at Di-Lane Plantation Wildlife Management Area Burke County Geor-gia JulyOctober 2001

Species Common Name Burned Unburned

Gastrophryne carolinensis Holbrook Eastern Narrowmouth Toad 42 76Bufo terrestris Bonnaterre Southern Toad 41 19Scaphiopus holbrookii Harlan Eastern Spadefoot 11 29Eumeces fasciatus Linnaeus Five-lined Skink 11 5Bufo americanus Holbrook American Toad 7 5Ambystoma opacum Gravenhorst Marbled Salamander 6 22Hyla versicolor LeConte Gray Treefrog 6 2Plethodon glutinosus Green Slimy Salamander 5 13Rana utricularia Schreber Southern Leopard Frog 4 13Scincella lateralis Say Ground Skink 4 1Rana catesbeiana Shaw Bullfrog 3 4Notophthalmus viridescens Rafinesque Red-spotted Newt 1 0Rana clamitans clamitans Latreille Bronze Frog 1 0Pseudacris ornata Holbrook Ornate Chorus Frog 1 1Eumeces laticeps Schneider Broadhead Skink 1 0Sceloporus undulatus Bosc and Daudin Fence Lizard 1 0Tantilla coronata Baird and Girard Southeastern Crowned Snake 1 0Elaphe obsoleta obsoleta Say Black Rat Snake 1 0

Diadophis punctatus punctatus Linnaeus Southern Ringneck Snake 1 1Hyla cinerea Schneider Green Treefrog 0 1

Anolis carolinensis Voigt Green Anole 0 6

KR Moseley SB Castleberry and SH Schweitzer2003 481

(Table 3) Percent bare ground cover was greater (P = 00029) in burned(2028 ) than unburned (117 ) stands All other habitat variableswere not different (P gt 005) between burned and unburned stands

Significant habitat variables from ANOVA including litter depthcanopy height and percent bare ground cover were examined in correla-tion analyses Abundance of eastern narrowmouth toads (Gastrophrynecarolinensis H) (r = -060 P = 00396) and salamander abundance (r =-062 P = 00303) were negatively correlated with percent bare groundcover No other significant relationships (P gt 005) were observed

DISCUSSION

Our results indicate that periodic winter surface fires in bottomlandhardwood stands of the southeastern Coastal Plain do not affect overallherpetofaunal amphibian salamander or anuran abundance diversityor richness Our findings are similar to those of Ford et al (1999) whofound a high intensity restoration fire in Southern Appalachian hard-wood stands to have a negligible effect on herpetofaunal communitiesbut suggested that alterations in vital microhabitat components due toprescribed fire may negatively impact woodland salamanders On ourstudy site most habitat variables measured were similar between burned

Table 2 Mean (plusmn SD) abundance Shannon-Weiner Index and total number of species foreach taxonomic category (total amphibian salamander anuran and reptile) of herpetofaunacaptured in three burned and three unburned bottomland hardwood stands at Di-LanePlantation Wildlife Management Area Burke County Georgia July-October 2001

Burned Unburned F14

P

Total HerpetofaunaAbundance 4900 plusmn 2346 6600 plusmn 2100 084 04111Shannon-Weiner Index 188 plusmn 026 177 plusmn 014 020 06755Richness 1067 plusmn 306 1033 plusmn 058 013 07344

AmphibiansAbundance 4267 plusmn 2303 6167 plusmn 2079 112 03486Shannon-Weiner Index 148 plusmn 032 154 plusmn 011 012 07466Richness 767 plusmn 208 800 plusmn 0 002 09019

SalamandersAbundance 200 plusmn 068 583 plusmn 114 622 00671Shannon-Weiner Index 021 plusmn 006 035 plusmn 004 649 00635Richness 117 plusmn 031 150 plusmn 022 057 04918

AnuransAbundance 3867 plusmn 2108 5000 plusmn 1609 055 05003Shannon-Weiner Index 124 plusmn 035 117 plusmn 018 011 07581Richness 600 plusmn 173 633 plusmn 058 002 08944

ReptilesAbundance 667 plusmn 058 433 plusmn 058 245 00080a

Shannon-Weiner Index 040 plusmn 006 023 plusmn 005 814 00462a

Richness 300 plusmn 100 233 plusmn 058 080 04216aMeans are significantly different (P le 005)

Southeastern Naturalist Vol 2 No 4482

and unburned stands probably due to the low intensity of burns Of thethree habitat variables that were different between treatments the mostprofound was reduction in litter layer depth which subsequently pro-duced an increase in bare ground cover in burned stands

Decrease in leaf litter depth results in a reduction of on-site mois-ture which can impair dermal respiratory ability in amphibians(Duellman and Trueb 1994) and has been cited as a major factor inreduction of amphibian abundance (Ash 1995 Ash 1997) We capturedalmost twice as many eastern narrowmouth toads in unburned standscompared to burned stands (76 and 42 respectively) and found that

Table 3 Mean (plusmn SD) of habitat variables measured within a 004-ha plot around trapclusters in three burned and three unburned bottomland hardwood stands at Di-LanePlantation Wildlife Management Area Burke County Georgia JulyOctober 2001

Habitat Variable Burned Unburned F14 P

Basal area m2ha (gt 716 cm DBH)Hardwood 2408 plusmn 316 2154 plusmn 343 002 09060

Pine 659 plusmn 221 763 plusmn 343 005 08364Snaga 093 plusmn 040 116 plusmn 058 046 05369Totala 3159 plusmn 446 3032 plusmn 25 026 06369

Midstory stemsha (lt 716 cm DBH)Total stems 134167 plusmn 41966 237083 plusmn 21461 382 01223

Course Woody Debris (m3 ha)

Class 1a 2901 plusmn 1629 2359 plusmn 1437 009 07787Class 2a 588 plusmn 406 4113 plusmn 382 260 01820Class 3a 2507 plusmn 873 6371 plusmn 409 008 07871Totala 5995 plusmn 197 12843 plusmn 7272 002 08921

Canopy height (m) 2114 plusmn 122 2431 plusmn 118 1230 00247b

Midstory height (m) 1339 plusmn 083 1212 plusmn 079 088 04023

Understory (0-155m) stems6m2

Grassa 3700 plusmn 3192 117 plusmn117 108 03582Shruba 7533 plusmn 3357 26 plusmn 1842 505 00880Vine 3033 plusmn 1052 3783 plusmn 1034 020 06785Woody 2583 plusmn 531 3117 plusmn 88 040 05617Other 117 plusmn 075 15 plusmn 096 004 08554Total stems 16967 plusmn 4242 9767 plusmn 1737 433 01059

Understory height (cm) 2641 plusmn 470 259 plusmn 48 001 09294

Percent ground coverPine litter 2436 plusmn 760 1986 plusmn 1027 007 08067Hardwood litter 4917 plusmn 970 7703 plusmn 996 217 02148Bare grounda 2028 plusmn 537 117 plusmn 094 4239 00029b

Woody debrisa 571 plusmn 242 194 plusmn 09 195 02349

Litter depth (cm) 093 plusmn 023 286 plusmn 034 1247 00242b

pH 433 plusmn 011 445 plusmn 016 040 05591

Distance to edge (m) 8650 plusmn 993 1315 plusmn 2155 331 0143

Distance to water (m) 74017 plusmn 18026 122617 plusmn 15381 172 02594aVariable was log10 transformedbMeans are significantly different (P le 005)

KR Moseley SB Castleberry and SH Schweitzer2003 483

abundance was negatively correlated with percent bare ground coverSalamander abundance also was negatively correlated with percent bareground cover Although we did not test for differences of individualspecies between treatments because of low sample sizes our resultssuggest that increases in bare ground cover as a result of leaf litterreductions may be associated with lower abundance of some amphib-ians Other studies have demonstrated similar results where reduction inleaf litter depth was a result of forest harvesting (Ash 1995 Ash 1997Grialou et al 2000 McLeod and Gates 1998 Pough et al 1987) Ourstudy apparently is the first to find such a relationship in bottomlandhardwoods as a result of prescribed fire

Several amphibian species captured possess behavioral characteris-tics important in avoiding desiccation Southern toads (Bufo terrestrisB) eastern spadefoot toads (Scaphiopus holbrooki H) and Americantoads (Bufo americanus Holbrook) together representing 461 and286 of amphibian captures in burned and unburned plots respec-tively are capable of burrowing into moist soil to avoid dry surfaceconditions (Conant and Collins 1998) The ability of these species toescape such conditions by burrowing may enable them to survive instands with a reduced litter layer

Total CWD volume and volume of all CWD decay classes did not differbetween burned and unburned stands Other studies have found decreasesin CWD and consequent decreases in amphibian abundance in areassubjected to prescribed fire (McLeod and Gates 1998) All species cap-tured in our study possess some dependence on CWD (Conant and Collins1998 Harmon et al 1986 Maidens et al 1998 Whiles and Grubaugh1996 Wilson 1995) For example eastern narrowmouth toads represent-ing 328 and 41 of amphibians captured in burned and unburnedstands respectively typically occupy habitats that provide moisture andshelter such as under CWD (Conant and Collins 1998) CWD has arelatively constant moisture regime and can remain moist during long dryperiods providing refuge for amphibians (Boddy 1983) Furthermorearthropods are abundant in and around CWD (Graham 1935 Hanula 1993Harmon et al 1986 Savely 1939) yielding an important food source forinsectivorous herpetofauna These characteristics of CWD may havecompensated to some degree for leaf litter reduction by providing adequateamounts of moisture and food for amphibians

Similar to our results previous studies examining fire effects onherpetofauna in southeastern Coastal Plain pine communities also foundreptiles to increase in diversity and abundance following application ofprescribed fire (Means and Campbell 1981 Mushinsky 1985) attributingthe increases to greater thermoregulatory opportunity within burnedstands Increased bare ground cover in burned stands resulted in a largerground surface area for thermoregulatory activity by reptiles When solar

Southeastern Naturalist Vol 2 No 4484

radiation declines in the evening lizards will leave their perches to flattenout against the warm sand to maintain optimal temperature (Pough et al2001) Many species of reptiles also utilize CWD as a basking substrate aswell as for shelter and nesting (Conant and Collins 1998 Pough et al 2001Wilson 1995) Numerous lizards were observed basking on logs at the edgeof burned stands during our study Five-lined skinks (Eumeces fasciatusL) representing 579 and 385 of reptile species captured in burnedand unburned stands respectively frequently utilize CWD as nesting sitesbecause of the relatively constant microclimate provided for egg develop-ment (Hecnar 1994) We recognize that the observed increase in reptilediversity and abundance in our study may be an underestimate because ourtrapping methods are relatively ineffective at catching large snakes whichwere likely underrepresented in our collections

We conclude that the use of low intensity prescribed fire had littleoverall effect on herpetofauna in our bottomland hardwood forest studyarea However reductions in leaf litter and associated increases in bareground cover may be associated with lower abundance of some amphib-ians Conversely the increased ground surface area available for ther-moregulatory activity may benefit some reptiles CWD was relativelyunaffected by the burning regime probably because of the low intensityof the fires In areas where fire intensity is greater the impacts on theherpetofaunal community may be more pronounced

ACKNOWLEDGMENTS

We thank V VanSant H Barnhill and J Bearden of the Georgia Depart-ment of Natural Resources Wildlife Resources Division for their assistance andaccess to Di-Lane WMA We also thank K Francl for GIS assistance

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Ash AN 1995 Effects of clear-cutting on litter parameters in the SouthernBlue Ridge mountains Castanea 60(2)8997

Ash AN 1997 Disappearance and return of Plethodontid salamanders toclearcut plots in the southern Blue Ridge mountains Conservation Biology11(4)983989

Bennett SH JW Gibbons and J Glanville 1980 Terrestrial activity abun-dance and diversity of amphibians in differently managed forest typesAmerican Midland Naturalist 103(2)412416

Boddy L 1983 Microclimate and moisture dynamics of wood decomposing interrestrial ecosystems Soil Biology and Biochemistry 15(2)149157

Boughton RG J Staiger and R Franz 2000 Use of PVC refugia as asampling technique for hylid treefrogs American Midland Naturalist144(1)168177

Conant R and JT Collins 1998 A Field Guide to Reptiles and Amphibians ofEasternCentral North America 3rd Edition Houghton Mifflin Co BostonMA 616 pp

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Corn PS 1994 Straight line drift fences and pitfall traps Pp 109117 InWR Heyer MA Donnelly RW McDiarmid LC Hayek and MSFoster (Eds) Measuring and Monitoring Biological Diversity StandardMethods for Amphibians Smithsonian Institute Press Washington DC364 pp

DeMaynadier PG and ML Hunter Jr 1995 The relationship between forestmanagement and amphibian ecology A review of the North Americanliterature Environmental Reviews 3(3-4)230261

Duellman WE and L Trueb 1994 Biology of Amphibians The JohnsHopkins University Press Baltimore MD 670 pp

Environmental Systems Research Institute 1999 ARCVIEW Version 32Environmental Systems Research Institute Redlands CA

Fellers GM and CA Drost 1994 Sampling with artificial cover Pp 146-150 In WR Heyer MA Donnelly RW McDiarmid LC Hayek andMS Foster (Eds) Measuring and Monitoring Biological Diversity Stan-dard Methods for Amphibians Smithsonian Institute Press WashingtonDC 364 pp

Ford WM MA Menzel DW McGill J Laerm and TS McCay 1999Effects of a community restoration fire on small mammals and herpetofaunain the Southern Appalachians Forest Ecology and Management 114(2-3)233243

Graham SA 1935 The felled tree trunk as an ecological unit Ecology6(4)397412

Greenberg CH DG Neary and LD Harris 1994 Effect of high-intensitywildfire and silvicultural treatments on reptile communities in sand-pinescrub Conservation Biology 8(4)10471057

Grialou JA SD West and RN Wilkins 2000 The effects of forest clearcutharvesting and thinning on terrestrial salamanders Journal of Wildlife Man-agement 64(1)105113

Hanula JL 1993 Relationship of wood-feeding insects and coarse woodydebris Pp 5565 In JW McMinn and DA Crossley Jr (Eds)Biodiversity of Coarse Woody Debris in Southern Forests Effects onBiodiversity Proceedings of the Workshop on Coarse Woody Debris inSouthern Forests US Department of Agriculture Forest Service GeneralTechnical Report SE-94 146 pp

Harmon ME JF Franklin FJ Swanson P Sollins SV Gregory JDLattin NH Anderson SP Cline NG Aumen JR Sedell GWLienkaemper K Cromack Jr KW Cummins 1986 Ecology of coarsewoody debris in temperate ecosystems Advances in Ecological Research15133302

Hecnar SJ 1994 Nest distribution site selection and brooding in the five-linedskink (Eumeces fasciatus) Canadian Journal of Zoology 72(8)15101516

Maidens DA MA Menzel and J Laerm 1998 Notes on the effect of sizeand level of decay of coarse woody debris on relative abundance of shrewsand salamanders in the southern Appalachian mountains Georgia Journal ofScience 56(4)226-233

McLeod RF and JE Gates 1998 Response of herpetofaunal communities toforest cutting and burning at Chesapeake Farms Maryland American Mid-land Naturalist 139(1)164177

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Means DB and HW Campbell 1981 Effects of prescribed burningonamphibians and reptiles Pp 8996 In GW Wood (Ed) Prescribed Fireand Wildlife in Southern Forests Belle Baruch Forest Science InstituteClemson University Georgetown SC 170 pp

Mushinsky HR 1985 Fire and the Florida sandhill herptofaunal communitywith special attention to responses of Cnemidophorus sexlineatusHerpetologica 41(3)333342

Nyland RD 1996 Silviculture Concepts and Applications McGraw-HillNew York NY 633 pp

Pielou EC 1977 Mathematical Ecology John Wiley and Sons New YorkNY 385 pp

Pough FH RM Andrews JE Cadle ML Crump AH Savitzky and KDWells 2001 Herpetology 2ndEdition Prentice Hall NJ 612 pp

Russell KR DH Van Lear and DC Guynn Jr 1999 Prescribed fire effectson herpetofauna Review and management implications Wildlife SocietyBulletin 27(2)374384

Savely HE 1939 Ecological relations of certain animals in dead pine and oaklogs Ecological Monographs 9(3)322377

SAS Institute 1997 SASSTAT software Changes and enhancements throughrelease 612 SAS Institute Cary NC

Smith DM BC Larson MJ Kelty and PMS Ashton 1997 The Practice ofSilviculture Applied Forest Ecology Wiley and Sons New York NY 537 pp

Spotila JR 1972 Role of temperature and water in the ecology of lunglesssalamanders Ecological Monographs 42(1)95125

Stoddard HL 1962 Use of fire in pine forests and game lands of the deepSoutheast Proceedings of the Tall Timbers Fire Ecology Conference TallTimbers Research Station Tallahassee FL 13142

Van Lear DH and TA Waldrop 1989 History uses and effects of fire in theAppalachians US Department of Agriculture Forest Service GeneralTechnical Report SE-54 20 pp

Walker LC 1999 The North American Forests Geography Ecology andSilviculture CRC Press Boca Raton FL 398 pp

Whiles MR and JW Grubaugh 1996 Importance of coarse woody debris tosouthern forest herpetofauna Pp 94100 In JW McMinn and DACrossley Jr (Eds) Biodiversity of Coarse Woody Debris in SouthernForests Effects on Biodiversity Proceedings of the Workshop on CoarseWoody Debris in Southern Forests US Department of Agriculture ForestService General Technical Report SE-94 146 pp

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KR Moseley SB Castleberry and SH Schweitzer2003 477

nigra L) laurel oak (Q laurifolia Michx) and sweetgum (Liquidam-bar styraciflua L) Midstory tree species included water oak laurel oaksweetgum and black gum (Nyssa sylvatica Marsh) The understoryconsisted of blueberry (Vaccinium spp) greenbrier (Smilax spp)grapevine (Vitis spp) American beautyberry (Callicarpa americana L)and other seedling hardwood species

In an effort to improve northern bobwhite (Colinus virginianus L)habitat on Di-Lane pine and hardwood stands were burned to increasecanopy openings and stimulate herbaceous growth Stands were burnedduring winter using aerial ignition Burns in bottomland hardwood standswere light due to moist soil and litter conditions and numerous ephemeralponds present during winter Treatment stands considered for the studywere those burned every two to three years for the past nine years withthe most recent burn occurring in January 2001 (J Bearden GeorgiaDepartment of Natural Resources pers comm June 2001) Three standswere randomly chosen from all stands meeting the criteria (Fig 1)Treatment stand sizes were 364 ha 445 ha and 931 ha Three standswith similar plant species composition hydrology and elevation to treat-ment stands but which had not been burned in at least 9 years wererandomly selected as reference stands All stands contained numerouspools of standing water from late fall 2000 through early spring 2001

Figure 1 Di-Lane Wildlife Management Area Burke County Georgia andthe locations of trap clusters in burned and unburned bottomland hardwoodstands

Southeastern Naturalist Vol 2 No 4478

METHODS

Two trap clusters each consisting of a drift fence pitfall array ply-wood coverboards and polyvinyl chloride (PVC) pipe refugia (Fig 2)were installed in each burned and unburned stand Y-shaped drift fencearrays consisting of four pitfall traps per array were constructed of 30-cm aluminum flashing Each leg of the array was five m with 189 Lbucket pitfall traps located at the end of each leg and one bucket in thecenter (Corn 1994) Four 39 x 122 cm plywood coverboards were ran-domly placed around each array (Fellers and Drost 1994) Three PVCpipes were attached to overstory hardwood trees around each array at 213m from the ground to sample tree frogs (Hylidae) (Boughton et al 2000)Each trap cluster was within a 004-ha area centered around the pitfallarray (Fig 2) Trap clusters were monitored for 29 days between 11 Julyand seven October 2001 for a total of 348 array nights where one arraynight was considered one open trap cluster for one night Traps werechecked every 2448 hrs Each animal captured was identified to speciestoe clipped weighed (g) measured (snout-vent length cm) and released

Habitat variables were measured within a 004-ha plot centered oneach array (n = 12) Species and diameter at breast height (DBH) were

Figure 2 Arrangement of trap clusters used to sample herpetofauna in burned inunburned bottomland hardwood stands at Di-Lane Plantation Wildlife Manage-ment Area Burke County Georgia JulyOctober 2001

KR Moseley SB Castleberry and SH Schweitzer2003 479

recorded for all trees ge 72 cm DBH Basal area was calculated for pinehardwood and snag components Midstory stems (lt 72 cm DBH andge155 m tall) were tallied by species Diameter and length of CWDgreater than 72 cm mean diameter were recorded to determine volumeof CWD All CWD measured was categorized based on degree ofdecomposition class one included logs with a hard outer covering andno soft parts class two included logs with hard or soft outer parts andclass three included logs with completely soft outer parts (Maidens et al1998) Canopy and midstory height (m) were determined using a cli-nometer One canopy and one midstory tree representative of the aver-age height in each plot were measured (m)

Within each 004-ha plot three transect lines 120o apart wereestablished beginning at the center of each array and extending out113 m (Fig 2) Two 1-m2 plots were randomly located along eachtransect line Within each plot understory stems (0155 m) werecounted and percentage of ground cover was estimated Understorystems were placed into one of five categories grass shrub vinewoody and other A stem representing average understory height (cm)within each 1-m2 plot was measured Percent ground cover of pinelitter hardwood litter bare ground and woody debris was estimatedAt three random points along each transect litter depth was measuredto the nearest mm by inserting a ruler into the litter layer Soil pH wasdetermined by taking a mixture of six random soil samples from withinthe 004-ha plot Soil samples were analyzed by the Georgia Coopera-tive Extension Service Athens Distance of array to habitat edge andnearest body of water was determined using ARCVIEW (Environmen-tal Systems Research Institute 1999)

Captured species were pooled into the following taxonomic catego-ries for analysis total herpetofauna amphibians salamanders anuransand reptiles Reptilian taxa were not examined separately because oflow sample sizes Abundance was expressed as the total number ofcaptures per trap cluster and diversity was calculated using the Shan-non-Weiner index (Pielou 1977) Recaptures were not included in abun-dance or diversity calculations A one-way analysis of variance(ANOVA) was used to test taxonomic categories for differences inabundance diversity and richness between treatments Because a par-ticular stand received only a single treatment (burn or no burn) theeffect of stand was nested within treatments and used as the error termwhen testing for treatment effects Habitat variables and distance ofarray to water and habitat edge also were analyzed as a one-wayANOVA with nested effects as described above Habitat variables thatwere not normally distributed were log10 transformed

Spearmans rank correlation was used to examine relationships be-tween habitat variables and abundance of individual species with gt 20

Southeastern Naturalist Vol 2 No 4480

total captures as well as abundance richness and diversity of taxo-nomic categories Only habitat variables that differed between burnedand unburned stands using ANOVA were examined in correlationanalysis All statistical analyses were conducted using SAS statisticalsoftware (SAS Institute 1997)

RESULTS

A total of 346 individuals representing 21 species was captured in348 array nights (Table 1) Of those captured 33 (eight species) werereptiles and 313 (13 species) were amphibians Seven species wererepresented by a single capture Over the trapping period 33 individualswere recaptured

Overall herpetofaunal abundance diversity and richness were notdifferent (P gt 005) between burned and unburned stands (Table 2) Nodifferences (P gt 005) were observed in amphibian salamander oranuran abundance diversity or richness between burned and unburnedstands (Table 2) Reptile abundance (P = 00080) and diversity (P =00462) were greater in burned than unburned stands However reptilespecies richness was not different (P = 04216) between burned andunburned stands (Table 2)

Of habitat variables examined litter depth (P = 00242) and canopyheight (P = 00247) were greater in unburned than in burned stands

Table 1 Numbers of herpetofauna species captured in burned and unburned bottomlandhardwood stands at Di-Lane Plantation Wildlife Management Area Burke County Geor-gia JulyOctober 2001

Species Common Name Burned Unburned

Gastrophryne carolinensis Holbrook Eastern Narrowmouth Toad 42 76Bufo terrestris Bonnaterre Southern Toad 41 19Scaphiopus holbrookii Harlan Eastern Spadefoot 11 29Eumeces fasciatus Linnaeus Five-lined Skink 11 5Bufo americanus Holbrook American Toad 7 5Ambystoma opacum Gravenhorst Marbled Salamander 6 22Hyla versicolor LeConte Gray Treefrog 6 2Plethodon glutinosus Green Slimy Salamander 5 13Rana utricularia Schreber Southern Leopard Frog 4 13Scincella lateralis Say Ground Skink 4 1Rana catesbeiana Shaw Bullfrog 3 4Notophthalmus viridescens Rafinesque Red-spotted Newt 1 0Rana clamitans clamitans Latreille Bronze Frog 1 0Pseudacris ornata Holbrook Ornate Chorus Frog 1 1Eumeces laticeps Schneider Broadhead Skink 1 0Sceloporus undulatus Bosc and Daudin Fence Lizard 1 0Tantilla coronata Baird and Girard Southeastern Crowned Snake 1 0Elaphe obsoleta obsoleta Say Black Rat Snake 1 0

Diadophis punctatus punctatus Linnaeus Southern Ringneck Snake 1 1Hyla cinerea Schneider Green Treefrog 0 1

Anolis carolinensis Voigt Green Anole 0 6

KR Moseley SB Castleberry and SH Schweitzer2003 481

(Table 3) Percent bare ground cover was greater (P = 00029) in burned(2028 ) than unburned (117 ) stands All other habitat variableswere not different (P gt 005) between burned and unburned stands

Significant habitat variables from ANOVA including litter depthcanopy height and percent bare ground cover were examined in correla-tion analyses Abundance of eastern narrowmouth toads (Gastrophrynecarolinensis H) (r = -060 P = 00396) and salamander abundance (r =-062 P = 00303) were negatively correlated with percent bare groundcover No other significant relationships (P gt 005) were observed

DISCUSSION

Our results indicate that periodic winter surface fires in bottomlandhardwood stands of the southeastern Coastal Plain do not affect overallherpetofaunal amphibian salamander or anuran abundance diversityor richness Our findings are similar to those of Ford et al (1999) whofound a high intensity restoration fire in Southern Appalachian hard-wood stands to have a negligible effect on herpetofaunal communitiesbut suggested that alterations in vital microhabitat components due toprescribed fire may negatively impact woodland salamanders On ourstudy site most habitat variables measured were similar between burned

Table 2 Mean (plusmn SD) abundance Shannon-Weiner Index and total number of species foreach taxonomic category (total amphibian salamander anuran and reptile) of herpetofaunacaptured in three burned and three unburned bottomland hardwood stands at Di-LanePlantation Wildlife Management Area Burke County Georgia July-October 2001

Burned Unburned F14

P

Total HerpetofaunaAbundance 4900 plusmn 2346 6600 plusmn 2100 084 04111Shannon-Weiner Index 188 plusmn 026 177 plusmn 014 020 06755Richness 1067 plusmn 306 1033 plusmn 058 013 07344

AmphibiansAbundance 4267 plusmn 2303 6167 plusmn 2079 112 03486Shannon-Weiner Index 148 plusmn 032 154 plusmn 011 012 07466Richness 767 plusmn 208 800 plusmn 0 002 09019

SalamandersAbundance 200 plusmn 068 583 plusmn 114 622 00671Shannon-Weiner Index 021 plusmn 006 035 plusmn 004 649 00635Richness 117 plusmn 031 150 plusmn 022 057 04918

AnuransAbundance 3867 plusmn 2108 5000 plusmn 1609 055 05003Shannon-Weiner Index 124 plusmn 035 117 plusmn 018 011 07581Richness 600 plusmn 173 633 plusmn 058 002 08944

ReptilesAbundance 667 plusmn 058 433 plusmn 058 245 00080a

Shannon-Weiner Index 040 plusmn 006 023 plusmn 005 814 00462a

Richness 300 plusmn 100 233 plusmn 058 080 04216aMeans are significantly different (P le 005)

Southeastern Naturalist Vol 2 No 4482

and unburned stands probably due to the low intensity of burns Of thethree habitat variables that were different between treatments the mostprofound was reduction in litter layer depth which subsequently pro-duced an increase in bare ground cover in burned stands

Decrease in leaf litter depth results in a reduction of on-site mois-ture which can impair dermal respiratory ability in amphibians(Duellman and Trueb 1994) and has been cited as a major factor inreduction of amphibian abundance (Ash 1995 Ash 1997) We capturedalmost twice as many eastern narrowmouth toads in unburned standscompared to burned stands (76 and 42 respectively) and found that

Table 3 Mean (plusmn SD) of habitat variables measured within a 004-ha plot around trapclusters in three burned and three unburned bottomland hardwood stands at Di-LanePlantation Wildlife Management Area Burke County Georgia JulyOctober 2001

Habitat Variable Burned Unburned F14 P

Basal area m2ha (gt 716 cm DBH)Hardwood 2408 plusmn 316 2154 plusmn 343 002 09060

Pine 659 plusmn 221 763 plusmn 343 005 08364Snaga 093 plusmn 040 116 plusmn 058 046 05369Totala 3159 plusmn 446 3032 plusmn 25 026 06369

Midstory stemsha (lt 716 cm DBH)Total stems 134167 plusmn 41966 237083 plusmn 21461 382 01223

Course Woody Debris (m3 ha)

Class 1a 2901 plusmn 1629 2359 plusmn 1437 009 07787Class 2a 588 plusmn 406 4113 plusmn 382 260 01820Class 3a 2507 plusmn 873 6371 plusmn 409 008 07871Totala 5995 plusmn 197 12843 plusmn 7272 002 08921

Canopy height (m) 2114 plusmn 122 2431 plusmn 118 1230 00247b

Midstory height (m) 1339 plusmn 083 1212 plusmn 079 088 04023

Understory (0-155m) stems6m2

Grassa 3700 plusmn 3192 117 plusmn117 108 03582Shruba 7533 plusmn 3357 26 plusmn 1842 505 00880Vine 3033 plusmn 1052 3783 plusmn 1034 020 06785Woody 2583 plusmn 531 3117 plusmn 88 040 05617Other 117 plusmn 075 15 plusmn 096 004 08554Total stems 16967 plusmn 4242 9767 plusmn 1737 433 01059

Understory height (cm) 2641 plusmn 470 259 plusmn 48 001 09294

Percent ground coverPine litter 2436 plusmn 760 1986 plusmn 1027 007 08067Hardwood litter 4917 plusmn 970 7703 plusmn 996 217 02148Bare grounda 2028 plusmn 537 117 plusmn 094 4239 00029b

Woody debrisa 571 plusmn 242 194 plusmn 09 195 02349

Litter depth (cm) 093 plusmn 023 286 plusmn 034 1247 00242b

pH 433 plusmn 011 445 plusmn 016 040 05591

Distance to edge (m) 8650 plusmn 993 1315 plusmn 2155 331 0143

Distance to water (m) 74017 plusmn 18026 122617 plusmn 15381 172 02594aVariable was log10 transformedbMeans are significantly different (P le 005)

KR Moseley SB Castleberry and SH Schweitzer2003 483

abundance was negatively correlated with percent bare ground coverSalamander abundance also was negatively correlated with percent bareground cover Although we did not test for differences of individualspecies between treatments because of low sample sizes our resultssuggest that increases in bare ground cover as a result of leaf litterreductions may be associated with lower abundance of some amphib-ians Other studies have demonstrated similar results where reduction inleaf litter depth was a result of forest harvesting (Ash 1995 Ash 1997Grialou et al 2000 McLeod and Gates 1998 Pough et al 1987) Ourstudy apparently is the first to find such a relationship in bottomlandhardwoods as a result of prescribed fire

Several amphibian species captured possess behavioral characteris-tics important in avoiding desiccation Southern toads (Bufo terrestrisB) eastern spadefoot toads (Scaphiopus holbrooki H) and Americantoads (Bufo americanus Holbrook) together representing 461 and286 of amphibian captures in burned and unburned plots respec-tively are capable of burrowing into moist soil to avoid dry surfaceconditions (Conant and Collins 1998) The ability of these species toescape such conditions by burrowing may enable them to survive instands with a reduced litter layer

Total CWD volume and volume of all CWD decay classes did not differbetween burned and unburned stands Other studies have found decreasesin CWD and consequent decreases in amphibian abundance in areassubjected to prescribed fire (McLeod and Gates 1998) All species cap-tured in our study possess some dependence on CWD (Conant and Collins1998 Harmon et al 1986 Maidens et al 1998 Whiles and Grubaugh1996 Wilson 1995) For example eastern narrowmouth toads represent-ing 328 and 41 of amphibians captured in burned and unburnedstands respectively typically occupy habitats that provide moisture andshelter such as under CWD (Conant and Collins 1998) CWD has arelatively constant moisture regime and can remain moist during long dryperiods providing refuge for amphibians (Boddy 1983) Furthermorearthropods are abundant in and around CWD (Graham 1935 Hanula 1993Harmon et al 1986 Savely 1939) yielding an important food source forinsectivorous herpetofauna These characteristics of CWD may havecompensated to some degree for leaf litter reduction by providing adequateamounts of moisture and food for amphibians

Similar to our results previous studies examining fire effects onherpetofauna in southeastern Coastal Plain pine communities also foundreptiles to increase in diversity and abundance following application ofprescribed fire (Means and Campbell 1981 Mushinsky 1985) attributingthe increases to greater thermoregulatory opportunity within burnedstands Increased bare ground cover in burned stands resulted in a largerground surface area for thermoregulatory activity by reptiles When solar

Southeastern Naturalist Vol 2 No 4484

radiation declines in the evening lizards will leave their perches to flattenout against the warm sand to maintain optimal temperature (Pough et al2001) Many species of reptiles also utilize CWD as a basking substrate aswell as for shelter and nesting (Conant and Collins 1998 Pough et al 2001Wilson 1995) Numerous lizards were observed basking on logs at the edgeof burned stands during our study Five-lined skinks (Eumeces fasciatusL) representing 579 and 385 of reptile species captured in burnedand unburned stands respectively frequently utilize CWD as nesting sitesbecause of the relatively constant microclimate provided for egg develop-ment (Hecnar 1994) We recognize that the observed increase in reptilediversity and abundance in our study may be an underestimate because ourtrapping methods are relatively ineffective at catching large snakes whichwere likely underrepresented in our collections

We conclude that the use of low intensity prescribed fire had littleoverall effect on herpetofauna in our bottomland hardwood forest studyarea However reductions in leaf litter and associated increases in bareground cover may be associated with lower abundance of some amphib-ians Conversely the increased ground surface area available for ther-moregulatory activity may benefit some reptiles CWD was relativelyunaffected by the burning regime probably because of the low intensityof the fires In areas where fire intensity is greater the impacts on theherpetofaunal community may be more pronounced

ACKNOWLEDGMENTS

We thank V VanSant H Barnhill and J Bearden of the Georgia Depart-ment of Natural Resources Wildlife Resources Division for their assistance andaccess to Di-Lane WMA We also thank K Francl for GIS assistance

LITERATURE CITED

Ash AN 1995 Effects of clear-cutting on litter parameters in the SouthernBlue Ridge mountains Castanea 60(2)8997

Ash AN 1997 Disappearance and return of Plethodontid salamanders toclearcut plots in the southern Blue Ridge mountains Conservation Biology11(4)983989

Bennett SH JW Gibbons and J Glanville 1980 Terrestrial activity abun-dance and diversity of amphibians in differently managed forest typesAmerican Midland Naturalist 103(2)412416

Boddy L 1983 Microclimate and moisture dynamics of wood decomposing interrestrial ecosystems Soil Biology and Biochemistry 15(2)149157

Boughton RG J Staiger and R Franz 2000 Use of PVC refugia as asampling technique for hylid treefrogs American Midland Naturalist144(1)168177

Conant R and JT Collins 1998 A Field Guide to Reptiles and Amphibians ofEasternCentral North America 3rd Edition Houghton Mifflin Co BostonMA 616 pp

KR Moseley SB Castleberry and SH Schweitzer2003 485

Corn PS 1994 Straight line drift fences and pitfall traps Pp 109117 InWR Heyer MA Donnelly RW McDiarmid LC Hayek and MSFoster (Eds) Measuring and Monitoring Biological Diversity StandardMethods for Amphibians Smithsonian Institute Press Washington DC364 pp

DeMaynadier PG and ML Hunter Jr 1995 The relationship between forestmanagement and amphibian ecology A review of the North Americanliterature Environmental Reviews 3(3-4)230261

Duellman WE and L Trueb 1994 Biology of Amphibians The JohnsHopkins University Press Baltimore MD 670 pp

Environmental Systems Research Institute 1999 ARCVIEW Version 32Environmental Systems Research Institute Redlands CA

Fellers GM and CA Drost 1994 Sampling with artificial cover Pp 146-150 In WR Heyer MA Donnelly RW McDiarmid LC Hayek andMS Foster (Eds) Measuring and Monitoring Biological Diversity Stan-dard Methods for Amphibians Smithsonian Institute Press WashingtonDC 364 pp

Ford WM MA Menzel DW McGill J Laerm and TS McCay 1999Effects of a community restoration fire on small mammals and herpetofaunain the Southern Appalachians Forest Ecology and Management 114(2-3)233243

Graham SA 1935 The felled tree trunk as an ecological unit Ecology6(4)397412

Greenberg CH DG Neary and LD Harris 1994 Effect of high-intensitywildfire and silvicultural treatments on reptile communities in sand-pinescrub Conservation Biology 8(4)10471057

Grialou JA SD West and RN Wilkins 2000 The effects of forest clearcutharvesting and thinning on terrestrial salamanders Journal of Wildlife Man-agement 64(1)105113

Hanula JL 1993 Relationship of wood-feeding insects and coarse woodydebris Pp 5565 In JW McMinn and DA Crossley Jr (Eds)Biodiversity of Coarse Woody Debris in Southern Forests Effects onBiodiversity Proceedings of the Workshop on Coarse Woody Debris inSouthern Forests US Department of Agriculture Forest Service GeneralTechnical Report SE-94 146 pp

Harmon ME JF Franklin FJ Swanson P Sollins SV Gregory JDLattin NH Anderson SP Cline NG Aumen JR Sedell GWLienkaemper K Cromack Jr KW Cummins 1986 Ecology of coarsewoody debris in temperate ecosystems Advances in Ecological Research15133302

Hecnar SJ 1994 Nest distribution site selection and brooding in the five-linedskink (Eumeces fasciatus) Canadian Journal of Zoology 72(8)15101516

Maidens DA MA Menzel and J Laerm 1998 Notes on the effect of sizeand level of decay of coarse woody debris on relative abundance of shrewsand salamanders in the southern Appalachian mountains Georgia Journal ofScience 56(4)226-233

McLeod RF and JE Gates 1998 Response of herpetofaunal communities toforest cutting and burning at Chesapeake Farms Maryland American Mid-land Naturalist 139(1)164177

Southeastern Naturalist Vol 2 No 4486

Means DB and HW Campbell 1981 Effects of prescribed burningonamphibians and reptiles Pp 8996 In GW Wood (Ed) Prescribed Fireand Wildlife in Southern Forests Belle Baruch Forest Science InstituteClemson University Georgetown SC 170 pp

Mushinsky HR 1985 Fire and the Florida sandhill herptofaunal communitywith special attention to responses of Cnemidophorus sexlineatusHerpetologica 41(3)333342

Nyland RD 1996 Silviculture Concepts and Applications McGraw-HillNew York NY 633 pp

Pielou EC 1977 Mathematical Ecology John Wiley and Sons New YorkNY 385 pp

Pough FH RM Andrews JE Cadle ML Crump AH Savitzky and KDWells 2001 Herpetology 2ndEdition Prentice Hall NJ 612 pp

Russell KR DH Van Lear and DC Guynn Jr 1999 Prescribed fire effectson herpetofauna Review and management implications Wildlife SocietyBulletin 27(2)374384

Savely HE 1939 Ecological relations of certain animals in dead pine and oaklogs Ecological Monographs 9(3)322377

SAS Institute 1997 SASSTAT software Changes and enhancements throughrelease 612 SAS Institute Cary NC

Smith DM BC Larson MJ Kelty and PMS Ashton 1997 The Practice ofSilviculture Applied Forest Ecology Wiley and Sons New York NY 537 pp

Spotila JR 1972 Role of temperature and water in the ecology of lunglesssalamanders Ecological Monographs 42(1)95125

Stoddard HL 1962 Use of fire in pine forests and game lands of the deepSoutheast Proceedings of the Tall Timbers Fire Ecology Conference TallTimbers Research Station Tallahassee FL 13142

Van Lear DH and TA Waldrop 1989 History uses and effects of fire in theAppalachians US Department of Agriculture Forest Service GeneralTechnical Report SE-54 20 pp

Walker LC 1999 The North American Forests Geography Ecology andSilviculture CRC Press Boca Raton FL 398 pp

Whiles MR and JW Grubaugh 1996 Importance of coarse woody debris tosouthern forest herpetofauna Pp 94100 In JW McMinn and DACrossley Jr (Eds) Biodiversity of Coarse Woody Debris in SouthernForests Effects on Biodiversity Proceedings of the Workshop on CoarseWoody Debris in Southern Forests US Department of Agriculture ForestService General Technical Report SE-94 146 pp

Wilson LA 1995 Land Manageriacutes Guide to the Amphibians and Reptiles ofthe SouthThe Nature Conservancy Southeastern Region Chapel Hill NC324 pp

Southeastern Naturalist Vol 2 No 4478

METHODS

Two trap clusters each consisting of a drift fence pitfall array ply-wood coverboards and polyvinyl chloride (PVC) pipe refugia (Fig 2)were installed in each burned and unburned stand Y-shaped drift fencearrays consisting of four pitfall traps per array were constructed of 30-cm aluminum flashing Each leg of the array was five m with 189 Lbucket pitfall traps located at the end of each leg and one bucket in thecenter (Corn 1994) Four 39 x 122 cm plywood coverboards were ran-domly placed around each array (Fellers and Drost 1994) Three PVCpipes were attached to overstory hardwood trees around each array at 213m from the ground to sample tree frogs (Hylidae) (Boughton et al 2000)Each trap cluster was within a 004-ha area centered around the pitfallarray (Fig 2) Trap clusters were monitored for 29 days between 11 Julyand seven October 2001 for a total of 348 array nights where one arraynight was considered one open trap cluster for one night Traps werechecked every 2448 hrs Each animal captured was identified to speciestoe clipped weighed (g) measured (snout-vent length cm) and released

Habitat variables were measured within a 004-ha plot centered oneach array (n = 12) Species and diameter at breast height (DBH) were

Figure 2 Arrangement of trap clusters used to sample herpetofauna in burned inunburned bottomland hardwood stands at Di-Lane Plantation Wildlife Manage-ment Area Burke County Georgia JulyOctober 2001

KR Moseley SB Castleberry and SH Schweitzer2003 479

recorded for all trees ge 72 cm DBH Basal area was calculated for pinehardwood and snag components Midstory stems (lt 72 cm DBH andge155 m tall) were tallied by species Diameter and length of CWDgreater than 72 cm mean diameter were recorded to determine volumeof CWD All CWD measured was categorized based on degree ofdecomposition class one included logs with a hard outer covering andno soft parts class two included logs with hard or soft outer parts andclass three included logs with completely soft outer parts (Maidens et al1998) Canopy and midstory height (m) were determined using a cli-nometer One canopy and one midstory tree representative of the aver-age height in each plot were measured (m)

Within each 004-ha plot three transect lines 120o apart wereestablished beginning at the center of each array and extending out113 m (Fig 2) Two 1-m2 plots were randomly located along eachtransect line Within each plot understory stems (0155 m) werecounted and percentage of ground cover was estimated Understorystems were placed into one of five categories grass shrub vinewoody and other A stem representing average understory height (cm)within each 1-m2 plot was measured Percent ground cover of pinelitter hardwood litter bare ground and woody debris was estimatedAt three random points along each transect litter depth was measuredto the nearest mm by inserting a ruler into the litter layer Soil pH wasdetermined by taking a mixture of six random soil samples from withinthe 004-ha plot Soil samples were analyzed by the Georgia Coopera-tive Extension Service Athens Distance of array to habitat edge andnearest body of water was determined using ARCVIEW (Environmen-tal Systems Research Institute 1999)

Captured species were pooled into the following taxonomic catego-ries for analysis total herpetofauna amphibians salamanders anuransand reptiles Reptilian taxa were not examined separately because oflow sample sizes Abundance was expressed as the total number ofcaptures per trap cluster and diversity was calculated using the Shan-non-Weiner index (Pielou 1977) Recaptures were not included in abun-dance or diversity calculations A one-way analysis of variance(ANOVA) was used to test taxonomic categories for differences inabundance diversity and richness between treatments Because a par-ticular stand received only a single treatment (burn or no burn) theeffect of stand was nested within treatments and used as the error termwhen testing for treatment effects Habitat variables and distance ofarray to water and habitat edge also were analyzed as a one-wayANOVA with nested effects as described above Habitat variables thatwere not normally distributed were log10 transformed

Spearmans rank correlation was used to examine relationships be-tween habitat variables and abundance of individual species with gt 20

Southeastern Naturalist Vol 2 No 4480

total captures as well as abundance richness and diversity of taxo-nomic categories Only habitat variables that differed between burnedand unburned stands using ANOVA were examined in correlationanalysis All statistical analyses were conducted using SAS statisticalsoftware (SAS Institute 1997)

RESULTS

A total of 346 individuals representing 21 species was captured in348 array nights (Table 1) Of those captured 33 (eight species) werereptiles and 313 (13 species) were amphibians Seven species wererepresented by a single capture Over the trapping period 33 individualswere recaptured

Overall herpetofaunal abundance diversity and richness were notdifferent (P gt 005) between burned and unburned stands (Table 2) Nodifferences (P gt 005) were observed in amphibian salamander oranuran abundance diversity or richness between burned and unburnedstands (Table 2) Reptile abundance (P = 00080) and diversity (P =00462) were greater in burned than unburned stands However reptilespecies richness was not different (P = 04216) between burned andunburned stands (Table 2)

Of habitat variables examined litter depth (P = 00242) and canopyheight (P = 00247) were greater in unburned than in burned stands

Table 1 Numbers of herpetofauna species captured in burned and unburned bottomlandhardwood stands at Di-Lane Plantation Wildlife Management Area Burke County Geor-gia JulyOctober 2001

Species Common Name Burned Unburned

Gastrophryne carolinensis Holbrook Eastern Narrowmouth Toad 42 76Bufo terrestris Bonnaterre Southern Toad 41 19Scaphiopus holbrookii Harlan Eastern Spadefoot 11 29Eumeces fasciatus Linnaeus Five-lined Skink 11 5Bufo americanus Holbrook American Toad 7 5Ambystoma opacum Gravenhorst Marbled Salamander 6 22Hyla versicolor LeConte Gray Treefrog 6 2Plethodon glutinosus Green Slimy Salamander 5 13Rana utricularia Schreber Southern Leopard Frog 4 13Scincella lateralis Say Ground Skink 4 1Rana catesbeiana Shaw Bullfrog 3 4Notophthalmus viridescens Rafinesque Red-spotted Newt 1 0Rana clamitans clamitans Latreille Bronze Frog 1 0Pseudacris ornata Holbrook Ornate Chorus Frog 1 1Eumeces laticeps Schneider Broadhead Skink 1 0Sceloporus undulatus Bosc and Daudin Fence Lizard 1 0Tantilla coronata Baird and Girard Southeastern Crowned Snake 1 0Elaphe obsoleta obsoleta Say Black Rat Snake 1 0

Diadophis punctatus punctatus Linnaeus Southern Ringneck Snake 1 1Hyla cinerea Schneider Green Treefrog 0 1

Anolis carolinensis Voigt Green Anole 0 6

KR Moseley SB Castleberry and SH Schweitzer2003 481

(Table 3) Percent bare ground cover was greater (P = 00029) in burned(2028 ) than unburned (117 ) stands All other habitat variableswere not different (P gt 005) between burned and unburned stands

Significant habitat variables from ANOVA including litter depthcanopy height and percent bare ground cover were examined in correla-tion analyses Abundance of eastern narrowmouth toads (Gastrophrynecarolinensis H) (r = -060 P = 00396) and salamander abundance (r =-062 P = 00303) were negatively correlated with percent bare groundcover No other significant relationships (P gt 005) were observed

DISCUSSION

Our results indicate that periodic winter surface fires in bottomlandhardwood stands of the southeastern Coastal Plain do not affect overallherpetofaunal amphibian salamander or anuran abundance diversityor richness Our findings are similar to those of Ford et al (1999) whofound a high intensity restoration fire in Southern Appalachian hard-wood stands to have a negligible effect on herpetofaunal communitiesbut suggested that alterations in vital microhabitat components due toprescribed fire may negatively impact woodland salamanders On ourstudy site most habitat variables measured were similar between burned

Table 2 Mean (plusmn SD) abundance Shannon-Weiner Index and total number of species foreach taxonomic category (total amphibian salamander anuran and reptile) of herpetofaunacaptured in three burned and three unburned bottomland hardwood stands at Di-LanePlantation Wildlife Management Area Burke County Georgia July-October 2001

Burned Unburned F14

P

Total HerpetofaunaAbundance 4900 plusmn 2346 6600 plusmn 2100 084 04111Shannon-Weiner Index 188 plusmn 026 177 plusmn 014 020 06755Richness 1067 plusmn 306 1033 plusmn 058 013 07344

AmphibiansAbundance 4267 plusmn 2303 6167 plusmn 2079 112 03486Shannon-Weiner Index 148 plusmn 032 154 plusmn 011 012 07466Richness 767 plusmn 208 800 plusmn 0 002 09019

SalamandersAbundance 200 plusmn 068 583 plusmn 114 622 00671Shannon-Weiner Index 021 plusmn 006 035 plusmn 004 649 00635Richness 117 plusmn 031 150 plusmn 022 057 04918

AnuransAbundance 3867 plusmn 2108 5000 plusmn 1609 055 05003Shannon-Weiner Index 124 plusmn 035 117 plusmn 018 011 07581Richness 600 plusmn 173 633 plusmn 058 002 08944

ReptilesAbundance 667 plusmn 058 433 plusmn 058 245 00080a

Shannon-Weiner Index 040 plusmn 006 023 plusmn 005 814 00462a

Richness 300 plusmn 100 233 plusmn 058 080 04216aMeans are significantly different (P le 005)

Southeastern Naturalist Vol 2 No 4482

and unburned stands probably due to the low intensity of burns Of thethree habitat variables that were different between treatments the mostprofound was reduction in litter layer depth which subsequently pro-duced an increase in bare ground cover in burned stands

Decrease in leaf litter depth results in a reduction of on-site mois-ture which can impair dermal respiratory ability in amphibians(Duellman and Trueb 1994) and has been cited as a major factor inreduction of amphibian abundance (Ash 1995 Ash 1997) We capturedalmost twice as many eastern narrowmouth toads in unburned standscompared to burned stands (76 and 42 respectively) and found that

Table 3 Mean (plusmn SD) of habitat variables measured within a 004-ha plot around trapclusters in three burned and three unburned bottomland hardwood stands at Di-LanePlantation Wildlife Management Area Burke County Georgia JulyOctober 2001

Habitat Variable Burned Unburned F14 P

Basal area m2ha (gt 716 cm DBH)Hardwood 2408 plusmn 316 2154 plusmn 343 002 09060

Pine 659 plusmn 221 763 plusmn 343 005 08364Snaga 093 plusmn 040 116 plusmn 058 046 05369Totala 3159 plusmn 446 3032 plusmn 25 026 06369

Midstory stemsha (lt 716 cm DBH)Total stems 134167 plusmn 41966 237083 plusmn 21461 382 01223

Course Woody Debris (m3 ha)

Class 1a 2901 plusmn 1629 2359 plusmn 1437 009 07787Class 2a 588 plusmn 406 4113 plusmn 382 260 01820Class 3a 2507 plusmn 873 6371 plusmn 409 008 07871Totala 5995 plusmn 197 12843 plusmn 7272 002 08921

Canopy height (m) 2114 plusmn 122 2431 plusmn 118 1230 00247b

Midstory height (m) 1339 plusmn 083 1212 plusmn 079 088 04023

Understory (0-155m) stems6m2

Grassa 3700 plusmn 3192 117 plusmn117 108 03582Shruba 7533 plusmn 3357 26 plusmn 1842 505 00880Vine 3033 plusmn 1052 3783 plusmn 1034 020 06785Woody 2583 plusmn 531 3117 plusmn 88 040 05617Other 117 plusmn 075 15 plusmn 096 004 08554Total stems 16967 plusmn 4242 9767 plusmn 1737 433 01059

Understory height (cm) 2641 plusmn 470 259 plusmn 48 001 09294

Percent ground coverPine litter 2436 plusmn 760 1986 plusmn 1027 007 08067Hardwood litter 4917 plusmn 970 7703 plusmn 996 217 02148Bare grounda 2028 plusmn 537 117 plusmn 094 4239 00029b

Woody debrisa 571 plusmn 242 194 plusmn 09 195 02349

Litter depth (cm) 093 plusmn 023 286 plusmn 034 1247 00242b

pH 433 plusmn 011 445 plusmn 016 040 05591

Distance to edge (m) 8650 plusmn 993 1315 plusmn 2155 331 0143

Distance to water (m) 74017 plusmn 18026 122617 plusmn 15381 172 02594aVariable was log10 transformedbMeans are significantly different (P le 005)

KR Moseley SB Castleberry and SH Schweitzer2003 483

abundance was negatively correlated with percent bare ground coverSalamander abundance also was negatively correlated with percent bareground cover Although we did not test for differences of individualspecies between treatments because of low sample sizes our resultssuggest that increases in bare ground cover as a result of leaf litterreductions may be associated with lower abundance of some amphib-ians Other studies have demonstrated similar results where reduction inleaf litter depth was a result of forest harvesting (Ash 1995 Ash 1997Grialou et al 2000 McLeod and Gates 1998 Pough et al 1987) Ourstudy apparently is the first to find such a relationship in bottomlandhardwoods as a result of prescribed fire

Several amphibian species captured possess behavioral characteris-tics important in avoiding desiccation Southern toads (Bufo terrestrisB) eastern spadefoot toads (Scaphiopus holbrooki H) and Americantoads (Bufo americanus Holbrook) together representing 461 and286 of amphibian captures in burned and unburned plots respec-tively are capable of burrowing into moist soil to avoid dry surfaceconditions (Conant and Collins 1998) The ability of these species toescape such conditions by burrowing may enable them to survive instands with a reduced litter layer

Total CWD volume and volume of all CWD decay classes did not differbetween burned and unburned stands Other studies have found decreasesin CWD and consequent decreases in amphibian abundance in areassubjected to prescribed fire (McLeod and Gates 1998) All species cap-tured in our study possess some dependence on CWD (Conant and Collins1998 Harmon et al 1986 Maidens et al 1998 Whiles and Grubaugh1996 Wilson 1995) For example eastern narrowmouth toads represent-ing 328 and 41 of amphibians captured in burned and unburnedstands respectively typically occupy habitats that provide moisture andshelter such as under CWD (Conant and Collins 1998) CWD has arelatively constant moisture regime and can remain moist during long dryperiods providing refuge for amphibians (Boddy 1983) Furthermorearthropods are abundant in and around CWD (Graham 1935 Hanula 1993Harmon et al 1986 Savely 1939) yielding an important food source forinsectivorous herpetofauna These characteristics of CWD may havecompensated to some degree for leaf litter reduction by providing adequateamounts of moisture and food for amphibians

Similar to our results previous studies examining fire effects onherpetofauna in southeastern Coastal Plain pine communities also foundreptiles to increase in diversity and abundance following application ofprescribed fire (Means and Campbell 1981 Mushinsky 1985) attributingthe increases to greater thermoregulatory opportunity within burnedstands Increased bare ground cover in burned stands resulted in a largerground surface area for thermoregulatory activity by reptiles When solar

Southeastern Naturalist Vol 2 No 4484

radiation declines in the evening lizards will leave their perches to flattenout against the warm sand to maintain optimal temperature (Pough et al2001) Many species of reptiles also utilize CWD as a basking substrate aswell as for shelter and nesting (Conant and Collins 1998 Pough et al 2001Wilson 1995) Numerous lizards were observed basking on logs at the edgeof burned stands during our study Five-lined skinks (Eumeces fasciatusL) representing 579 and 385 of reptile species captured in burnedand unburned stands respectively frequently utilize CWD as nesting sitesbecause of the relatively constant microclimate provided for egg develop-ment (Hecnar 1994) We recognize that the observed increase in reptilediversity and abundance in our study may be an underestimate because ourtrapping methods are relatively ineffective at catching large snakes whichwere likely underrepresented in our collections

We conclude that the use of low intensity prescribed fire had littleoverall effect on herpetofauna in our bottomland hardwood forest studyarea However reductions in leaf litter and associated increases in bareground cover may be associated with lower abundance of some amphib-ians Conversely the increased ground surface area available for ther-moregulatory activity may benefit some reptiles CWD was relativelyunaffected by the burning regime probably because of the low intensityof the fires In areas where fire intensity is greater the impacts on theherpetofaunal community may be more pronounced

ACKNOWLEDGMENTS

We thank V VanSant H Barnhill and J Bearden of the Georgia Depart-ment of Natural Resources Wildlife Resources Division for their assistance andaccess to Di-Lane WMA We also thank K Francl for GIS assistance

LITERATURE CITED

Ash AN 1995 Effects of clear-cutting on litter parameters in the SouthernBlue Ridge mountains Castanea 60(2)8997

Ash AN 1997 Disappearance and return of Plethodontid salamanders toclearcut plots in the southern Blue Ridge mountains Conservation Biology11(4)983989

Bennett SH JW Gibbons and J Glanville 1980 Terrestrial activity abun-dance and diversity of amphibians in differently managed forest typesAmerican Midland Naturalist 103(2)412416

Boddy L 1983 Microclimate and moisture dynamics of wood decomposing interrestrial ecosystems Soil Biology and Biochemistry 15(2)149157

Boughton RG J Staiger and R Franz 2000 Use of PVC refugia as asampling technique for hylid treefrogs American Midland Naturalist144(1)168177

Conant R and JT Collins 1998 A Field Guide to Reptiles and Amphibians ofEasternCentral North America 3rd Edition Houghton Mifflin Co BostonMA 616 pp

KR Moseley SB Castleberry and SH Schweitzer2003 485

Corn PS 1994 Straight line drift fences and pitfall traps Pp 109117 InWR Heyer MA Donnelly RW McDiarmid LC Hayek and MSFoster (Eds) Measuring and Monitoring Biological Diversity StandardMethods for Amphibians Smithsonian Institute Press Washington DC364 pp

DeMaynadier PG and ML Hunter Jr 1995 The relationship between forestmanagement and amphibian ecology A review of the North Americanliterature Environmental Reviews 3(3-4)230261

Duellman WE and L Trueb 1994 Biology of Amphibians The JohnsHopkins University Press Baltimore MD 670 pp

Environmental Systems Research Institute 1999 ARCVIEW Version 32Environmental Systems Research Institute Redlands CA

Fellers GM and CA Drost 1994 Sampling with artificial cover Pp 146-150 In WR Heyer MA Donnelly RW McDiarmid LC Hayek andMS Foster (Eds) Measuring and Monitoring Biological Diversity Stan-dard Methods for Amphibians Smithsonian Institute Press WashingtonDC 364 pp

Ford WM MA Menzel DW McGill J Laerm and TS McCay 1999Effects of a community restoration fire on small mammals and herpetofaunain the Southern Appalachians Forest Ecology and Management 114(2-3)233243

Graham SA 1935 The felled tree trunk as an ecological unit Ecology6(4)397412

Greenberg CH DG Neary and LD Harris 1994 Effect of high-intensitywildfire and silvicultural treatments on reptile communities in sand-pinescrub Conservation Biology 8(4)10471057

Grialou JA SD West and RN Wilkins 2000 The effects of forest clearcutharvesting and thinning on terrestrial salamanders Journal of Wildlife Man-agement 64(1)105113

Hanula JL 1993 Relationship of wood-feeding insects and coarse woodydebris Pp 5565 In JW McMinn and DA Crossley Jr (Eds)Biodiversity of Coarse Woody Debris in Southern Forests Effects onBiodiversity Proceedings of the Workshop on Coarse Woody Debris inSouthern Forests US Department of Agriculture Forest Service GeneralTechnical Report SE-94 146 pp

Harmon ME JF Franklin FJ Swanson P Sollins SV Gregory JDLattin NH Anderson SP Cline NG Aumen JR Sedell GWLienkaemper K Cromack Jr KW Cummins 1986 Ecology of coarsewoody debris in temperate ecosystems Advances in Ecological Research15133302

Hecnar SJ 1994 Nest distribution site selection and brooding in the five-linedskink (Eumeces fasciatus) Canadian Journal of Zoology 72(8)15101516

Maidens DA MA Menzel and J Laerm 1998 Notes on the effect of sizeand level of decay of coarse woody debris on relative abundance of shrewsand salamanders in the southern Appalachian mountains Georgia Journal ofScience 56(4)226-233

McLeod RF and JE Gates 1998 Response of herpetofaunal communities toforest cutting and burning at Chesapeake Farms Maryland American Mid-land Naturalist 139(1)164177

Southeastern Naturalist Vol 2 No 4486

Means DB and HW Campbell 1981 Effects of prescribed burningonamphibians and reptiles Pp 8996 In GW Wood (Ed) Prescribed Fireand Wildlife in Southern Forests Belle Baruch Forest Science InstituteClemson University Georgetown SC 170 pp

Mushinsky HR 1985 Fire and the Florida sandhill herptofaunal communitywith special attention to responses of Cnemidophorus sexlineatusHerpetologica 41(3)333342

Nyland RD 1996 Silviculture Concepts and Applications McGraw-HillNew York NY 633 pp

Pielou EC 1977 Mathematical Ecology John Wiley and Sons New YorkNY 385 pp

Pough FH RM Andrews JE Cadle ML Crump AH Savitzky and KDWells 2001 Herpetology 2ndEdition Prentice Hall NJ 612 pp

Russell KR DH Van Lear and DC Guynn Jr 1999 Prescribed fire effectson herpetofauna Review and management implications Wildlife SocietyBulletin 27(2)374384

Savely HE 1939 Ecological relations of certain animals in dead pine and oaklogs Ecological Monographs 9(3)322377

SAS Institute 1997 SASSTAT software Changes and enhancements throughrelease 612 SAS Institute Cary NC

Smith DM BC Larson MJ Kelty and PMS Ashton 1997 The Practice ofSilviculture Applied Forest Ecology Wiley and Sons New York NY 537 pp

Spotila JR 1972 Role of temperature and water in the ecology of lunglesssalamanders Ecological Monographs 42(1)95125

Stoddard HL 1962 Use of fire in pine forests and game lands of the deepSoutheast Proceedings of the Tall Timbers Fire Ecology Conference TallTimbers Research Station Tallahassee FL 13142

Van Lear DH and TA Waldrop 1989 History uses and effects of fire in theAppalachians US Department of Agriculture Forest Service GeneralTechnical Report SE-54 20 pp

Walker LC 1999 The North American Forests Geography Ecology andSilviculture CRC Press Boca Raton FL 398 pp

Whiles MR and JW Grubaugh 1996 Importance of coarse woody debris tosouthern forest herpetofauna Pp 94100 In JW McMinn and DACrossley Jr (Eds) Biodiversity of Coarse Woody Debris in SouthernForests Effects on Biodiversity Proceedings of the Workshop on CoarseWoody Debris in Southern Forests US Department of Agriculture ForestService General Technical Report SE-94 146 pp

Wilson LA 1995 Land Manageriacutes Guide to the Amphibians and Reptiles ofthe SouthThe Nature Conservancy Southeastern Region Chapel Hill NC324 pp

KR Moseley SB Castleberry and SH Schweitzer2003 479

recorded for all trees ge 72 cm DBH Basal area was calculated for pinehardwood and snag components Midstory stems (lt 72 cm DBH andge155 m tall) were tallied by species Diameter and length of CWDgreater than 72 cm mean diameter were recorded to determine volumeof CWD All CWD measured was categorized based on degree ofdecomposition class one included logs with a hard outer covering andno soft parts class two included logs with hard or soft outer parts andclass three included logs with completely soft outer parts (Maidens et al1998) Canopy and midstory height (m) were determined using a cli-nometer One canopy and one midstory tree representative of the aver-age height in each plot were measured (m)

Within each 004-ha plot three transect lines 120o apart wereestablished beginning at the center of each array and extending out113 m (Fig 2) Two 1-m2 plots were randomly located along eachtransect line Within each plot understory stems (0155 m) werecounted and percentage of ground cover was estimated Understorystems were placed into one of five categories grass shrub vinewoody and other A stem representing average understory height (cm)within each 1-m2 plot was measured Percent ground cover of pinelitter hardwood litter bare ground and woody debris was estimatedAt three random points along each transect litter depth was measuredto the nearest mm by inserting a ruler into the litter layer Soil pH wasdetermined by taking a mixture of six random soil samples from withinthe 004-ha plot Soil samples were analyzed by the Georgia Coopera-tive Extension Service Athens Distance of array to habitat edge andnearest body of water was determined using ARCVIEW (Environmen-tal Systems Research Institute 1999)

Captured species were pooled into the following taxonomic catego-ries for analysis total herpetofauna amphibians salamanders anuransand reptiles Reptilian taxa were not examined separately because oflow sample sizes Abundance was expressed as the total number ofcaptures per trap cluster and diversity was calculated using the Shan-non-Weiner index (Pielou 1977) Recaptures were not included in abun-dance or diversity calculations A one-way analysis of variance(ANOVA) was used to test taxonomic categories for differences inabundance diversity and richness between treatments Because a par-ticular stand received only a single treatment (burn or no burn) theeffect of stand was nested within treatments and used as the error termwhen testing for treatment effects Habitat variables and distance ofarray to water and habitat edge also were analyzed as a one-wayANOVA with nested effects as described above Habitat variables thatwere not normally distributed were log10 transformed

Spearmans rank correlation was used to examine relationships be-tween habitat variables and abundance of individual species with gt 20

Southeastern Naturalist Vol 2 No 4480

total captures as well as abundance richness and diversity of taxo-nomic categories Only habitat variables that differed between burnedand unburned stands using ANOVA were examined in correlationanalysis All statistical analyses were conducted using SAS statisticalsoftware (SAS Institute 1997)

RESULTS

A total of 346 individuals representing 21 species was captured in348 array nights (Table 1) Of those captured 33 (eight species) werereptiles and 313 (13 species) were amphibians Seven species wererepresented by a single capture Over the trapping period 33 individualswere recaptured

Overall herpetofaunal abundance diversity and richness were notdifferent (P gt 005) between burned and unburned stands (Table 2) Nodifferences (P gt 005) were observed in amphibian salamander oranuran abundance diversity or richness between burned and unburnedstands (Table 2) Reptile abundance (P = 00080) and diversity (P =00462) were greater in burned than unburned stands However reptilespecies richness was not different (P = 04216) between burned andunburned stands (Table 2)

Of habitat variables examined litter depth (P = 00242) and canopyheight (P = 00247) were greater in unburned than in burned stands

Table 1 Numbers of herpetofauna species captured in burned and unburned bottomlandhardwood stands at Di-Lane Plantation Wildlife Management Area Burke County Geor-gia JulyOctober 2001

Species Common Name Burned Unburned

Gastrophryne carolinensis Holbrook Eastern Narrowmouth Toad 42 76Bufo terrestris Bonnaterre Southern Toad 41 19Scaphiopus holbrookii Harlan Eastern Spadefoot 11 29Eumeces fasciatus Linnaeus Five-lined Skink 11 5Bufo americanus Holbrook American Toad 7 5Ambystoma opacum Gravenhorst Marbled Salamander 6 22Hyla versicolor LeConte Gray Treefrog 6 2Plethodon glutinosus Green Slimy Salamander 5 13Rana utricularia Schreber Southern Leopard Frog 4 13Scincella lateralis Say Ground Skink 4 1Rana catesbeiana Shaw Bullfrog 3 4Notophthalmus viridescens Rafinesque Red-spotted Newt 1 0Rana clamitans clamitans Latreille Bronze Frog 1 0Pseudacris ornata Holbrook Ornate Chorus Frog 1 1Eumeces laticeps Schneider Broadhead Skink 1 0Sceloporus undulatus Bosc and Daudin Fence Lizard 1 0Tantilla coronata Baird and Girard Southeastern Crowned Snake 1 0Elaphe obsoleta obsoleta Say Black Rat Snake 1 0

Diadophis punctatus punctatus Linnaeus Southern Ringneck Snake 1 1Hyla cinerea Schneider Green Treefrog 0 1

Anolis carolinensis Voigt Green Anole 0 6

KR Moseley SB Castleberry and SH Schweitzer2003 481

(Table 3) Percent bare ground cover was greater (P = 00029) in burned(2028 ) than unburned (117 ) stands All other habitat variableswere not different (P gt 005) between burned and unburned stands

Significant habitat variables from ANOVA including litter depthcanopy height and percent bare ground cover were examined in correla-tion analyses Abundance of eastern narrowmouth toads (Gastrophrynecarolinensis H) (r = -060 P = 00396) and salamander abundance (r =-062 P = 00303) were negatively correlated with percent bare groundcover No other significant relationships (P gt 005) were observed

DISCUSSION

Our results indicate that periodic winter surface fires in bottomlandhardwood stands of the southeastern Coastal Plain do not affect overallherpetofaunal amphibian salamander or anuran abundance diversityor richness Our findings are similar to those of Ford et al (1999) whofound a high intensity restoration fire in Southern Appalachian hard-wood stands to have a negligible effect on herpetofaunal communitiesbut suggested that alterations in vital microhabitat components due toprescribed fire may negatively impact woodland salamanders On ourstudy site most habitat variables measured were similar between burned

Table 2 Mean (plusmn SD) abundance Shannon-Weiner Index and total number of species foreach taxonomic category (total amphibian salamander anuran and reptile) of herpetofaunacaptured in three burned and three unburned bottomland hardwood stands at Di-LanePlantation Wildlife Management Area Burke County Georgia July-October 2001

Burned Unburned F14

P

Total HerpetofaunaAbundance 4900 plusmn 2346 6600 plusmn 2100 084 04111Shannon-Weiner Index 188 plusmn 026 177 plusmn 014 020 06755Richness 1067 plusmn 306 1033 plusmn 058 013 07344

AmphibiansAbundance 4267 plusmn 2303 6167 plusmn 2079 112 03486Shannon-Weiner Index 148 plusmn 032 154 plusmn 011 012 07466Richness 767 plusmn 208 800 plusmn 0 002 09019

SalamandersAbundance 200 plusmn 068 583 plusmn 114 622 00671Shannon-Weiner Index 021 plusmn 006 035 plusmn 004 649 00635Richness 117 plusmn 031 150 plusmn 022 057 04918

AnuransAbundance 3867 plusmn 2108 5000 plusmn 1609 055 05003Shannon-Weiner Index 124 plusmn 035 117 plusmn 018 011 07581Richness 600 plusmn 173 633 plusmn 058 002 08944

ReptilesAbundance 667 plusmn 058 433 plusmn 058 245 00080a

Shannon-Weiner Index 040 plusmn 006 023 plusmn 005 814 00462a

Richness 300 plusmn 100 233 plusmn 058 080 04216aMeans are significantly different (P le 005)

Southeastern Naturalist Vol 2 No 4482

and unburned stands probably due to the low intensity of burns Of thethree habitat variables that were different between treatments the mostprofound was reduction in litter layer depth which subsequently pro-duced an increase in bare ground cover in burned stands

Decrease in leaf litter depth results in a reduction of on-site mois-ture which can impair dermal respiratory ability in amphibians(Duellman and Trueb 1994) and has been cited as a major factor inreduction of amphibian abundance (Ash 1995 Ash 1997) We capturedalmost twice as many eastern narrowmouth toads in unburned standscompared to burned stands (76 and 42 respectively) and found that

Table 3 Mean (plusmn SD) of habitat variables measured within a 004-ha plot around trapclusters in three burned and three unburned bottomland hardwood stands at Di-LanePlantation Wildlife Management Area Burke County Georgia JulyOctober 2001

Habitat Variable Burned Unburned F14 P

Basal area m2ha (gt 716 cm DBH)Hardwood 2408 plusmn 316 2154 plusmn 343 002 09060

Pine 659 plusmn 221 763 plusmn 343 005 08364Snaga 093 plusmn 040 116 plusmn 058 046 05369Totala 3159 plusmn 446 3032 plusmn 25 026 06369

Midstory stemsha (lt 716 cm DBH)Total stems 134167 plusmn 41966 237083 plusmn 21461 382 01223

Course Woody Debris (m3 ha)

Class 1a 2901 plusmn 1629 2359 plusmn 1437 009 07787Class 2a 588 plusmn 406 4113 plusmn 382 260 01820Class 3a 2507 plusmn 873 6371 plusmn 409 008 07871Totala 5995 plusmn 197 12843 plusmn 7272 002 08921

Canopy height (m) 2114 plusmn 122 2431 plusmn 118 1230 00247b

Midstory height (m) 1339 plusmn 083 1212 plusmn 079 088 04023

Understory (0-155m) stems6m2

Grassa 3700 plusmn 3192 117 plusmn117 108 03582Shruba 7533 plusmn 3357 26 plusmn 1842 505 00880Vine 3033 plusmn 1052 3783 plusmn 1034 020 06785Woody 2583 plusmn 531 3117 plusmn 88 040 05617Other 117 plusmn 075 15 plusmn 096 004 08554Total stems 16967 plusmn 4242 9767 plusmn 1737 433 01059

Understory height (cm) 2641 plusmn 470 259 plusmn 48 001 09294

Percent ground coverPine litter 2436 plusmn 760 1986 plusmn 1027 007 08067Hardwood litter 4917 plusmn 970 7703 plusmn 996 217 02148Bare grounda 2028 plusmn 537 117 plusmn 094 4239 00029b

Woody debrisa 571 plusmn 242 194 plusmn 09 195 02349

Litter depth (cm) 093 plusmn 023 286 plusmn 034 1247 00242b

pH 433 plusmn 011 445 plusmn 016 040 05591

Distance to edge (m) 8650 plusmn 993 1315 plusmn 2155 331 0143

Distance to water (m) 74017 plusmn 18026 122617 plusmn 15381 172 02594aVariable was log10 transformedbMeans are significantly different (P le 005)

KR Moseley SB Castleberry and SH Schweitzer2003 483

abundance was negatively correlated with percent bare ground coverSalamander abundance also was negatively correlated with percent bareground cover Although we did not test for differences of individualspecies between treatments because of low sample sizes our resultssuggest that increases in bare ground cover as a result of leaf litterreductions may be associated with lower abundance of some amphib-ians Other studies have demonstrated similar results where reduction inleaf litter depth was a result of forest harvesting (Ash 1995 Ash 1997Grialou et al 2000 McLeod and Gates 1998 Pough et al 1987) Ourstudy apparently is the first to find such a relationship in bottomlandhardwoods as a result of prescribed fire

Several amphibian species captured possess behavioral characteris-tics important in avoiding desiccation Southern toads (Bufo terrestrisB) eastern spadefoot toads (Scaphiopus holbrooki H) and Americantoads (Bufo americanus Holbrook) together representing 461 and286 of amphibian captures in burned and unburned plots respec-tively are capable of burrowing into moist soil to avoid dry surfaceconditions (Conant and Collins 1998) The ability of these species toescape such conditions by burrowing may enable them to survive instands with a reduced litter layer

Total CWD volume and volume of all CWD decay classes did not differbetween burned and unburned stands Other studies have found decreasesin CWD and consequent decreases in amphibian abundance in areassubjected to prescribed fire (McLeod and Gates 1998) All species cap-tured in our study possess some dependence on CWD (Conant and Collins1998 Harmon et al 1986 Maidens et al 1998 Whiles and Grubaugh1996 Wilson 1995) For example eastern narrowmouth toads represent-ing 328 and 41 of amphibians captured in burned and unburnedstands respectively typically occupy habitats that provide moisture andshelter such as under CWD (Conant and Collins 1998) CWD has arelatively constant moisture regime and can remain moist during long dryperiods providing refuge for amphibians (Boddy 1983) Furthermorearthropods are abundant in and around CWD (Graham 1935 Hanula 1993Harmon et al 1986 Savely 1939) yielding an important food source forinsectivorous herpetofauna These characteristics of CWD may havecompensated to some degree for leaf litter reduction by providing adequateamounts of moisture and food for amphibians

Similar to our results previous studies examining fire effects onherpetofauna in southeastern Coastal Plain pine communities also foundreptiles to increase in diversity and abundance following application ofprescribed fire (Means and Campbell 1981 Mushinsky 1985) attributingthe increases to greater thermoregulatory opportunity within burnedstands Increased bare ground cover in burned stands resulted in a largerground surface area for thermoregulatory activity by reptiles When solar

Southeastern Naturalist Vol 2 No 4484

radiation declines in the evening lizards will leave their perches to flattenout against the warm sand to maintain optimal temperature (Pough et al2001) Many species of reptiles also utilize CWD as a basking substrate aswell as for shelter and nesting (Conant and Collins 1998 Pough et al 2001Wilson 1995) Numerous lizards were observed basking on logs at the edgeof burned stands during our study Five-lined skinks (Eumeces fasciatusL) representing 579 and 385 of reptile species captured in burnedand unburned stands respectively frequently utilize CWD as nesting sitesbecause of the relatively constant microclimate provided for egg develop-ment (Hecnar 1994) We recognize that the observed increase in reptilediversity and abundance in our study may be an underestimate because ourtrapping methods are relatively ineffective at catching large snakes whichwere likely underrepresented in our collections

We conclude that the use of low intensity prescribed fire had littleoverall effect on herpetofauna in our bottomland hardwood forest studyarea However reductions in leaf litter and associated increases in bareground cover may be associated with lower abundance of some amphib-ians Conversely the increased ground surface area available for ther-moregulatory activity may benefit some reptiles CWD was relativelyunaffected by the burning regime probably because of the low intensityof the fires In areas where fire intensity is greater the impacts on theherpetofaunal community may be more pronounced

ACKNOWLEDGMENTS

We thank V VanSant H Barnhill and J Bearden of the Georgia Depart-ment of Natural Resources Wildlife Resources Division for their assistance andaccess to Di-Lane WMA We also thank K Francl for GIS assistance

LITERATURE CITED

Ash AN 1995 Effects of clear-cutting on litter parameters in the SouthernBlue Ridge mountains Castanea 60(2)8997

Ash AN 1997 Disappearance and return of Plethodontid salamanders toclearcut plots in the southern Blue Ridge mountains Conservation Biology11(4)983989

Bennett SH JW Gibbons and J Glanville 1980 Terrestrial activity abun-dance and diversity of amphibians in differently managed forest typesAmerican Midland Naturalist 103(2)412416

Boddy L 1983 Microclimate and moisture dynamics of wood decomposing interrestrial ecosystems Soil Biology and Biochemistry 15(2)149157

Boughton RG J Staiger and R Franz 2000 Use of PVC refugia as asampling technique for hylid treefrogs American Midland Naturalist144(1)168177

Conant R and JT Collins 1998 A Field Guide to Reptiles and Amphibians ofEasternCentral North America 3rd Edition Houghton Mifflin Co BostonMA 616 pp

KR Moseley SB Castleberry and SH Schweitzer2003 485

Corn PS 1994 Straight line drift fences and pitfall traps Pp 109117 InWR Heyer MA Donnelly RW McDiarmid LC Hayek and MSFoster (Eds) Measuring and Monitoring Biological Diversity StandardMethods for Amphibians Smithsonian Institute Press Washington DC364 pp

DeMaynadier PG and ML Hunter Jr 1995 The relationship between forestmanagement and amphibian ecology A review of the North Americanliterature Environmental Reviews 3(3-4)230261

Duellman WE and L Trueb 1994 Biology of Amphibians The JohnsHopkins University Press Baltimore MD 670 pp

Environmental Systems Research Institute 1999 ARCVIEW Version 32Environmental Systems Research Institute Redlands CA

Fellers GM and CA Drost 1994 Sampling with artificial cover Pp 146-150 In WR Heyer MA Donnelly RW McDiarmid LC Hayek andMS Foster (Eds) Measuring and Monitoring Biological Diversity Stan-dard Methods for Amphibians Smithsonian Institute Press WashingtonDC 364 pp

Ford WM MA Menzel DW McGill J Laerm and TS McCay 1999Effects of a community restoration fire on small mammals and herpetofaunain the Southern Appalachians Forest Ecology and Management 114(2-3)233243

Graham SA 1935 The felled tree trunk as an ecological unit Ecology6(4)397412

Greenberg CH DG Neary and LD Harris 1994 Effect of high-intensitywildfire and silvicultural treatments on reptile communities in sand-pinescrub Conservation Biology 8(4)10471057

Grialou JA SD West and RN Wilkins 2000 The effects of forest clearcutharvesting and thinning on terrestrial salamanders Journal of Wildlife Man-agement 64(1)105113

Hanula JL 1993 Relationship of wood-feeding insects and coarse woodydebris Pp 5565 In JW McMinn and DA Crossley Jr (Eds)Biodiversity of Coarse Woody Debris in Southern Forests Effects onBiodiversity Proceedings of the Workshop on Coarse Woody Debris inSouthern Forests US Department of Agriculture Forest Service GeneralTechnical Report SE-94 146 pp

Harmon ME JF Franklin FJ Swanson P Sollins SV Gregory JDLattin NH Anderson SP Cline NG Aumen JR Sedell GWLienkaemper K Cromack Jr KW Cummins 1986 Ecology of coarsewoody debris in temperate ecosystems Advances in Ecological Research15133302

Hecnar SJ 1994 Nest distribution site selection and brooding in the five-linedskink (Eumeces fasciatus) Canadian Journal of Zoology 72(8)15101516

Maidens DA MA Menzel and J Laerm 1998 Notes on the effect of sizeand level of decay of coarse woody debris on relative abundance of shrewsand salamanders in the southern Appalachian mountains Georgia Journal ofScience 56(4)226-233

McLeod RF and JE Gates 1998 Response of herpetofaunal communities toforest cutting and burning at Chesapeake Farms Maryland American Mid-land Naturalist 139(1)164177

Southeastern Naturalist Vol 2 No 4486

Means DB and HW Campbell 1981 Effects of prescribed burningonamphibians and reptiles Pp 8996 In GW Wood (Ed) Prescribed Fireand Wildlife in Southern Forests Belle Baruch Forest Science InstituteClemson University Georgetown SC 170 pp

Mushinsky HR 1985 Fire and the Florida sandhill herptofaunal communitywith special attention to responses of Cnemidophorus sexlineatusHerpetologica 41(3)333342

Nyland RD 1996 Silviculture Concepts and Applications McGraw-HillNew York NY 633 pp

Pielou EC 1977 Mathematical Ecology John Wiley and Sons New YorkNY 385 pp

Pough FH RM Andrews JE Cadle ML Crump AH Savitzky and KDWells 2001 Herpetology 2ndEdition Prentice Hall NJ 612 pp

Russell KR DH Van Lear and DC Guynn Jr 1999 Prescribed fire effectson herpetofauna Review and management implications Wildlife SocietyBulletin 27(2)374384

Savely HE 1939 Ecological relations of certain animals in dead pine and oaklogs Ecological Monographs 9(3)322377

SAS Institute 1997 SASSTAT software Changes and enhancements throughrelease 612 SAS Institute Cary NC

Smith DM BC Larson MJ Kelty and PMS Ashton 1997 The Practice ofSilviculture Applied Forest Ecology Wiley and Sons New York NY 537 pp

Spotila JR 1972 Role of temperature and water in the ecology of lunglesssalamanders Ecological Monographs 42(1)95125

Stoddard HL 1962 Use of fire in pine forests and game lands of the deepSoutheast Proceedings of the Tall Timbers Fire Ecology Conference TallTimbers Research Station Tallahassee FL 13142

Van Lear DH and TA Waldrop 1989 History uses and effects of fire in theAppalachians US Department of Agriculture Forest Service GeneralTechnical Report SE-54 20 pp

Walker LC 1999 The North American Forests Geography Ecology andSilviculture CRC Press Boca Raton FL 398 pp

Whiles MR and JW Grubaugh 1996 Importance of coarse woody debris tosouthern forest herpetofauna Pp 94100 In JW McMinn and DACrossley Jr (Eds) Biodiversity of Coarse Woody Debris in SouthernForests Effects on Biodiversity Proceedings of the Workshop on CoarseWoody Debris in Southern Forests US Department of Agriculture ForestService General Technical Report SE-94 146 pp

Wilson LA 1995 Land Manageriacutes Guide to the Amphibians and Reptiles ofthe SouthThe Nature Conservancy Southeastern Region Chapel Hill NC324 pp

Southeastern Naturalist Vol 2 No 4480

total captures as well as abundance richness and diversity of taxo-nomic categories Only habitat variables that differed between burnedand unburned stands using ANOVA were examined in correlationanalysis All statistical analyses were conducted using SAS statisticalsoftware (SAS Institute 1997)

RESULTS

A total of 346 individuals representing 21 species was captured in348 array nights (Table 1) Of those captured 33 (eight species) werereptiles and 313 (13 species) were amphibians Seven species wererepresented by a single capture Over the trapping period 33 individualswere recaptured

Overall herpetofaunal abundance diversity and richness were notdifferent (P gt 005) between burned and unburned stands (Table 2) Nodifferences (P gt 005) were observed in amphibian salamander oranuran abundance diversity or richness between burned and unburnedstands (Table 2) Reptile abundance (P = 00080) and diversity (P =00462) were greater in burned than unburned stands However reptilespecies richness was not different (P = 04216) between burned andunburned stands (Table 2)

Of habitat variables examined litter depth (P = 00242) and canopyheight (P = 00247) were greater in unburned than in burned stands

Table 1 Numbers of herpetofauna species captured in burned and unburned bottomlandhardwood stands at Di-Lane Plantation Wildlife Management Area Burke County Geor-gia JulyOctober 2001

Species Common Name Burned Unburned

Gastrophryne carolinensis Holbrook Eastern Narrowmouth Toad 42 76Bufo terrestris Bonnaterre Southern Toad 41 19Scaphiopus holbrookii Harlan Eastern Spadefoot 11 29Eumeces fasciatus Linnaeus Five-lined Skink 11 5Bufo americanus Holbrook American Toad 7 5Ambystoma opacum Gravenhorst Marbled Salamander 6 22Hyla versicolor LeConte Gray Treefrog 6 2Plethodon glutinosus Green Slimy Salamander 5 13Rana utricularia Schreber Southern Leopard Frog 4 13Scincella lateralis Say Ground Skink 4 1Rana catesbeiana Shaw Bullfrog 3 4Notophthalmus viridescens Rafinesque Red-spotted Newt 1 0Rana clamitans clamitans Latreille Bronze Frog 1 0Pseudacris ornata Holbrook Ornate Chorus Frog 1 1Eumeces laticeps Schneider Broadhead Skink 1 0Sceloporus undulatus Bosc and Daudin Fence Lizard 1 0Tantilla coronata Baird and Girard Southeastern Crowned Snake 1 0Elaphe obsoleta obsoleta Say Black Rat Snake 1 0

Diadophis punctatus punctatus Linnaeus Southern Ringneck Snake 1 1Hyla cinerea Schneider Green Treefrog 0 1

Anolis carolinensis Voigt Green Anole 0 6

KR Moseley SB Castleberry and SH Schweitzer2003 481

(Table 3) Percent bare ground cover was greater (P = 00029) in burned(2028 ) than unburned (117 ) stands All other habitat variableswere not different (P gt 005) between burned and unburned stands

Significant habitat variables from ANOVA including litter depthcanopy height and percent bare ground cover were examined in correla-tion analyses Abundance of eastern narrowmouth toads (Gastrophrynecarolinensis H) (r = -060 P = 00396) and salamander abundance (r =-062 P = 00303) were negatively correlated with percent bare groundcover No other significant relationships (P gt 005) were observed

DISCUSSION

Our results indicate that periodic winter surface fires in bottomlandhardwood stands of the southeastern Coastal Plain do not affect overallherpetofaunal amphibian salamander or anuran abundance diversityor richness Our findings are similar to those of Ford et al (1999) whofound a high intensity restoration fire in Southern Appalachian hard-wood stands to have a negligible effect on herpetofaunal communitiesbut suggested that alterations in vital microhabitat components due toprescribed fire may negatively impact woodland salamanders On ourstudy site most habitat variables measured were similar between burned

Table 2 Mean (plusmn SD) abundance Shannon-Weiner Index and total number of species foreach taxonomic category (total amphibian salamander anuran and reptile) of herpetofaunacaptured in three burned and three unburned bottomland hardwood stands at Di-LanePlantation Wildlife Management Area Burke County Georgia July-October 2001

Burned Unburned F14

P

Total HerpetofaunaAbundance 4900 plusmn 2346 6600 plusmn 2100 084 04111Shannon-Weiner Index 188 plusmn 026 177 plusmn 014 020 06755Richness 1067 plusmn 306 1033 plusmn 058 013 07344

AmphibiansAbundance 4267 plusmn 2303 6167 plusmn 2079 112 03486Shannon-Weiner Index 148 plusmn 032 154 plusmn 011 012 07466Richness 767 plusmn 208 800 plusmn 0 002 09019

SalamandersAbundance 200 plusmn 068 583 plusmn 114 622 00671Shannon-Weiner Index 021 plusmn 006 035 plusmn 004 649 00635Richness 117 plusmn 031 150 plusmn 022 057 04918

AnuransAbundance 3867 plusmn 2108 5000 plusmn 1609 055 05003Shannon-Weiner Index 124 plusmn 035 117 plusmn 018 011 07581Richness 600 plusmn 173 633 plusmn 058 002 08944

ReptilesAbundance 667 plusmn 058 433 plusmn 058 245 00080a

Shannon-Weiner Index 040 plusmn 006 023 plusmn 005 814 00462a

Richness 300 plusmn 100 233 plusmn 058 080 04216aMeans are significantly different (P le 005)

Southeastern Naturalist Vol 2 No 4482

and unburned stands probably due to the low intensity of burns Of thethree habitat variables that were different between treatments the mostprofound was reduction in litter layer depth which subsequently pro-duced an increase in bare ground cover in burned stands

Decrease in leaf litter depth results in a reduction of on-site mois-ture which can impair dermal respiratory ability in amphibians(Duellman and Trueb 1994) and has been cited as a major factor inreduction of amphibian abundance (Ash 1995 Ash 1997) We capturedalmost twice as many eastern narrowmouth toads in unburned standscompared to burned stands (76 and 42 respectively) and found that

Table 3 Mean (plusmn SD) of habitat variables measured within a 004-ha plot around trapclusters in three burned and three unburned bottomland hardwood stands at Di-LanePlantation Wildlife Management Area Burke County Georgia JulyOctober 2001

Habitat Variable Burned Unburned F14 P

Basal area m2ha (gt 716 cm DBH)Hardwood 2408 plusmn 316 2154 plusmn 343 002 09060

Pine 659 plusmn 221 763 plusmn 343 005 08364Snaga 093 plusmn 040 116 plusmn 058 046 05369Totala 3159 plusmn 446 3032 plusmn 25 026 06369

Midstory stemsha (lt 716 cm DBH)Total stems 134167 plusmn 41966 237083 plusmn 21461 382 01223

Course Woody Debris (m3 ha)

Class 1a 2901 plusmn 1629 2359 plusmn 1437 009 07787Class 2a 588 plusmn 406 4113 plusmn 382 260 01820Class 3a 2507 plusmn 873 6371 plusmn 409 008 07871Totala 5995 plusmn 197 12843 plusmn 7272 002 08921

Canopy height (m) 2114 plusmn 122 2431 plusmn 118 1230 00247b

Midstory height (m) 1339 plusmn 083 1212 plusmn 079 088 04023

Understory (0-155m) stems6m2

Grassa 3700 plusmn 3192 117 plusmn117 108 03582Shruba 7533 plusmn 3357 26 plusmn 1842 505 00880Vine 3033 plusmn 1052 3783 plusmn 1034 020 06785Woody 2583 plusmn 531 3117 plusmn 88 040 05617Other 117 plusmn 075 15 plusmn 096 004 08554Total stems 16967 plusmn 4242 9767 plusmn 1737 433 01059

Understory height (cm) 2641 plusmn 470 259 plusmn 48 001 09294

Percent ground coverPine litter 2436 plusmn 760 1986 plusmn 1027 007 08067Hardwood litter 4917 plusmn 970 7703 plusmn 996 217 02148Bare grounda 2028 plusmn 537 117 plusmn 094 4239 00029b

Woody debrisa 571 plusmn 242 194 plusmn 09 195 02349

Litter depth (cm) 093 plusmn 023 286 plusmn 034 1247 00242b

pH 433 plusmn 011 445 plusmn 016 040 05591

Distance to edge (m) 8650 plusmn 993 1315 plusmn 2155 331 0143

Distance to water (m) 74017 plusmn 18026 122617 plusmn 15381 172 02594aVariable was log10 transformedbMeans are significantly different (P le 005)

KR Moseley SB Castleberry and SH Schweitzer2003 483

abundance was negatively correlated with percent bare ground coverSalamander abundance also was negatively correlated with percent bareground cover Although we did not test for differences of individualspecies between treatments because of low sample sizes our resultssuggest that increases in bare ground cover as a result of leaf litterreductions may be associated with lower abundance of some amphib-ians Other studies have demonstrated similar results where reduction inleaf litter depth was a result of forest harvesting (Ash 1995 Ash 1997Grialou et al 2000 McLeod and Gates 1998 Pough et al 1987) Ourstudy apparently is the first to find such a relationship in bottomlandhardwoods as a result of prescribed fire

Several amphibian species captured possess behavioral characteris-tics important in avoiding desiccation Southern toads (Bufo terrestrisB) eastern spadefoot toads (Scaphiopus holbrooki H) and Americantoads (Bufo americanus Holbrook) together representing 461 and286 of amphibian captures in burned and unburned plots respec-tively are capable of burrowing into moist soil to avoid dry surfaceconditions (Conant and Collins 1998) The ability of these species toescape such conditions by burrowing may enable them to survive instands with a reduced litter layer

Total CWD volume and volume of all CWD decay classes did not differbetween burned and unburned stands Other studies have found decreasesin CWD and consequent decreases in amphibian abundance in areassubjected to prescribed fire (McLeod and Gates 1998) All species cap-tured in our study possess some dependence on CWD (Conant and Collins1998 Harmon et al 1986 Maidens et al 1998 Whiles and Grubaugh1996 Wilson 1995) For example eastern narrowmouth toads represent-ing 328 and 41 of amphibians captured in burned and unburnedstands respectively typically occupy habitats that provide moisture andshelter such as under CWD (Conant and Collins 1998) CWD has arelatively constant moisture regime and can remain moist during long dryperiods providing refuge for amphibians (Boddy 1983) Furthermorearthropods are abundant in and around CWD (Graham 1935 Hanula 1993Harmon et al 1986 Savely 1939) yielding an important food source forinsectivorous herpetofauna These characteristics of CWD may havecompensated to some degree for leaf litter reduction by providing adequateamounts of moisture and food for amphibians

Similar to our results previous studies examining fire effects onherpetofauna in southeastern Coastal Plain pine communities also foundreptiles to increase in diversity and abundance following application ofprescribed fire (Means and Campbell 1981 Mushinsky 1985) attributingthe increases to greater thermoregulatory opportunity within burnedstands Increased bare ground cover in burned stands resulted in a largerground surface area for thermoregulatory activity by reptiles When solar

Southeastern Naturalist Vol 2 No 4484

radiation declines in the evening lizards will leave their perches to flattenout against the warm sand to maintain optimal temperature (Pough et al2001) Many species of reptiles also utilize CWD as a basking substrate aswell as for shelter and nesting (Conant and Collins 1998 Pough et al 2001Wilson 1995) Numerous lizards were observed basking on logs at the edgeof burned stands during our study Five-lined skinks (Eumeces fasciatusL) representing 579 and 385 of reptile species captured in burnedand unburned stands respectively frequently utilize CWD as nesting sitesbecause of the relatively constant microclimate provided for egg develop-ment (Hecnar 1994) We recognize that the observed increase in reptilediversity and abundance in our study may be an underestimate because ourtrapping methods are relatively ineffective at catching large snakes whichwere likely underrepresented in our collections

We conclude that the use of low intensity prescribed fire had littleoverall effect on herpetofauna in our bottomland hardwood forest studyarea However reductions in leaf litter and associated increases in bareground cover may be associated with lower abundance of some amphib-ians Conversely the increased ground surface area available for ther-moregulatory activity may benefit some reptiles CWD was relativelyunaffected by the burning regime probably because of the low intensityof the fires In areas where fire intensity is greater the impacts on theherpetofaunal community may be more pronounced

ACKNOWLEDGMENTS

We thank V VanSant H Barnhill and J Bearden of the Georgia Depart-ment of Natural Resources Wildlife Resources Division for their assistance andaccess to Di-Lane WMA We also thank K Francl for GIS assistance

LITERATURE CITED

Ash AN 1995 Effects of clear-cutting on litter parameters in the SouthernBlue Ridge mountains Castanea 60(2)8997

Ash AN 1997 Disappearance and return of Plethodontid salamanders toclearcut plots in the southern Blue Ridge mountains Conservation Biology11(4)983989

Bennett SH JW Gibbons and J Glanville 1980 Terrestrial activity abun-dance and diversity of amphibians in differently managed forest typesAmerican Midland Naturalist 103(2)412416

Boddy L 1983 Microclimate and moisture dynamics of wood decomposing interrestrial ecosystems Soil Biology and Biochemistry 15(2)149157

Boughton RG J Staiger and R Franz 2000 Use of PVC refugia as asampling technique for hylid treefrogs American Midland Naturalist144(1)168177

Conant R and JT Collins 1998 A Field Guide to Reptiles and Amphibians ofEasternCentral North America 3rd Edition Houghton Mifflin Co BostonMA 616 pp

KR Moseley SB Castleberry and SH Schweitzer2003 485

Corn PS 1994 Straight line drift fences and pitfall traps Pp 109117 InWR Heyer MA Donnelly RW McDiarmid LC Hayek and MSFoster (Eds) Measuring and Monitoring Biological Diversity StandardMethods for Amphibians Smithsonian Institute Press Washington DC364 pp

DeMaynadier PG and ML Hunter Jr 1995 The relationship between forestmanagement and amphibian ecology A review of the North Americanliterature Environmental Reviews 3(3-4)230261

Duellman WE and L Trueb 1994 Biology of Amphibians The JohnsHopkins University Press Baltimore MD 670 pp

Environmental Systems Research Institute 1999 ARCVIEW Version 32Environmental Systems Research Institute Redlands CA

Fellers GM and CA Drost 1994 Sampling with artificial cover Pp 146-150 In WR Heyer MA Donnelly RW McDiarmid LC Hayek andMS Foster (Eds) Measuring and Monitoring Biological Diversity Stan-dard Methods for Amphibians Smithsonian Institute Press WashingtonDC 364 pp

Ford WM MA Menzel DW McGill J Laerm and TS McCay 1999Effects of a community restoration fire on small mammals and herpetofaunain the Southern Appalachians Forest Ecology and Management 114(2-3)233243

Graham SA 1935 The felled tree trunk as an ecological unit Ecology6(4)397412

Greenberg CH DG Neary and LD Harris 1994 Effect of high-intensitywildfire and silvicultural treatments on reptile communities in sand-pinescrub Conservation Biology 8(4)10471057

Grialou JA SD West and RN Wilkins 2000 The effects of forest clearcutharvesting and thinning on terrestrial salamanders Journal of Wildlife Man-agement 64(1)105113

Hanula JL 1993 Relationship of wood-feeding insects and coarse woodydebris Pp 5565 In JW McMinn and DA Crossley Jr (Eds)Biodiversity of Coarse Woody Debris in Southern Forests Effects onBiodiversity Proceedings of the Workshop on Coarse Woody Debris inSouthern Forests US Department of Agriculture Forest Service GeneralTechnical Report SE-94 146 pp

Harmon ME JF Franklin FJ Swanson P Sollins SV Gregory JDLattin NH Anderson SP Cline NG Aumen JR Sedell GWLienkaemper K Cromack Jr KW Cummins 1986 Ecology of coarsewoody debris in temperate ecosystems Advances in Ecological Research15133302

Hecnar SJ 1994 Nest distribution site selection and brooding in the five-linedskink (Eumeces fasciatus) Canadian Journal of Zoology 72(8)15101516

Maidens DA MA Menzel and J Laerm 1998 Notes on the effect of sizeand level of decay of coarse woody debris on relative abundance of shrewsand salamanders in the southern Appalachian mountains Georgia Journal ofScience 56(4)226-233

McLeod RF and JE Gates 1998 Response of herpetofaunal communities toforest cutting and burning at Chesapeake Farms Maryland American Mid-land Naturalist 139(1)164177

Southeastern Naturalist Vol 2 No 4486

Means DB and HW Campbell 1981 Effects of prescribed burningonamphibians and reptiles Pp 8996 In GW Wood (Ed) Prescribed Fireand Wildlife in Southern Forests Belle Baruch Forest Science InstituteClemson University Georgetown SC 170 pp

Mushinsky HR 1985 Fire and the Florida sandhill herptofaunal communitywith special attention to responses of Cnemidophorus sexlineatusHerpetologica 41(3)333342

Nyland RD 1996 Silviculture Concepts and Applications McGraw-HillNew York NY 633 pp

Pielou EC 1977 Mathematical Ecology John Wiley and Sons New YorkNY 385 pp

Pough FH RM Andrews JE Cadle ML Crump AH Savitzky and KDWells 2001 Herpetology 2ndEdition Prentice Hall NJ 612 pp

Russell KR DH Van Lear and DC Guynn Jr 1999 Prescribed fire effectson herpetofauna Review and management implications Wildlife SocietyBulletin 27(2)374384

Savely HE 1939 Ecological relations of certain animals in dead pine and oaklogs Ecological Monographs 9(3)322377

SAS Institute 1997 SASSTAT software Changes and enhancements throughrelease 612 SAS Institute Cary NC

Smith DM BC Larson MJ Kelty and PMS Ashton 1997 The Practice ofSilviculture Applied Forest Ecology Wiley and Sons New York NY 537 pp

Spotila JR 1972 Role of temperature and water in the ecology of lunglesssalamanders Ecological Monographs 42(1)95125

Stoddard HL 1962 Use of fire in pine forests and game lands of the deepSoutheast Proceedings of the Tall Timbers Fire Ecology Conference TallTimbers Research Station Tallahassee FL 13142

Van Lear DH and TA Waldrop 1989 History uses and effects of fire in theAppalachians US Department of Agriculture Forest Service GeneralTechnical Report SE-54 20 pp

Walker LC 1999 The North American Forests Geography Ecology andSilviculture CRC Press Boca Raton FL 398 pp

Whiles MR and JW Grubaugh 1996 Importance of coarse woody debris tosouthern forest herpetofauna Pp 94100 In JW McMinn and DACrossley Jr (Eds) Biodiversity of Coarse Woody Debris in SouthernForests Effects on Biodiversity Proceedings of the Workshop on CoarseWoody Debris in Southern Forests US Department of Agriculture ForestService General Technical Report SE-94 146 pp

Wilson LA 1995 Land Manageriacutes Guide to the Amphibians and Reptiles ofthe SouthThe Nature Conservancy Southeastern Region Chapel Hill NC324 pp

KR Moseley SB Castleberry and SH Schweitzer2003 481

(Table 3) Percent bare ground cover was greater (P = 00029) in burned(2028 ) than unburned (117 ) stands All other habitat variableswere not different (P gt 005) between burned and unburned stands

Significant habitat variables from ANOVA including litter depthcanopy height and percent bare ground cover were examined in correla-tion analyses Abundance of eastern narrowmouth toads (Gastrophrynecarolinensis H) (r = -060 P = 00396) and salamander abundance (r =-062 P = 00303) were negatively correlated with percent bare groundcover No other significant relationships (P gt 005) were observed

DISCUSSION

Our results indicate that periodic winter surface fires in bottomlandhardwood stands of the southeastern Coastal Plain do not affect overallherpetofaunal amphibian salamander or anuran abundance diversityor richness Our findings are similar to those of Ford et al (1999) whofound a high intensity restoration fire in Southern Appalachian hard-wood stands to have a negligible effect on herpetofaunal communitiesbut suggested that alterations in vital microhabitat components due toprescribed fire may negatively impact woodland salamanders On ourstudy site most habitat variables measured were similar between burned

Table 2 Mean (plusmn SD) abundance Shannon-Weiner Index and total number of species foreach taxonomic category (total amphibian salamander anuran and reptile) of herpetofaunacaptured in three burned and three unburned bottomland hardwood stands at Di-LanePlantation Wildlife Management Area Burke County Georgia July-October 2001

Burned Unburned F14

P

Total HerpetofaunaAbundance 4900 plusmn 2346 6600 plusmn 2100 084 04111Shannon-Weiner Index 188 plusmn 026 177 plusmn 014 020 06755Richness 1067 plusmn 306 1033 plusmn 058 013 07344

AmphibiansAbundance 4267 plusmn 2303 6167 plusmn 2079 112 03486Shannon-Weiner Index 148 plusmn 032 154 plusmn 011 012 07466Richness 767 plusmn 208 800 plusmn 0 002 09019

SalamandersAbundance 200 plusmn 068 583 plusmn 114 622 00671Shannon-Weiner Index 021 plusmn 006 035 plusmn 004 649 00635Richness 117 plusmn 031 150 plusmn 022 057 04918

AnuransAbundance 3867 plusmn 2108 5000 plusmn 1609 055 05003Shannon-Weiner Index 124 plusmn 035 117 plusmn 018 011 07581Richness 600 plusmn 173 633 plusmn 058 002 08944

ReptilesAbundance 667 plusmn 058 433 plusmn 058 245 00080a

Shannon-Weiner Index 040 plusmn 006 023 plusmn 005 814 00462a

Richness 300 plusmn 100 233 plusmn 058 080 04216aMeans are significantly different (P le 005)

Southeastern Naturalist Vol 2 No 4482

and unburned stands probably due to the low intensity of burns Of thethree habitat variables that were different between treatments the mostprofound was reduction in litter layer depth which subsequently pro-duced an increase in bare ground cover in burned stands

Decrease in leaf litter depth results in a reduction of on-site mois-ture which can impair dermal respiratory ability in amphibians(Duellman and Trueb 1994) and has been cited as a major factor inreduction of amphibian abundance (Ash 1995 Ash 1997) We capturedalmost twice as many eastern narrowmouth toads in unburned standscompared to burned stands (76 and 42 respectively) and found that

Table 3 Mean (plusmn SD) of habitat variables measured within a 004-ha plot around trapclusters in three burned and three unburned bottomland hardwood stands at Di-LanePlantation Wildlife Management Area Burke County Georgia JulyOctober 2001

Habitat Variable Burned Unburned F14 P

Basal area m2ha (gt 716 cm DBH)Hardwood 2408 plusmn 316 2154 plusmn 343 002 09060

Pine 659 plusmn 221 763 plusmn 343 005 08364Snaga 093 plusmn 040 116 plusmn 058 046 05369Totala 3159 plusmn 446 3032 plusmn 25 026 06369

Midstory stemsha (lt 716 cm DBH)Total stems 134167 plusmn 41966 237083 plusmn 21461 382 01223

Course Woody Debris (m3 ha)

Class 1a 2901 plusmn 1629 2359 plusmn 1437 009 07787Class 2a 588 plusmn 406 4113 plusmn 382 260 01820Class 3a 2507 plusmn 873 6371 plusmn 409 008 07871Totala 5995 plusmn 197 12843 plusmn 7272 002 08921

Canopy height (m) 2114 plusmn 122 2431 plusmn 118 1230 00247b

Midstory height (m) 1339 plusmn 083 1212 plusmn 079 088 04023

Understory (0-155m) stems6m2

Grassa 3700 plusmn 3192 117 plusmn117 108 03582Shruba 7533 plusmn 3357 26 plusmn 1842 505 00880Vine 3033 plusmn 1052 3783 plusmn 1034 020 06785Woody 2583 plusmn 531 3117 plusmn 88 040 05617Other 117 plusmn 075 15 plusmn 096 004 08554Total stems 16967 plusmn 4242 9767 plusmn 1737 433 01059

Understory height (cm) 2641 plusmn 470 259 plusmn 48 001 09294

Percent ground coverPine litter 2436 plusmn 760 1986 plusmn 1027 007 08067Hardwood litter 4917 plusmn 970 7703 plusmn 996 217 02148Bare grounda 2028 plusmn 537 117 plusmn 094 4239 00029b

Woody debrisa 571 plusmn 242 194 plusmn 09 195 02349

Litter depth (cm) 093 plusmn 023 286 plusmn 034 1247 00242b

pH 433 plusmn 011 445 plusmn 016 040 05591

Distance to edge (m) 8650 plusmn 993 1315 plusmn 2155 331 0143

Distance to water (m) 74017 plusmn 18026 122617 plusmn 15381 172 02594aVariable was log10 transformedbMeans are significantly different (P le 005)

KR Moseley SB Castleberry and SH Schweitzer2003 483

abundance was negatively correlated with percent bare ground coverSalamander abundance also was negatively correlated with percent bareground cover Although we did not test for differences of individualspecies between treatments because of low sample sizes our resultssuggest that increases in bare ground cover as a result of leaf litterreductions may be associated with lower abundance of some amphib-ians Other studies have demonstrated similar results where reduction inleaf litter depth was a result of forest harvesting (Ash 1995 Ash 1997Grialou et al 2000 McLeod and Gates 1998 Pough et al 1987) Ourstudy apparently is the first to find such a relationship in bottomlandhardwoods as a result of prescribed fire

Several amphibian species captured possess behavioral characteris-tics important in avoiding desiccation Southern toads (Bufo terrestrisB) eastern spadefoot toads (Scaphiopus holbrooki H) and Americantoads (Bufo americanus Holbrook) together representing 461 and286 of amphibian captures in burned and unburned plots respec-tively are capable of burrowing into moist soil to avoid dry surfaceconditions (Conant and Collins 1998) The ability of these species toescape such conditions by burrowing may enable them to survive instands with a reduced litter layer

Total CWD volume and volume of all CWD decay classes did not differbetween burned and unburned stands Other studies have found decreasesin CWD and consequent decreases in amphibian abundance in areassubjected to prescribed fire (McLeod and Gates 1998) All species cap-tured in our study possess some dependence on CWD (Conant and Collins1998 Harmon et al 1986 Maidens et al 1998 Whiles and Grubaugh1996 Wilson 1995) For example eastern narrowmouth toads represent-ing 328 and 41 of amphibians captured in burned and unburnedstands respectively typically occupy habitats that provide moisture andshelter such as under CWD (Conant and Collins 1998) CWD has arelatively constant moisture regime and can remain moist during long dryperiods providing refuge for amphibians (Boddy 1983) Furthermorearthropods are abundant in and around CWD (Graham 1935 Hanula 1993Harmon et al 1986 Savely 1939) yielding an important food source forinsectivorous herpetofauna These characteristics of CWD may havecompensated to some degree for leaf litter reduction by providing adequateamounts of moisture and food for amphibians

Similar to our results previous studies examining fire effects onherpetofauna in southeastern Coastal Plain pine communities also foundreptiles to increase in diversity and abundance following application ofprescribed fire (Means and Campbell 1981 Mushinsky 1985) attributingthe increases to greater thermoregulatory opportunity within burnedstands Increased bare ground cover in burned stands resulted in a largerground surface area for thermoregulatory activity by reptiles When solar

Southeastern Naturalist Vol 2 No 4484

radiation declines in the evening lizards will leave their perches to flattenout against the warm sand to maintain optimal temperature (Pough et al2001) Many species of reptiles also utilize CWD as a basking substrate aswell as for shelter and nesting (Conant and Collins 1998 Pough et al 2001Wilson 1995) Numerous lizards were observed basking on logs at the edgeof burned stands during our study Five-lined skinks (Eumeces fasciatusL) representing 579 and 385 of reptile species captured in burnedand unburned stands respectively frequently utilize CWD as nesting sitesbecause of the relatively constant microclimate provided for egg develop-ment (Hecnar 1994) We recognize that the observed increase in reptilediversity and abundance in our study may be an underestimate because ourtrapping methods are relatively ineffective at catching large snakes whichwere likely underrepresented in our collections

We conclude that the use of low intensity prescribed fire had littleoverall effect on herpetofauna in our bottomland hardwood forest studyarea However reductions in leaf litter and associated increases in bareground cover may be associated with lower abundance of some amphib-ians Conversely the increased ground surface area available for ther-moregulatory activity may benefit some reptiles CWD was relativelyunaffected by the burning regime probably because of the low intensityof the fires In areas where fire intensity is greater the impacts on theherpetofaunal community may be more pronounced

ACKNOWLEDGMENTS

We thank V VanSant H Barnhill and J Bearden of the Georgia Depart-ment of Natural Resources Wildlife Resources Division for their assistance andaccess to Di-Lane WMA We also thank K Francl for GIS assistance

LITERATURE CITED

Ash AN 1995 Effects of clear-cutting on litter parameters in the SouthernBlue Ridge mountains Castanea 60(2)8997

Ash AN 1997 Disappearance and return of Plethodontid salamanders toclearcut plots in the southern Blue Ridge mountains Conservation Biology11(4)983989

Bennett SH JW Gibbons and J Glanville 1980 Terrestrial activity abun-dance and diversity of amphibians in differently managed forest typesAmerican Midland Naturalist 103(2)412416

Boddy L 1983 Microclimate and moisture dynamics of wood decomposing interrestrial ecosystems Soil Biology and Biochemistry 15(2)149157

Boughton RG J Staiger and R Franz 2000 Use of PVC refugia as asampling technique for hylid treefrogs American Midland Naturalist144(1)168177

Conant R and JT Collins 1998 A Field Guide to Reptiles and Amphibians ofEasternCentral North America 3rd Edition Houghton Mifflin Co BostonMA 616 pp

KR Moseley SB Castleberry and SH Schweitzer2003 485

Corn PS 1994 Straight line drift fences and pitfall traps Pp 109117 InWR Heyer MA Donnelly RW McDiarmid LC Hayek and MSFoster (Eds) Measuring and Monitoring Biological Diversity StandardMethods for Amphibians Smithsonian Institute Press Washington DC364 pp

DeMaynadier PG and ML Hunter Jr 1995 The relationship between forestmanagement and amphibian ecology A review of the North Americanliterature Environmental Reviews 3(3-4)230261

Duellman WE and L Trueb 1994 Biology of Amphibians The JohnsHopkins University Press Baltimore MD 670 pp

Environmental Systems Research Institute 1999 ARCVIEW Version 32Environmental Systems Research Institute Redlands CA

Fellers GM and CA Drost 1994 Sampling with artificial cover Pp 146-150 In WR Heyer MA Donnelly RW McDiarmid LC Hayek andMS Foster (Eds) Measuring and Monitoring Biological Diversity Stan-dard Methods for Amphibians Smithsonian Institute Press WashingtonDC 364 pp

Ford WM MA Menzel DW McGill J Laerm and TS McCay 1999Effects of a community restoration fire on small mammals and herpetofaunain the Southern Appalachians Forest Ecology and Management 114(2-3)233243

Graham SA 1935 The felled tree trunk as an ecological unit Ecology6(4)397412

Greenberg CH DG Neary and LD Harris 1994 Effect of high-intensitywildfire and silvicultural treatments on reptile communities in sand-pinescrub Conservation Biology 8(4)10471057

Grialou JA SD West and RN Wilkins 2000 The effects of forest clearcutharvesting and thinning on terrestrial salamanders Journal of Wildlife Man-agement 64(1)105113

Hanula JL 1993 Relationship of wood-feeding insects and coarse woodydebris Pp 5565 In JW McMinn and DA Crossley Jr (Eds)Biodiversity of Coarse Woody Debris in Southern Forests Effects onBiodiversity Proceedings of the Workshop on Coarse Woody Debris inSouthern Forests US Department of Agriculture Forest Service GeneralTechnical Report SE-94 146 pp

Harmon ME JF Franklin FJ Swanson P Sollins SV Gregory JDLattin NH Anderson SP Cline NG Aumen JR Sedell GWLienkaemper K Cromack Jr KW Cummins 1986 Ecology of coarsewoody debris in temperate ecosystems Advances in Ecological Research15133302

Hecnar SJ 1994 Nest distribution site selection and brooding in the five-linedskink (Eumeces fasciatus) Canadian Journal of Zoology 72(8)15101516

Maidens DA MA Menzel and J Laerm 1998 Notes on the effect of sizeand level of decay of coarse woody debris on relative abundance of shrewsand salamanders in the southern Appalachian mountains Georgia Journal ofScience 56(4)226-233

McLeod RF and JE Gates 1998 Response of herpetofaunal communities toforest cutting and burning at Chesapeake Farms Maryland American Mid-land Naturalist 139(1)164177

Southeastern Naturalist Vol 2 No 4486

Means DB and HW Campbell 1981 Effects of prescribed burningonamphibians and reptiles Pp 8996 In GW Wood (Ed) Prescribed Fireand Wildlife in Southern Forests Belle Baruch Forest Science InstituteClemson University Georgetown SC 170 pp

Mushinsky HR 1985 Fire and the Florida sandhill herptofaunal communitywith special attention to responses of Cnemidophorus sexlineatusHerpetologica 41(3)333342

Nyland RD 1996 Silviculture Concepts and Applications McGraw-HillNew York NY 633 pp

Pielou EC 1977 Mathematical Ecology John Wiley and Sons New YorkNY 385 pp

Pough FH RM Andrews JE Cadle ML Crump AH Savitzky and KDWells 2001 Herpetology 2ndEdition Prentice Hall NJ 612 pp

Russell KR DH Van Lear and DC Guynn Jr 1999 Prescribed fire effectson herpetofauna Review and management implications Wildlife SocietyBulletin 27(2)374384

Savely HE 1939 Ecological relations of certain animals in dead pine and oaklogs Ecological Monographs 9(3)322377

SAS Institute 1997 SASSTAT software Changes and enhancements throughrelease 612 SAS Institute Cary NC

Smith DM BC Larson MJ Kelty and PMS Ashton 1997 The Practice ofSilviculture Applied Forest Ecology Wiley and Sons New York NY 537 pp

Spotila JR 1972 Role of temperature and water in the ecology of lunglesssalamanders Ecological Monographs 42(1)95125

Stoddard HL 1962 Use of fire in pine forests and game lands of the deepSoutheast Proceedings of the Tall Timbers Fire Ecology Conference TallTimbers Research Station Tallahassee FL 13142

Van Lear DH and TA Waldrop 1989 History uses and effects of fire in theAppalachians US Department of Agriculture Forest Service GeneralTechnical Report SE-54 20 pp

Walker LC 1999 The North American Forests Geography Ecology andSilviculture CRC Press Boca Raton FL 398 pp

Whiles MR and JW Grubaugh 1996 Importance of coarse woody debris tosouthern forest herpetofauna Pp 94100 In JW McMinn and DACrossley Jr (Eds) Biodiversity of Coarse Woody Debris in SouthernForests Effects on Biodiversity Proceedings of the Workshop on CoarseWoody Debris in Southern Forests US Department of Agriculture ForestService General Technical Report SE-94 146 pp

Wilson LA 1995 Land Manageriacutes Guide to the Amphibians and Reptiles ofthe SouthThe Nature Conservancy Southeastern Region Chapel Hill NC324 pp

Southeastern Naturalist Vol 2 No 4482

and unburned stands probably due to the low intensity of burns Of thethree habitat variables that were different between treatments the mostprofound was reduction in litter layer depth which subsequently pro-duced an increase in bare ground cover in burned stands

Decrease in leaf litter depth results in a reduction of on-site mois-ture which can impair dermal respiratory ability in amphibians(Duellman and Trueb 1994) and has been cited as a major factor inreduction of amphibian abundance (Ash 1995 Ash 1997) We capturedalmost twice as many eastern narrowmouth toads in unburned standscompared to burned stands (76 and 42 respectively) and found that

Table 3 Mean (plusmn SD) of habitat variables measured within a 004-ha plot around trapclusters in three burned and three unburned bottomland hardwood stands at Di-LanePlantation Wildlife Management Area Burke County Georgia JulyOctober 2001

Habitat Variable Burned Unburned F14 P

Basal area m2ha (gt 716 cm DBH)Hardwood 2408 plusmn 316 2154 plusmn 343 002 09060

Pine 659 plusmn 221 763 plusmn 343 005 08364Snaga 093 plusmn 040 116 plusmn 058 046 05369Totala 3159 plusmn 446 3032 plusmn 25 026 06369

Midstory stemsha (lt 716 cm DBH)Total stems 134167 plusmn 41966 237083 plusmn 21461 382 01223

Course Woody Debris (m3 ha)

Class 1a 2901 plusmn 1629 2359 plusmn 1437 009 07787Class 2a 588 plusmn 406 4113 plusmn 382 260 01820Class 3a 2507 plusmn 873 6371 plusmn 409 008 07871Totala 5995 plusmn 197 12843 plusmn 7272 002 08921

Canopy height (m) 2114 plusmn 122 2431 plusmn 118 1230 00247b

Midstory height (m) 1339 plusmn 083 1212 plusmn 079 088 04023

Understory (0-155m) stems6m2

Grassa 3700 plusmn 3192 117 plusmn117 108 03582Shruba 7533 plusmn 3357 26 plusmn 1842 505 00880Vine 3033 plusmn 1052 3783 plusmn 1034 020 06785Woody 2583 plusmn 531 3117 plusmn 88 040 05617Other 117 plusmn 075 15 plusmn 096 004 08554Total stems 16967 plusmn 4242 9767 plusmn 1737 433 01059

Understory height (cm) 2641 plusmn 470 259 plusmn 48 001 09294

Percent ground coverPine litter 2436 plusmn 760 1986 plusmn 1027 007 08067Hardwood litter 4917 plusmn 970 7703 plusmn 996 217 02148Bare grounda 2028 plusmn 537 117 plusmn 094 4239 00029b

Woody debrisa 571 plusmn 242 194 plusmn 09 195 02349

Litter depth (cm) 093 plusmn 023 286 plusmn 034 1247 00242b

pH 433 plusmn 011 445 plusmn 016 040 05591

Distance to edge (m) 8650 plusmn 993 1315 plusmn 2155 331 0143

Distance to water (m) 74017 plusmn 18026 122617 plusmn 15381 172 02594aVariable was log10 transformedbMeans are significantly different (P le 005)

KR Moseley SB Castleberry and SH Schweitzer2003 483

abundance was negatively correlated with percent bare ground coverSalamander abundance also was negatively correlated with percent bareground cover Although we did not test for differences of individualspecies between treatments because of low sample sizes our resultssuggest that increases in bare ground cover as a result of leaf litterreductions may be associated with lower abundance of some amphib-ians Other studies have demonstrated similar results where reduction inleaf litter depth was a result of forest harvesting (Ash 1995 Ash 1997Grialou et al 2000 McLeod and Gates 1998 Pough et al 1987) Ourstudy apparently is the first to find such a relationship in bottomlandhardwoods as a result of prescribed fire

Several amphibian species captured possess behavioral characteris-tics important in avoiding desiccation Southern toads (Bufo terrestrisB) eastern spadefoot toads (Scaphiopus holbrooki H) and Americantoads (Bufo americanus Holbrook) together representing 461 and286 of amphibian captures in burned and unburned plots respec-tively are capable of burrowing into moist soil to avoid dry surfaceconditions (Conant and Collins 1998) The ability of these species toescape such conditions by burrowing may enable them to survive instands with a reduced litter layer

Total CWD volume and volume of all CWD decay classes did not differbetween burned and unburned stands Other studies have found decreasesin CWD and consequent decreases in amphibian abundance in areassubjected to prescribed fire (McLeod and Gates 1998) All species cap-tured in our study possess some dependence on CWD (Conant and Collins1998 Harmon et al 1986 Maidens et al 1998 Whiles and Grubaugh1996 Wilson 1995) For example eastern narrowmouth toads represent-ing 328 and 41 of amphibians captured in burned and unburnedstands respectively typically occupy habitats that provide moisture andshelter such as under CWD (Conant and Collins 1998) CWD has arelatively constant moisture regime and can remain moist during long dryperiods providing refuge for amphibians (Boddy 1983) Furthermorearthropods are abundant in and around CWD (Graham 1935 Hanula 1993Harmon et al 1986 Savely 1939) yielding an important food source forinsectivorous herpetofauna These characteristics of CWD may havecompensated to some degree for leaf litter reduction by providing adequateamounts of moisture and food for amphibians

Similar to our results previous studies examining fire effects onherpetofauna in southeastern Coastal Plain pine communities also foundreptiles to increase in diversity and abundance following application ofprescribed fire (Means and Campbell 1981 Mushinsky 1985) attributingthe increases to greater thermoregulatory opportunity within burnedstands Increased bare ground cover in burned stands resulted in a largerground surface area for thermoregulatory activity by reptiles When solar

Southeastern Naturalist Vol 2 No 4484

radiation declines in the evening lizards will leave their perches to flattenout against the warm sand to maintain optimal temperature (Pough et al2001) Many species of reptiles also utilize CWD as a basking substrate aswell as for shelter and nesting (Conant and Collins 1998 Pough et al 2001Wilson 1995) Numerous lizards were observed basking on logs at the edgeof burned stands during our study Five-lined skinks (Eumeces fasciatusL) representing 579 and 385 of reptile species captured in burnedand unburned stands respectively frequently utilize CWD as nesting sitesbecause of the relatively constant microclimate provided for egg develop-ment (Hecnar 1994) We recognize that the observed increase in reptilediversity and abundance in our study may be an underestimate because ourtrapping methods are relatively ineffective at catching large snakes whichwere likely underrepresented in our collections

We conclude that the use of low intensity prescribed fire had littleoverall effect on herpetofauna in our bottomland hardwood forest studyarea However reductions in leaf litter and associated increases in bareground cover may be associated with lower abundance of some amphib-ians Conversely the increased ground surface area available for ther-moregulatory activity may benefit some reptiles CWD was relativelyunaffected by the burning regime probably because of the low intensityof the fires In areas where fire intensity is greater the impacts on theherpetofaunal community may be more pronounced

ACKNOWLEDGMENTS

We thank V VanSant H Barnhill and J Bearden of the Georgia Depart-ment of Natural Resources Wildlife Resources Division for their assistance andaccess to Di-Lane WMA We also thank K Francl for GIS assistance

LITERATURE CITED

Ash AN 1995 Effects of clear-cutting on litter parameters in the SouthernBlue Ridge mountains Castanea 60(2)8997

Ash AN 1997 Disappearance and return of Plethodontid salamanders toclearcut plots in the southern Blue Ridge mountains Conservation Biology11(4)983989

Bennett SH JW Gibbons and J Glanville 1980 Terrestrial activity abun-dance and diversity of amphibians in differently managed forest typesAmerican Midland Naturalist 103(2)412416

Boddy L 1983 Microclimate and moisture dynamics of wood decomposing interrestrial ecosystems Soil Biology and Biochemistry 15(2)149157

Boughton RG J Staiger and R Franz 2000 Use of PVC refugia as asampling technique for hylid treefrogs American Midland Naturalist144(1)168177

Conant R and JT Collins 1998 A Field Guide to Reptiles and Amphibians ofEasternCentral North America 3rd Edition Houghton Mifflin Co BostonMA 616 pp

KR Moseley SB Castleberry and SH Schweitzer2003 485

Corn PS 1994 Straight line drift fences and pitfall traps Pp 109117 InWR Heyer MA Donnelly RW McDiarmid LC Hayek and MSFoster (Eds) Measuring and Monitoring Biological Diversity StandardMethods for Amphibians Smithsonian Institute Press Washington DC364 pp

DeMaynadier PG and ML Hunter Jr 1995 The relationship between forestmanagement and amphibian ecology A review of the North Americanliterature Environmental Reviews 3(3-4)230261

Duellman WE and L Trueb 1994 Biology of Amphibians The JohnsHopkins University Press Baltimore MD 670 pp

Environmental Systems Research Institute 1999 ARCVIEW Version 32Environmental Systems Research Institute Redlands CA

Fellers GM and CA Drost 1994 Sampling with artificial cover Pp 146-150 In WR Heyer MA Donnelly RW McDiarmid LC Hayek andMS Foster (Eds) Measuring and Monitoring Biological Diversity Stan-dard Methods for Amphibians Smithsonian Institute Press WashingtonDC 364 pp

Ford WM MA Menzel DW McGill J Laerm and TS McCay 1999Effects of a community restoration fire on small mammals and herpetofaunain the Southern Appalachians Forest Ecology and Management 114(2-3)233243

Graham SA 1935 The felled tree trunk as an ecological unit Ecology6(4)397412

Greenberg CH DG Neary and LD Harris 1994 Effect of high-intensitywildfire and silvicultural treatments on reptile communities in sand-pinescrub Conservation Biology 8(4)10471057

Grialou JA SD West and RN Wilkins 2000 The effects of forest clearcutharvesting and thinning on terrestrial salamanders Journal of Wildlife Man-agement 64(1)105113

Hanula JL 1993 Relationship of wood-feeding insects and coarse woodydebris Pp 5565 In JW McMinn and DA Crossley Jr (Eds)Biodiversity of Coarse Woody Debris in Southern Forests Effects onBiodiversity Proceedings of the Workshop on Coarse Woody Debris inSouthern Forests US Department of Agriculture Forest Service GeneralTechnical Report SE-94 146 pp

Harmon ME JF Franklin FJ Swanson P Sollins SV Gregory JDLattin NH Anderson SP Cline NG Aumen JR Sedell GWLienkaemper K Cromack Jr KW Cummins 1986 Ecology of coarsewoody debris in temperate ecosystems Advances in Ecological Research15133302

Hecnar SJ 1994 Nest distribution site selection and brooding in the five-linedskink (Eumeces fasciatus) Canadian Journal of Zoology 72(8)15101516

Maidens DA MA Menzel and J Laerm 1998 Notes on the effect of sizeand level of decay of coarse woody debris on relative abundance of shrewsand salamanders in the southern Appalachian mountains Georgia Journal ofScience 56(4)226-233

McLeod RF and JE Gates 1998 Response of herpetofaunal communities toforest cutting and burning at Chesapeake Farms Maryland American Mid-land Naturalist 139(1)164177

Southeastern Naturalist Vol 2 No 4486

Means DB and HW Campbell 1981 Effects of prescribed burningonamphibians and reptiles Pp 8996 In GW Wood (Ed) Prescribed Fireand Wildlife in Southern Forests Belle Baruch Forest Science InstituteClemson University Georgetown SC 170 pp

Mushinsky HR 1985 Fire and the Florida sandhill herptofaunal communitywith special attention to responses of Cnemidophorus sexlineatusHerpetologica 41(3)333342

Nyland RD 1996 Silviculture Concepts and Applications McGraw-HillNew York NY 633 pp

Pielou EC 1977 Mathematical Ecology John Wiley and Sons New YorkNY 385 pp

Pough FH RM Andrews JE Cadle ML Crump AH Savitzky and KDWells 2001 Herpetology 2ndEdition Prentice Hall NJ 612 pp

Russell KR DH Van Lear and DC Guynn Jr 1999 Prescribed fire effectson herpetofauna Review and management implications Wildlife SocietyBulletin 27(2)374384

Savely HE 1939 Ecological relations of certain animals in dead pine and oaklogs Ecological Monographs 9(3)322377

SAS Institute 1997 SASSTAT software Changes and enhancements throughrelease 612 SAS Institute Cary NC

Smith DM BC Larson MJ Kelty and PMS Ashton 1997 The Practice ofSilviculture Applied Forest Ecology Wiley and Sons New York NY 537 pp

Spotila JR 1972 Role of temperature and water in the ecology of lunglesssalamanders Ecological Monographs 42(1)95125

Stoddard HL 1962 Use of fire in pine forests and game lands of the deepSoutheast Proceedings of the Tall Timbers Fire Ecology Conference TallTimbers Research Station Tallahassee FL 13142

Van Lear DH and TA Waldrop 1989 History uses and effects of fire in theAppalachians US Department of Agriculture Forest Service GeneralTechnical Report SE-54 20 pp

Walker LC 1999 The North American Forests Geography Ecology andSilviculture CRC Press Boca Raton FL 398 pp

Whiles MR and JW Grubaugh 1996 Importance of coarse woody debris tosouthern forest herpetofauna Pp 94100 In JW McMinn and DACrossley Jr (Eds) Biodiversity of Coarse Woody Debris in SouthernForests Effects on Biodiversity Proceedings of the Workshop on CoarseWoody Debris in Southern Forests US Department of Agriculture ForestService General Technical Report SE-94 146 pp

Wilson LA 1995 Land Manageriacutes Guide to the Amphibians and Reptiles ofthe SouthThe Nature Conservancy Southeastern Region Chapel Hill NC324 pp

KR Moseley SB Castleberry and SH Schweitzer2003 483

abundance was negatively correlated with percent bare ground coverSalamander abundance also was negatively correlated with percent bareground cover Although we did not test for differences of individualspecies between treatments because of low sample sizes our resultssuggest that increases in bare ground cover as a result of leaf litterreductions may be associated with lower abundance of some amphib-ians Other studies have demonstrated similar results where reduction inleaf litter depth was a result of forest harvesting (Ash 1995 Ash 1997Grialou et al 2000 McLeod and Gates 1998 Pough et al 1987) Ourstudy apparently is the first to find such a relationship in bottomlandhardwoods as a result of prescribed fire

Several amphibian species captured possess behavioral characteris-tics important in avoiding desiccation Southern toads (Bufo terrestrisB) eastern spadefoot toads (Scaphiopus holbrooki H) and Americantoads (Bufo americanus Holbrook) together representing 461 and286 of amphibian captures in burned and unburned plots respec-tively are capable of burrowing into moist soil to avoid dry surfaceconditions (Conant and Collins 1998) The ability of these species toescape such conditions by burrowing may enable them to survive instands with a reduced litter layer

Total CWD volume and volume of all CWD decay classes did not differbetween burned and unburned stands Other studies have found decreasesin CWD and consequent decreases in amphibian abundance in areassubjected to prescribed fire (McLeod and Gates 1998) All species cap-tured in our study possess some dependence on CWD (Conant and Collins1998 Harmon et al 1986 Maidens et al 1998 Whiles and Grubaugh1996 Wilson 1995) For example eastern narrowmouth toads represent-ing 328 and 41 of amphibians captured in burned and unburnedstands respectively typically occupy habitats that provide moisture andshelter such as under CWD (Conant and Collins 1998) CWD has arelatively constant moisture regime and can remain moist during long dryperiods providing refuge for amphibians (Boddy 1983) Furthermorearthropods are abundant in and around CWD (Graham 1935 Hanula 1993Harmon et al 1986 Savely 1939) yielding an important food source forinsectivorous herpetofauna These characteristics of CWD may havecompensated to some degree for leaf litter reduction by providing adequateamounts of moisture and food for amphibians

Similar to our results previous studies examining fire effects onherpetofauna in southeastern Coastal Plain pine communities also foundreptiles to increase in diversity and abundance following application ofprescribed fire (Means and Campbell 1981 Mushinsky 1985) attributingthe increases to greater thermoregulatory opportunity within burnedstands Increased bare ground cover in burned stands resulted in a largerground surface area for thermoregulatory activity by reptiles When solar

Southeastern Naturalist Vol 2 No 4484

radiation declines in the evening lizards will leave their perches to flattenout against the warm sand to maintain optimal temperature (Pough et al2001) Many species of reptiles also utilize CWD as a basking substrate aswell as for shelter and nesting (Conant and Collins 1998 Pough et al 2001Wilson 1995) Numerous lizards were observed basking on logs at the edgeof burned stands during our study Five-lined skinks (Eumeces fasciatusL) representing 579 and 385 of reptile species captured in burnedand unburned stands respectively frequently utilize CWD as nesting sitesbecause of the relatively constant microclimate provided for egg develop-ment (Hecnar 1994) We recognize that the observed increase in reptilediversity and abundance in our study may be an underestimate because ourtrapping methods are relatively ineffective at catching large snakes whichwere likely underrepresented in our collections

We conclude that the use of low intensity prescribed fire had littleoverall effect on herpetofauna in our bottomland hardwood forest studyarea However reductions in leaf litter and associated increases in bareground cover may be associated with lower abundance of some amphib-ians Conversely the increased ground surface area available for ther-moregulatory activity may benefit some reptiles CWD was relativelyunaffected by the burning regime probably because of the low intensityof the fires In areas where fire intensity is greater the impacts on theherpetofaunal community may be more pronounced

ACKNOWLEDGMENTS

We thank V VanSant H Barnhill and J Bearden of the Georgia Depart-ment of Natural Resources Wildlife Resources Division for their assistance andaccess to Di-Lane WMA We also thank K Francl for GIS assistance

LITERATURE CITED

Ash AN 1995 Effects of clear-cutting on litter parameters in the SouthernBlue Ridge mountains Castanea 60(2)8997

Ash AN 1997 Disappearance and return of Plethodontid salamanders toclearcut plots in the southern Blue Ridge mountains Conservation Biology11(4)983989

Bennett SH JW Gibbons and J Glanville 1980 Terrestrial activity abun-dance and diversity of amphibians in differently managed forest typesAmerican Midland Naturalist 103(2)412416

Boddy L 1983 Microclimate and moisture dynamics of wood decomposing interrestrial ecosystems Soil Biology and Biochemistry 15(2)149157

Boughton RG J Staiger and R Franz 2000 Use of PVC refugia as asampling technique for hylid treefrogs American Midland Naturalist144(1)168177

Conant R and JT Collins 1998 A Field Guide to Reptiles and Amphibians ofEasternCentral North America 3rd Edition Houghton Mifflin Co BostonMA 616 pp

KR Moseley SB Castleberry and SH Schweitzer2003 485

Corn PS 1994 Straight line drift fences and pitfall traps Pp 109117 InWR Heyer MA Donnelly RW McDiarmid LC Hayek and MSFoster (Eds) Measuring and Monitoring Biological Diversity StandardMethods for Amphibians Smithsonian Institute Press Washington DC364 pp

DeMaynadier PG and ML Hunter Jr 1995 The relationship between forestmanagement and amphibian ecology A review of the North Americanliterature Environmental Reviews 3(3-4)230261

Duellman WE and L Trueb 1994 Biology of Amphibians The JohnsHopkins University Press Baltimore MD 670 pp

Environmental Systems Research Institute 1999 ARCVIEW Version 32Environmental Systems Research Institute Redlands CA

Fellers GM and CA Drost 1994 Sampling with artificial cover Pp 146-150 In WR Heyer MA Donnelly RW McDiarmid LC Hayek andMS Foster (Eds) Measuring and Monitoring Biological Diversity Stan-dard Methods for Amphibians Smithsonian Institute Press WashingtonDC 364 pp

Ford WM MA Menzel DW McGill J Laerm and TS McCay 1999Effects of a community restoration fire on small mammals and herpetofaunain the Southern Appalachians Forest Ecology and Management 114(2-3)233243

Graham SA 1935 The felled tree trunk as an ecological unit Ecology6(4)397412

Greenberg CH DG Neary and LD Harris 1994 Effect of high-intensitywildfire and silvicultural treatments on reptile communities in sand-pinescrub Conservation Biology 8(4)10471057

Grialou JA SD West and RN Wilkins 2000 The effects of forest clearcutharvesting and thinning on terrestrial salamanders Journal of Wildlife Man-agement 64(1)105113

Hanula JL 1993 Relationship of wood-feeding insects and coarse woodydebris Pp 5565 In JW McMinn and DA Crossley Jr (Eds)Biodiversity of Coarse Woody Debris in Southern Forests Effects onBiodiversity Proceedings of the Workshop on Coarse Woody Debris inSouthern Forests US Department of Agriculture Forest Service GeneralTechnical Report SE-94 146 pp

Harmon ME JF Franklin FJ Swanson P Sollins SV Gregory JDLattin NH Anderson SP Cline NG Aumen JR Sedell GWLienkaemper K Cromack Jr KW Cummins 1986 Ecology of coarsewoody debris in temperate ecosystems Advances in Ecological Research15133302

Hecnar SJ 1994 Nest distribution site selection and brooding in the five-linedskink (Eumeces fasciatus) Canadian Journal of Zoology 72(8)15101516

Maidens DA MA Menzel and J Laerm 1998 Notes on the effect of sizeand level of decay of coarse woody debris on relative abundance of shrewsand salamanders in the southern Appalachian mountains Georgia Journal ofScience 56(4)226-233

McLeod RF and JE Gates 1998 Response of herpetofaunal communities toforest cutting and burning at Chesapeake Farms Maryland American Mid-land Naturalist 139(1)164177

Southeastern Naturalist Vol 2 No 4486

Means DB and HW Campbell 1981 Effects of prescribed burningonamphibians and reptiles Pp 8996 In GW Wood (Ed) Prescribed Fireand Wildlife in Southern Forests Belle Baruch Forest Science InstituteClemson University Georgetown SC 170 pp

Mushinsky HR 1985 Fire and the Florida sandhill herptofaunal communitywith special attention to responses of Cnemidophorus sexlineatusHerpetologica 41(3)333342

Nyland RD 1996 Silviculture Concepts and Applications McGraw-HillNew York NY 633 pp

Pielou EC 1977 Mathematical Ecology John Wiley and Sons New YorkNY 385 pp

Pough FH RM Andrews JE Cadle ML Crump AH Savitzky and KDWells 2001 Herpetology 2ndEdition Prentice Hall NJ 612 pp

Russell KR DH Van Lear and DC Guynn Jr 1999 Prescribed fire effectson herpetofauna Review and management implications Wildlife SocietyBulletin 27(2)374384

Savely HE 1939 Ecological relations of certain animals in dead pine and oaklogs Ecological Monographs 9(3)322377

SAS Institute 1997 SASSTAT software Changes and enhancements throughrelease 612 SAS Institute Cary NC

Smith DM BC Larson MJ Kelty and PMS Ashton 1997 The Practice ofSilviculture Applied Forest Ecology Wiley and Sons New York NY 537 pp

Spotila JR 1972 Role of temperature and water in the ecology of lunglesssalamanders Ecological Monographs 42(1)95125

Stoddard HL 1962 Use of fire in pine forests and game lands of the deepSoutheast Proceedings of the Tall Timbers Fire Ecology Conference TallTimbers Research Station Tallahassee FL 13142

Van Lear DH and TA Waldrop 1989 History uses and effects of fire in theAppalachians US Department of Agriculture Forest Service GeneralTechnical Report SE-54 20 pp

Walker LC 1999 The North American Forests Geography Ecology andSilviculture CRC Press Boca Raton FL 398 pp

Whiles MR and JW Grubaugh 1996 Importance of coarse woody debris tosouthern forest herpetofauna Pp 94100 In JW McMinn and DACrossley Jr (Eds) Biodiversity of Coarse Woody Debris in SouthernForests Effects on Biodiversity Proceedings of the Workshop on CoarseWoody Debris in Southern Forests US Department of Agriculture ForestService General Technical Report SE-94 146 pp

Wilson LA 1995 Land Manageriacutes Guide to the Amphibians and Reptiles ofthe SouthThe Nature Conservancy Southeastern Region Chapel Hill NC324 pp

Southeastern Naturalist Vol 2 No 4484

radiation declines in the evening lizards will leave their perches to flattenout against the warm sand to maintain optimal temperature (Pough et al2001) Many species of reptiles also utilize CWD as a basking substrate aswell as for shelter and nesting (Conant and Collins 1998 Pough et al 2001Wilson 1995) Numerous lizards were observed basking on logs at the edgeof burned stands during our study Five-lined skinks (Eumeces fasciatusL) representing 579 and 385 of reptile species captured in burnedand unburned stands respectively frequently utilize CWD as nesting sitesbecause of the relatively constant microclimate provided for egg develop-ment (Hecnar 1994) We recognize that the observed increase in reptilediversity and abundance in our study may be an underestimate because ourtrapping methods are relatively ineffective at catching large snakes whichwere likely underrepresented in our collections

We conclude that the use of low intensity prescribed fire had littleoverall effect on herpetofauna in our bottomland hardwood forest studyarea However reductions in leaf litter and associated increases in bareground cover may be associated with lower abundance of some amphib-ians Conversely the increased ground surface area available for ther-moregulatory activity may benefit some reptiles CWD was relativelyunaffected by the burning regime probably because of the low intensityof the fires In areas where fire intensity is greater the impacts on theherpetofaunal community may be more pronounced

ACKNOWLEDGMENTS

We thank V VanSant H Barnhill and J Bearden of the Georgia Depart-ment of Natural Resources Wildlife Resources Division for their assistance andaccess to Di-Lane WMA We also thank K Francl for GIS assistance

LITERATURE CITED

Ash AN 1995 Effects of clear-cutting on litter parameters in the SouthernBlue Ridge mountains Castanea 60(2)8997

Ash AN 1997 Disappearance and return of Plethodontid salamanders toclearcut plots in the southern Blue Ridge mountains Conservation Biology11(4)983989

Bennett SH JW Gibbons and J Glanville 1980 Terrestrial activity abun-dance and diversity of amphibians in differently managed forest typesAmerican Midland Naturalist 103(2)412416

Boddy L 1983 Microclimate and moisture dynamics of wood decomposing interrestrial ecosystems Soil Biology and Biochemistry 15(2)149157

Boughton RG J Staiger and R Franz 2000 Use of PVC refugia as asampling technique for hylid treefrogs American Midland Naturalist144(1)168177

Conant R and JT Collins 1998 A Field Guide to Reptiles and Amphibians ofEasternCentral North America 3rd Edition Houghton Mifflin Co BostonMA 616 pp

KR Moseley SB Castleberry and SH Schweitzer2003 485

Corn PS 1994 Straight line drift fences and pitfall traps Pp 109117 InWR Heyer MA Donnelly RW McDiarmid LC Hayek and MSFoster (Eds) Measuring and Monitoring Biological Diversity StandardMethods for Amphibians Smithsonian Institute Press Washington DC364 pp

DeMaynadier PG and ML Hunter Jr 1995 The relationship between forestmanagement and amphibian ecology A review of the North Americanliterature Environmental Reviews 3(3-4)230261

Duellman WE and L Trueb 1994 Biology of Amphibians The JohnsHopkins University Press Baltimore MD 670 pp

Environmental Systems Research Institute 1999 ARCVIEW Version 32Environmental Systems Research Institute Redlands CA

Fellers GM and CA Drost 1994 Sampling with artificial cover Pp 146-150 In WR Heyer MA Donnelly RW McDiarmid LC Hayek andMS Foster (Eds) Measuring and Monitoring Biological Diversity Stan-dard Methods for Amphibians Smithsonian Institute Press WashingtonDC 364 pp

Ford WM MA Menzel DW McGill J Laerm and TS McCay 1999Effects of a community restoration fire on small mammals and herpetofaunain the Southern Appalachians Forest Ecology and Management 114(2-3)233243

Graham SA 1935 The felled tree trunk as an ecological unit Ecology6(4)397412

Greenberg CH DG Neary and LD Harris 1994 Effect of high-intensitywildfire and silvicultural treatments on reptile communities in sand-pinescrub Conservation Biology 8(4)10471057

Grialou JA SD West and RN Wilkins 2000 The effects of forest clearcutharvesting and thinning on terrestrial salamanders Journal of Wildlife Man-agement 64(1)105113

Hanula JL 1993 Relationship of wood-feeding insects and coarse woodydebris Pp 5565 In JW McMinn and DA Crossley Jr (Eds)Biodiversity of Coarse Woody Debris in Southern Forests Effects onBiodiversity Proceedings of the Workshop on Coarse Woody Debris inSouthern Forests US Department of Agriculture Forest Service GeneralTechnical Report SE-94 146 pp

Harmon ME JF Franklin FJ Swanson P Sollins SV Gregory JDLattin NH Anderson SP Cline NG Aumen JR Sedell GWLienkaemper K Cromack Jr KW Cummins 1986 Ecology of coarsewoody debris in temperate ecosystems Advances in Ecological Research15133302

Hecnar SJ 1994 Nest distribution site selection and brooding in the five-linedskink (Eumeces fasciatus) Canadian Journal of Zoology 72(8)15101516

Maidens DA MA Menzel and J Laerm 1998 Notes on the effect of sizeand level of decay of coarse woody debris on relative abundance of shrewsand salamanders in the southern Appalachian mountains Georgia Journal ofScience 56(4)226-233

McLeod RF and JE Gates 1998 Response of herpetofaunal communities toforest cutting and burning at Chesapeake Farms Maryland American Mid-land Naturalist 139(1)164177

Southeastern Naturalist Vol 2 No 4486

Means DB and HW Campbell 1981 Effects of prescribed burningonamphibians and reptiles Pp 8996 In GW Wood (Ed) Prescribed Fireand Wildlife in Southern Forests Belle Baruch Forest Science InstituteClemson University Georgetown SC 170 pp

Mushinsky HR 1985 Fire and the Florida sandhill herptofaunal communitywith special attention to responses of Cnemidophorus sexlineatusHerpetologica 41(3)333342

Nyland RD 1996 Silviculture Concepts and Applications McGraw-HillNew York NY 633 pp

Pielou EC 1977 Mathematical Ecology John Wiley and Sons New YorkNY 385 pp

Pough FH RM Andrews JE Cadle ML Crump AH Savitzky and KDWells 2001 Herpetology 2ndEdition Prentice Hall NJ 612 pp

Russell KR DH Van Lear and DC Guynn Jr 1999 Prescribed fire effectson herpetofauna Review and management implications Wildlife SocietyBulletin 27(2)374384

Savely HE 1939 Ecological relations of certain animals in dead pine and oaklogs Ecological Monographs 9(3)322377

SAS Institute 1997 SASSTAT software Changes and enhancements throughrelease 612 SAS Institute Cary NC

Smith DM BC Larson MJ Kelty and PMS Ashton 1997 The Practice ofSilviculture Applied Forest Ecology Wiley and Sons New York NY 537 pp

Spotila JR 1972 Role of temperature and water in the ecology of lunglesssalamanders Ecological Monographs 42(1)95125

Stoddard HL 1962 Use of fire in pine forests and game lands of the deepSoutheast Proceedings of the Tall Timbers Fire Ecology Conference TallTimbers Research Station Tallahassee FL 13142

Van Lear DH and TA Waldrop 1989 History uses and effects of fire in theAppalachians US Department of Agriculture Forest Service GeneralTechnical Report SE-54 20 pp

Walker LC 1999 The North American Forests Geography Ecology andSilviculture CRC Press Boca Raton FL 398 pp

Whiles MR and JW Grubaugh 1996 Importance of coarse woody debris tosouthern forest herpetofauna Pp 94100 In JW McMinn and DACrossley Jr (Eds) Biodiversity of Coarse Woody Debris in SouthernForests Effects on Biodiversity Proceedings of the Workshop on CoarseWoody Debris in Southern Forests US Department of Agriculture ForestService General Technical Report SE-94 146 pp

Wilson LA 1995 Land Manageriacutes Guide to the Amphibians and Reptiles ofthe SouthThe Nature Conservancy Southeastern Region Chapel Hill NC324 pp

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Hanula JL 1993 Relationship of wood-feeding insects and coarse woodydebris Pp 5565 In JW McMinn and DA Crossley Jr (Eds)Biodiversity of Coarse Woody Debris in Southern Forests Effects onBiodiversity Proceedings of the Workshop on Coarse Woody Debris inSouthern Forests US Department of Agriculture Forest Service GeneralTechnical Report SE-94 146 pp

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Hecnar SJ 1994 Nest distribution site selection and brooding in the five-linedskink (Eumeces fasciatus) Canadian Journal of Zoology 72(8)15101516

Maidens DA MA Menzel and J Laerm 1998 Notes on the effect of sizeand level of decay of coarse woody debris on relative abundance of shrewsand salamanders in the southern Appalachian mountains Georgia Journal ofScience 56(4)226-233

McLeod RF and JE Gates 1998 Response of herpetofaunal communities toforest cutting and burning at Chesapeake Farms Maryland American Mid-land Naturalist 139(1)164177

Southeastern Naturalist Vol 2 No 4486

Means DB and HW Campbell 1981 Effects of prescribed burningonamphibians and reptiles Pp 8996 In GW Wood (Ed) Prescribed Fireand Wildlife in Southern Forests Belle Baruch Forest Science InstituteClemson University Georgetown SC 170 pp

Mushinsky HR 1985 Fire and the Florida sandhill herptofaunal communitywith special attention to responses of Cnemidophorus sexlineatusHerpetologica 41(3)333342

Nyland RD 1996 Silviculture Concepts and Applications McGraw-HillNew York NY 633 pp

Pielou EC 1977 Mathematical Ecology John Wiley and Sons New YorkNY 385 pp

Pough FH RM Andrews JE Cadle ML Crump AH Savitzky and KDWells 2001 Herpetology 2ndEdition Prentice Hall NJ 612 pp

Russell KR DH Van Lear and DC Guynn Jr 1999 Prescribed fire effectson herpetofauna Review and management implications Wildlife SocietyBulletin 27(2)374384

Savely HE 1939 Ecological relations of certain animals in dead pine and oaklogs Ecological Monographs 9(3)322377

SAS Institute 1997 SASSTAT software Changes and enhancements throughrelease 612 SAS Institute Cary NC

Smith DM BC Larson MJ Kelty and PMS Ashton 1997 The Practice ofSilviculture Applied Forest Ecology Wiley and Sons New York NY 537 pp

Spotila JR 1972 Role of temperature and water in the ecology of lunglesssalamanders Ecological Monographs 42(1)95125

Stoddard HL 1962 Use of fire in pine forests and game lands of the deepSoutheast Proceedings of the Tall Timbers Fire Ecology Conference TallTimbers Research Station Tallahassee FL 13142

Van Lear DH and TA Waldrop 1989 History uses and effects of fire in theAppalachians US Department of Agriculture Forest Service GeneralTechnical Report SE-54 20 pp

Walker LC 1999 The North American Forests Geography Ecology andSilviculture CRC Press Boca Raton FL 398 pp

Whiles MR and JW Grubaugh 1996 Importance of coarse woody debris tosouthern forest herpetofauna Pp 94100 In JW McMinn and DACrossley Jr (Eds) Biodiversity of Coarse Woody Debris in SouthernForests Effects on Biodiversity Proceedings of the Workshop on CoarseWoody Debris in Southern Forests US Department of Agriculture ForestService General Technical Report SE-94 146 pp

Wilson LA 1995 Land Manageriacutes Guide to the Amphibians and Reptiles ofthe SouthThe Nature Conservancy Southeastern Region Chapel Hill NC324 pp

Southeastern Naturalist Vol 2 No 4486

Means DB and HW Campbell 1981 Effects of prescribed burningonamphibians and reptiles Pp 8996 In GW Wood (Ed) Prescribed Fireand Wildlife in Southern Forests Belle Baruch Forest Science InstituteClemson University Georgetown SC 170 pp

Mushinsky HR 1985 Fire and the Florida sandhill herptofaunal communitywith special attention to responses of Cnemidophorus sexlineatusHerpetologica 41(3)333342

Nyland RD 1996 Silviculture Concepts and Applications McGraw-HillNew York NY 633 pp

Pielou EC 1977 Mathematical Ecology John Wiley and Sons New YorkNY 385 pp

Pough FH RM Andrews JE Cadle ML Crump AH Savitzky and KDWells 2001 Herpetology 2ndEdition Prentice Hall NJ 612 pp

Russell KR DH Van Lear and DC Guynn Jr 1999 Prescribed fire effectson herpetofauna Review and management implications Wildlife SocietyBulletin 27(2)374384

Savely HE 1939 Ecological relations of certain animals in dead pine and oaklogs Ecological Monographs 9(3)322377

SAS Institute 1997 SASSTAT software Changes and enhancements throughrelease 612 SAS Institute Cary NC

Smith DM BC Larson MJ Kelty and PMS Ashton 1997 The Practice ofSilviculture Applied Forest Ecology Wiley and Sons New York NY 537 pp

Spotila JR 1972 Role of temperature and water in the ecology of lunglesssalamanders Ecological Monographs 42(1)95125

Stoddard HL 1962 Use of fire in pine forests and game lands of the deepSoutheast Proceedings of the Tall Timbers Fire Ecology Conference TallTimbers Research Station Tallahassee FL 13142

Van Lear DH and TA Waldrop 1989 History uses and effects of fire in theAppalachians US Department of Agriculture Forest Service GeneralTechnical Report SE-54 20 pp

Walker LC 1999 The North American Forests Geography Ecology andSilviculture CRC Press Boca Raton FL 398 pp

Whiles MR and JW Grubaugh 1996 Importance of coarse woody debris tosouthern forest herpetofauna Pp 94100 In JW McMinn and DACrossley Jr (Eds) Biodiversity of Coarse Woody Debris in SouthernForests Effects on Biodiversity Proceedings of the Workshop on CoarseWoody Debris in Southern Forests US Department of Agriculture ForestService General Technical Report SE-94 146 pp

Wilson LA 1995 Land Manageriacutes Guide to the Amphibians and Reptiles ofthe SouthThe Nature Conservancy Southeastern Region Chapel Hill NC324 pp