Chapter 5 (TERRA-5): Maintaining Species in the South170 amphibians, 197 reptiles, 595 birds, and...

Chapter 5: Maintaining Species in the South 113TERRESTRIAL

mammals of concern include theCarolina and Virginia northernflying squirrels, the river otter,and several rodents.

■ Twenty species of bats inhabitthe South. Four are listed asendangered: the gray bat, Indianabat, and Ozark and Virginia big-eared bats. Human disturbanceto hibernation and maternity coloniesis a major factor in their decline.

■ The South is the center ofamphibian biodiversity in theNation. However, there are growingconcerns about amphibian declines.Potential causes include habitatdestruction, exotic species, waterpollution, ozone depletion leadingto excessive ultraviolet radiation,acid deposition, synthetic chemicals,and prolonged drought conditions.

■ Seven species of amphibians arelisted as threatened or endangeredby the U.S. Fish and WildlifeService: the Houston toad, Flatwoodssalamander, San Marcos salamander,Barton Springs salamander, RedHills salamander, ShenandoahMountain salamander, and Texasblind salamander. These speciesare imperiled due to physiologicalconstraints that limit them to moisthabitats, relatively small ranges,and highly specific sites.

■ Reptile species of concern includethe Louisiana pine snake, easternindigo snake, crocodile, glass lizard,bluetail mole skink, gopher tortoise,and bog turtle. General problemsfaced by reptiles include habitatdestruction, pet trade, negative publicattitudes, degradation of aquatichabitats, and fire suppression or thelack of sufficient prescribed burning.

■ Many reptiles and amphibiansare long-lived and late maturing,and have restricted geographicranges. Managing for these specieswill require different strategies thanthose in place for birds and mammals.The paucity of monitoring datafurther inhibits their management.

Introduction

The biodiversity of the South isimpressive. Factors contributing tothat diversity include regional gradientsin climate, geologic and edaphic siteconditions, topographic variation,natural disturbance processes, andthe activities of Native Americansand European settlers (Boyce andMartin 1993, Delcourt and others1993, Healy 1985). These factorshave contributed to the diversity ofseveral species groups: salamanders,snakes, and turtles (White and others1998). The evolution of plants andanimals, combined with the isolationthat characterizes some habitats,produced remarkable levels ofendemism—species that arerestricted to special habitats.

The terrestrial vertebrate fauna ofthe South, including the entire Statesof Texas and Oklahoma, consistsof 1,208 species. This total includes170 amphibians, 197 reptiles, 595birds, and 246 mammals (NatureServe2000). Species richness is highest inTexas, Florida, North Carolina, andGeorgia (fig. 5.1). North Carolinaleads in amphibian diversity,while Texas leads in the richnessof mammals, birds, and reptiles.

The variation in species richnessamong States is influenced by

Key Findings

■ Geographic patterns of diversityin the South indicate that speciesrichness is highest in Texas, Florida,North Carolina, and Georgia. Texasleads in the richness of mammals,birds, and reptiles; North Carolinaleads in amphibian diversity. Texasdominates vertebrate richness byvirtue of its large size and thevariety of its ecosystems.

■ Loss of habitat is the primarycause of endangerment of terrestrialvertebrates. Forests, grasslands,shrublands, and wetlands havebeen converted to urban, industrial,and agricultural uses. Other factorsinclude environmental contaminants,commercial exploitation, coastaldevelopment, fire suppression,river and stream modification,and wetland degradation.

■ Species that are federally listed asthreatened or endangered consist of22 birds, 33 mammals, 7 amphibians,and 17 reptiles. Florida leads withthe number of threatened (16) andendangered (26) vertebrates; Texasis second in endangered species (23);while Mississippi is second in thenumber of threatened species (11).

■ Birds of high concern includethe red-cockaded woodpecker,bald eagle, piping plover, whoopingcrane, wood stork, black-cappedvireo, Florida scrub jay, and theroseate and least terns.

■ Habitat destruction and the paucityof large tracts of undisturbed landthreaten far-ranging mammals suchas the Florida panther, red wolf, andthe Louisiana black bear. Other

Chapter 5:Maintaining Speciesin the South

Margaret Katherine Trani (Griep)Southern Region, USDA Forest Service

What conditions will beneeded to maintain

animal speciesassociations in the South?

Southern Forest Resource Assessment114

TERRESTRIAL

KentuckyTennessee

ArkansasLouisiana

MississippiSouth Carolina

VirginiaAlabama

OklahomaGeorgia

North CarolinaFloridaTexas

0 100 200 300 400 500 600 700 800 900 1000

Amphibians Reptiles Mammals Birds

differences in size, geographic location,and environmental complexity (Steinand others 2000). Texas leads theregion with 911 vertebrate species;diversity there is influenced by theState’s large size and its diversity ofhabitats (NatureServe 2000). Florida,North Carolina, and Georgia eachsupport over 600 vertebrate species.The smallest number of species (487)occurs in Kentucky. Texas and Floridasupport species typical of Latin Americaand the Caribbean that reach theirnorthern limits there (Stein and others2000). For example, the northern limitfor the American crocodile is in theFlorida Keys and south Florida.

This diverse array of vertebratespecies is found in a variety of habitats.A habitat is comprised of the physicaland biological resources that allow aspecies to survive and reproduce. Thehabitat requirements for some speciesmay be quite narrow, while those foranother may be rather broad.

A species may require a certainhabitat structure such as vegetationheight, percent canopy cover, floristics,seral stage, patch size, or diversity andinterspersion of plant communities.Some species are constrained byabiotic factors such as the precisecave temperatures required bymany bat species. These features ofhabitat influence the distribution andabundance of species (Dickson 2001).

The habitat conditions for southernspecies have been modified by severalfactors (Buckner 1989). Habitat lossand degradation are serious threats tothe region’s fauna (Noss and others1995, Williams 1989). The rapid

growth of the human population hasresulted in land use conversion, urbansprawl, and habitat fragmentation(White and others 1998). Landscapemodification has been accompaniedby habitat isolation, water and airpollution, and altered disturbanceregimes (Lorimer 2001, Trani andothers 2001). In addition, southernwildlife has been influenced by theintroduction of exotic species andthe overexploitation of native species.Of particular concern is collectionof species for the pet trade andoverharvest of commercial species(Flather and others 1998). Thesefactors have influenced species andtheir habitats in different ways.

This chapter provides an overviewof the habitat associations of birds,mammals, reptiles, and amphibians inthe South. The focus is on vertebratesbecause information on the regionalbiogeography of many terrestrialinvertebrate groups is lacking(Echternacht and Harris 1993).Additional information on plantand animal associations is providedin chapters 1, 2, and 23.

Taxa groups are described, andgeneral habitat associations for eachare summarized. The status, distri-bution, and habitat requirementsare provided for selected species ofconcern. Finally, conservation andmanagement actions are suggestedfor enhancing habitat associationsand mitigating known threats.

The following sections discuss theconditions needed to maintain andenhance conditions for species thatoccupy the terrestrial habitats of the

South. Scientific names are providedin the chapter tables and the masterspecies list in the Assessment appendix;therefore, only common names areprovided in the text.

Methods andData Sources

Data on the status of threatened orendangered vertebrate species of theSouth were compiled from the U.S.Department of the Interior (2000). Thatagency provided information on thedistribution of listed species by State.Its recovery plans and other agencypublications were used to compileinformation on life history, ecology,and management of individual species.

Regional species richness in eachvertebrate taxon was compiledfrom State Natural Heritage offices(NatureServe 2000). This database isan inventory of all known occurrencesfor species of conservation concern.Information was derived from thedatabase to determine geographicpatterns of diversity by State in theSouth. The system was also used toverify the status and distribution ofspecies included in the fauna accounts.

Information on bird habitatassociations was obtained from Partnersin Flight (2000) conservation plans.These plans highlight the factors thatimperil bird species in physiographicareas and recommend managementactions. The conservation plans wereused to identify species of conservationconcern (Pashley and others 2000).

Habitat associations for herpe-tofauna (reptiles and amphibians)were summarized from the com-prehensive review conducted byWilson (1995). Additional literaturereviews and reference materialssupplied information on reptileand amphibian ecology.

State agency bear biologists weresurveyed for information about thecurrent status, habitat needs, andmanagement concerns about blackbears. Nine States responded withinformation: Alabama, Arkansas,Florida, Kentucky, Mississippi,North Carolina, Oklahoma, Texas,and Virginia.

Information on mammal habitatrelationships was compiled fromextensive literature searches, field

Figure 5.1—Geographic patterns of diversity by State withinthe South (NatureServe 2000).


Other birds183 (31%)

Raptors30 (5%)

Waterfowl45 (8%)

Wading birds21 (4%)

Shorebirds50 (8%)

Perching birds266 (44%)

56

55

55

54

33

08

53

0706

42

1905

04

14

21 10

11

12

13 03

23

02

01

guides, and texts on southern wildlife.Research stations and universitiesthroughout the South were contactedto obtain additional information onselected species.

Results

BirdsThe moderate climate and diverse

forests across the South supportabundant and diverse communitiesof breeding, wintering, and migratingbirds. This vertebrate group comprises17 major orders and 55 families(Echternacht and Harris 1993). Theorder Passeriformes dominates theregion’s avifauna in the number ofdifferent families (19) and species(127). These include the flycatchers,crows, swallows, jays, titmice, wrens,vireos, grackles, orioles, finches,sparrows, and warblers among others.

The South supports 595 avian species(NatureServe 2000). The number ofbird species ranges from 505 in Texasto 296 in Tennessee. Florida has 419;North Carolina, 390; Oklahoma, 359;and Alabama, 355. These totals includeperching birds, shorebirds, wadingbirds, waterfowl, raptors, and otherbirds (fig. 5.2).

Perching birds, which include thepasserines mentioned above, comprisethe majority of bird species. Examplesof shorebirds include plovers andcurlews, while wading birds includesandhill cranes and flamingos. Mottledducks, Canada geese, wood ducks,hooded merganser, and mallardsrepresent waterfowl. Eagles, hawks,kites, vultures, and owls are someof the species classified as raptors.The Natural Heritage designationof “other birds” includes gamebirds,such as bobwhite quail, ruffed grouse,American woodcock, wild turkey, andseveral dove species. This group alsoincludes woodpeckers; open oceanbirds such as cormorants, petrels, andpelicans; rails; and many other species.

The coastal wetlands support thegreatest number of species. In fact,the South supports the largest numberof wading species in the United States(White and others 1998). Thirty-onespecies occur solely at high elevationsin the Appalachian Mountains.

The South also provides habitat forsummer breeding populations, birds

that overwinter in the region, and birdsthat migrate to South America. Coastalhabitats, maritime forests, and longleafpine savanna are all important tomigrating species.

Twenty-one species of birds are listedas threatened or endangered (table 5.1).Several of these species inhabit theCoastal Plain. In addition, several birdsare classified as imperiled or vulnerableby the Natural Heritage agencies(chapter 1). These species are injeopardy due to habitat loss, habitatfragmentation, or coastal development(Hall 1995). The dependence onbreeding and staging areas has madeshorebird populations vulnerable todisturbance. Colonial waterbirds havedeclined as a result of habitatdegradation.

In contrast, the status of other specieshas improved during the past decade.The status of the brown pelican as wellas several species of raptors (ospreys,

bald eagles, and peregrine falcons) hasimproved due to habitat protection andrestrictions on the use of DDT(Fuller and others 1995).

There is a substantial body of infor-mation on bird-habitat relationships,and extensive long-term monitoringprograms have been in place forseveral decades. The distributionand composition of bird communitiesis influenced by local habitat andlandscape conditions. Local habitatfeatures include forest type, understory,number of foliage layers, canopystructure, and successional stage.Landscape conditions influencingbird populations include patchsize, interspersion of vegetativecommunities, forest fragmentation,edge length, interpatch distance,interior forest, adjacent land use,and spatial heterogeneity.

The following section discussesbird-habitat associations in the South.Species of concern are identified,and recommendations for theirmanagement are provided.

Partners in Flight physiographicareas—Partners in Flight (PIF) isan organization formed to promotebird conservation. It is comprisedof Federal and State agencies,conservation groups, and forestindustry. PIF uses physiographicareas as conservation planning unitsfor evaluating population trends,habitat conditions, land use practices,and emerging conservation issues(fig. 5.3). Boundaries defined bygeomorphology, topography, andvegetative communities are based

Figure 5.3—The distribution of Partners in Flight physiographicregions within the South (Partners in Flight 2000).

Figure 5.2—Species richness by majorsubgroups of avian taxa occurring withinthe South (NatureServe 2000).


TERRESTRIAL

Table 5.1—Bird species in the South that are listed as threatened or endangered

Scientific name Common name Areas of occurrence

Wading birdsGrus americana Whooping crane (E) FL, OK, TXGrus canadensis pulla Mississippi sandhill crane (E) MS

RaptorsFalco femoralis Northern aplomado falcon (E) TX

septentrionalisHaliaeetus leucocephalus Bald eagle (T) AL, AR, FL, GA, KY, LA, MS, NC,

OK, SC, TN, TX, VAPolyborus plancus Audubon’s crested caracara (T) FL

auduboniiRostrhamus sociabilis Everglade snail kite (E) FL

plumbeus

ShorebirdsCharadrius melodus Piping plover (T) AL, AR, FL, GA, KY, LA, MS, NC,

OK, SC, TN, TX, VAMycteria americana Wood stork (E) AL, FL, GA, SCNumenus borealis Eskimo curlew (E) OK, TX

Perching birdsAmmodramus maritimus Cape sable seaside sparrow (E) FL

mirabilisAmmodramus savannarum Florida grasshopper sparrow (E) FL

floridanusAphelocoma coerulescens Florida scrub-jay (E) FLDendroica chrysoparia Golden-cheeked warbler (E) TXEmpidonax traillii extimus Southern willow flycatcher (E) TXVireo atricapillus Black-capped vireo (E) LA, MS, OK, TX

Other birdsPelecanus occidentalis Brown pelican (E) LA, MS, TXPicoides borealis Red-cockaded woodpecker (E) AL, AR, FL, GA, KY, LA, MS, NC,

OK, SC, TN, TX, VASterna antillarum Least tern (E) AR, LA, MS, OK, TN, TXSterna dougallii dougallii Roseate tern (T, Ea) FL, GA, KY, NC, SC, VAStrix occidentalis Spotted owl (T) TXTympanuchus cupido Attwater’s greater prairie chicken (E) TX

attwateri

T = Threatened; E = endangered.a Threatened in the United States where not listed as endangered.Source: U.S. Department of the Interior (2000).

upon physiographic strata establishedby the North American Breeding BirdSurvey (Peterjohn and others 1995).Physiographic areas are distinguishedby having distinct species assemblages,land uses, and conservation issues.

Bird conservation plans prepared foreach physiographic area identify speciesand habitats of conservation concern.Seventeen physiographic areas liepredominately in the South (table 5.2).

All of the plans are available online atwww.blm.gov/wildlife/pifplans.htm.

The conservation plans prioritizebirds of concern and their habitatusing several criteria for rankinga species’ vulnerability: relativeabundance, size of breeding andnonbreeding ranges, threats duringbreeding and nonbreeding seasons,population trends, and relative density.Numerical scores are given for eachcriterion, with higher scores reflecting

higher vulnerability. Species of concernare represented by scores of 22 andabove; these species are the focus of thephysiographic area conservation plans.

Table 5.3 presents a summary ofthe birds of concern for the southernphysiographic areas. Species of concernthat occur in several physiographicareas include the swallow-tailed kite,red-cockaded woodpecker, Acadianflycatcher, Bell’s vireo, brown-headednuthatch, wood thrush, prairie


Table 5.2—Species richness by physiographic area for birds of the South (Partners in Flight 2000)

Physiographic area State(s) Total species Species of concern

No. % a

01 – Subtropical Florida FL 103 14 13.202 – Peninsular Florida FL 128 21 15.203 – South Atlantic Coastal Plain FL, GA, SC, NC 161 26 15.504 – East Gulf Coastal Plain FL, AL, MS, LA, TN 161 20 12.205 – Mississippi Alluvial Valley MS, LA, AR 143 17 11.906 – Coastal Prairies LA, TX 168 20 11.508 – Oaks and Prairies TX, OK 147 13 8.710 – Mid-Atlantic Piedmont VA 137 11 8.011 – Southern Piedmont AL, GA, SC, NC 125 14 11.212 – Mid-Atlantic Ridge and Valley VA 166 14 8.413 – Southern Ridge and Valley AL, GA, TN 131 21 16.014 – Interior Low Plateaus AL, TN, KY 159 15 9.419 – Ozark-Ouachita Plateau AR, OK 151 17 11.221 – Northern Cumberland Plateau AL, TN, KY, VA 144 18 12.523 – Southern Blue Ridge GA, SC, NC, VA 156 20 12.842 – West Gulf Coastal Plain LA, AR, TX, OK 130 18 13.844 – Mid-Atlantic Coastal Plain VA 185 20 10.6

a Species of concern represented by scores of 22 and above.

warbler, cerulean warbler, prothon-otary warbler, worm-eating warbler,Swainson’s warbler, Louisiana water-thrush, Kentucky warbler, Bachman’ssparrow, and Henslow’s sparrow. Thesespecies and the physiographic areasthey inhabit are described below.Management recommendations fromthe plans follow Pashley and others(2000) unless otherwise cited.

PIF physiographic areas: mid-Atlantic Coastal Plain—Thisphysiographic area extends from theAtlantic Ocean south of Long Islandto the Virginia-North Carolina border.The landscape is dominated by forestedwetlands, salt marshes, and barrierislands. Upland forests grade from pine-dominated areas on the outer CoastalPlain to hardwood forests on the inlandareas. This landscape has been alteredby human settlement for approximatelyfour centuries. Human populationgrowth is expected to continue, placingfurther demands on the region’snatural resources.

The mid-Atlantic Coastal Plainsupports 185 bird species; 20 (11percent) are of concern. Among thosespecies, the prairie warbler occupiespine savanna habitat, while theBachman’s sparrow occurs in grassyunderstories. Salt marshes supportimportant breeding and wintering

populations of the black duck, blackrail, salt marsh sharp-tailed sparrow,and seaside sparrow. The Acadianflycatcher, cerulean warbler, andprothonotary warbler inhabit forestedwetlands. Mixed upland forest supportsthe wood thrush in well-developedmidstories and the worm-eatingwarbler and Kentucky warbler indense understories. Henslow’s sparrowsmay also occur along the edges of saltmarsh habitat, in areas of regeneratingpines, and on former grasslands.

Conservation issues center onmanaging human population growthwhile maintaining functioningecosystems. The extensive forestedhabitat is heavily fragmented; main-taining blocks large enough to supporta diversity of breeding birds is apriority. Protection of critical sites forwintering species must be integratedwith conservation plans for breedinghabitats. Specific recommendationsinclude restoration of pine savannaconditions through prescribedburning; protection of barrier dunes tominimize losses in species productivity;protection of sites with greater than125 acres of high marsh; identificationof forest areas that support significantpopulations of prothonotary andcerulean warblers; and the restoration

of open lands greater than 125 acreswith Henslow’s sparrow potential.

PIF physiographic areas: mid-Atlantic Piedmont—The mid-AtlanticPiedmont is separated from thesouthern Piedmont at the NorthCarolina-Virginia line. It extends norththrough Virginia, Maryland, andPennsylvania before terminating innorthern New Jersey. The rollingtopography formerly supportedextensive hardwood forests includingoak-hickory, Appalachian oak, andloblolly-shortleaf pine. Approximately45 percent of the physiographic areais presently forested, 45 percent isin agricultural production, and theremainder is in urban areas.

The mid-Atlantic Piedmont supports137 bird species; 11 (8 percent) are ofconcern. Deciduous and mixed foresthabitats support the wood thrush,cerulean warbler, Louisiana water-thrush (in riparian forest buffers),and Kentucky warbler (in denseunderstory). The shrub-scrub areasand barrens support the bobwhitequail (in decline). The Americanwoodcock (also in decline) requiresan interspersion of forest clearingsand second-growth hardwoods.Agricultural pastureland supportsa large population of grasshoppersparrows and other grassland species.


TERRESTRIAL

Table 5.3—Bird species of concern in the South (Partners in Flight 2000)

Scientific name Common name Physiographic areasa

Egretta rufescens Reddish egret 02, 04, 08Eudocimus albus White ibis 01, 02Anas rubripes American black duck 44Anas fulvigula Mottled duck 01, 02, 06Elanoides forficatus Swallow-tailed kite 01, 02, 04, 05, 06, 08, 42Rostrhamus sociabilis Snail kite 01, 02Buteo brachyurus Short-tailed hawk 01, 02, 03Tympanuchus cupido Greater prairie chicken 06, 08Colinus virginianus Northern bobwhite 03, 04, 08, 11Laterallus jamaicensis Black rail 02, 03, 08, 44Rallus longirostris Clapper rail 02, 03, 06Grus Canadensis Sandhill crane 02Charadrius alexandrinus Snowy plover 02Charadrius wilsonia Wilson’s plover 02, 03, 06Charadrius melodus Piping plover 03, 44Haematopus palliatus American oystercatcher 02, 03, 04, 08Sterna forsteri Forster’s tern 06Columba leucocephala White-crowned pigeon 01Coccyzus americanus Yellow-billed cuckoo 04, 05, 08, 13, 14, 42Coccyzus minor Mangrove cuckoo 02Caprimulgus carolinensis Chuck-will’s-widow 04, 13, 42Caprimulgus vociferus Whip poor will 10, 11, 12, 14Chaetura pelagica Chimney swift 14Amazilia yucatanensis Buff-bellied hummingbird 06Picoides borealis Red-cockaded woodpecker 01, 02, 03, 04, 11, 13, 19, 21, 23, 42, 44Campephilus principalis Ivory-billed woodpecker 03Contopus virens Eastern wood-pewee 12, 44Empidonax virescens Acadian flycatcher 10, 13, 19, 21, 23, 44Tyrannus dominicensis Gray kingbird 01, 02Tyrannus forficatus Scissor-tailed flycatcher 06, 08, 42Lanius ludovicianus Loggerhead shrike 02Vireo griseus White-eyed vireo 05, 42Vireo bellii Bell’s vireo 04, 05, 06, 08, 14, 19, 42Vireo flavifrons Yellow-throated vireo 12, 13, 21, 23, 44Vireo altiloquus Black-whiskered vireo 01, 02Aphelocoma coerulescens Florida scrub-jay 01, 02, 03Tachycineta cyaneoviridis Bahama swallow 01Petrochelidon fulva Cave swallow 01, 06, 08Sitta pusilla Brown-headed nuthatch 02, 03, 04, 05, 10, 11, 13, 19, 23, 42, 44Thryomanes bewickii Bewick’s wren 11, 12Hylocichla mustelina Wood thrush 03, 05, 10, 12, 13, 19, 21, 23, 44Toxostoma longirostre Long-billed thrasher 06Vermivora bachmanii Bachman’s warbler 03, 04, 05Vermivora pinus Blue-winged warbler 05, 13, 14, 44Vermivora chrysoptera Golden-winged warbler 12, 13, 21, 23Parula americana Northern parula 03, 05, 12Dendroica pensylvanica Chestnut-sided warbler 23Dendroica caerulescens Black-throated blue warbler 04, 05, 12, 13, 21, 23Dendroica fusca Blackburnian warbler 23Dendroica dominica Yellow-throated warbler 03, 13, 23Dendroica discolor Prairie warbler 01, 03, 04, 10, 11, 12, 13, 14, 19, 21, 23,

42, 44continued


Table 5.3—Bird species of concern in the South (Partners in Flight 2000) (continued)

Scientific name Common name Physiographic areasa

Dendroica cerulea Cerulean warbler 03, 04, 05, 10, 11, 12, 13, 14, 19, 21, 23,42, 44

Mniotilta varia Black-and-white warbler 23Protonotaria citrea Prothonotary warbler 03, 04, 05, 06, 11, 13, 19, 21, 42, 44Helmitheros vermivorus Worm-eating warbler 03, 04, 05, 08, 10, 11, 12, 13, 14, 19, 21,

23, 42, 44Limnothlypis swainsonii Swainson’s warbler 03, 04, 05,06, 08, 11, 12, 13, 14, 19, 21,

23, 42, 44Seiurus motacilla Louisiana waterthrush 12, 13, 14, 19, 21, 23, 42Oporornis formosus Kentucky warbler 04, 05, 06, 10, 11, 13, 14, 19, 21, 23, 42,

44Wilsonia citrina Hooded warbler 03, 21, 23, 42Wilsonia canadensis Canada warbler 23Piranga rubra Summer tanager 21Aimophila aestivalis Bachman’s sparrow 02, 03, 04, 06, 10, 11, 13, 14, 19, 21, 23,

42, 44Spizella pusilla Field sparrow 10, 11, 13, 14, 19, 21Ammodramus henslowii Henslow’s sparrow 03, 06, 10, 11, 12, 14, 19, 21, 44Ammodramus caudacutus Saltmarsh sharp-tailed sparrow 03, 44Ammodramus maritimus Seaside sparrow 01, 02, 03, 04, 06, 44Passerina ciris Painted bunting 03, 05, 06, 08, 19Spiza americana Dickcissel 06, 14, 19Icterus spurius Orchard oriole 04, 05, 13, 42Icterus graduacauda Audubon’s oriole 06

a Physiographic areas: 01 – Subtropical Florida, 02 – Peninsular Florida, 03 – South Atlantic Coastal Plain, 04 – East Gulf Coastal Plain,05 – Mississippi Alluvial Valley, 06 – Coastal prairies, 08 – Oaks and prairies, 10 – Mid-Atlantic Piedmont, 11 – Southern Piedmont,12 – Mid-Atlantic Ridge and Valley, 13 – Southern ridge and valley, 14 – Interior Low Plateaus, 19 – Ozark-Ouachita Plateau,21 – Northern Cumberland Plateau, 23 – Southern Blue Ridge, 42 – west Gulf Coastal Plain, and 44 – Mid-Atlantic Coastal Plain.

Conservation issues center on themanagement of human populationgrowth and protection of conservationareas. Enhancement of grasslandhabitat also is a priority. Specificrecommendations include managementof areas that support significantpopulations of cerulean and Kentuckywarblers, restoration of naturalbarrens that support shrub-nestingspecies, and monitoring priorityspecies in disturbed areas.

PIF physiographic areas: mid-Atlantic Ridge and Valley—Thisphysiographic area extends fromwestern Maryland through themountains of Virginia. Consistingof mountain ridges and interveningvalleys, the predominant forest typeis oak-hickory. Relict patches of spruce-fir occur on high mountain ridges.Agricultural production and urbandevelopment dominate in the lowervalleys. Human populations arerelatively sparse and confined to thevalleys, while coal extraction occurs on

public and private forests. Disease andinsect pests are important disturbancefactors; the pesticides used for gypsymoth control impact other foliageinsects that are important bird food(Hunter and others 2001).

The mid-Atlantic Ridge andValley supports 166 bird species;14 (8 percent) are of concern. Earlysuccessional shrub habitat (includingbarrens and disturbed sites) supportsthe whippoorwill, golden-wingedwarbler, and prairie warbler. Thewood thrush and worm-eating warbleroccupy mature deciduous forest, whilethe Louisiana waterthrush is found inlate successional stands near streams.The black-throated blue warbler andthe blackburnian warbler use northernhardwood and spruce-fir forests.

Conservation issues center on long-term planning on public land to meetthe habitat needs of species requiringspecific seral stages. On public land,it is important to balance the needs ofearly successional species with those

requiring mature forest (Trani andothers 2001). Specific actions neededfor this physiographic area includemanagement of high-elevationspruce-fir habitat, intensive surveysfor Appalachian Bewick’s wren,identification of breeding sites forgolden-winged warbler, and themaintenance of composition andstructural diversity.

PIF physiographic areas:northern Cumberland Plateau—The Cumberland Plateau is apredominantly forested, gently rollingtableland bordered by the eastern rimof the Interior Low Plateaus and theCumberland Mountains (fig. 5.3). Thearea includes eastern Kentucky andTennessee, southwestern West Virginia,and a small area in western Virginia.Forests dominated by oaks andhickories are common. Various pinespecies are dominant on drier sites.

The northern Cumberland Plateausupports 144 bird species; 18 (12percent) are of concern. Among species


TERRESTRIAL

of concern, the Acadian flycatcher,wood thrush, worm-eating warbler,and Swainson’s warbler inhabit mixedmesophytic forests. Coniferous forestssupport the red-cockaded woodpecker(low-elevation yellow pine) andBachman’s sparrow. Bewick’s wrenand golden-winged warbler use earlysuccessional habitat, while Henslow’ssparrow occurs in grassland areas. Bothhabitats exist only due to disturbance.

Conservation issues center on themaintenance of species compositionand vegetation structure. Widespreadtimber harvesting and fire suppressionhave reduced both old-growth andyoung forest habitats. The currentstructure of the mid-seral forest maynot be optimal for many midstory andunderstory breeding birds. As a resultof diminishing habitat quality, severalhigh-priority birds have undergonesignificant population declines.The northern Cumberland Plateauis one of the most heavily forestedphysiographic areas in the South.Specific recommendations includemanagement of 12 to 15 percent offorests for long-rotation sawtimberor old growth, increased use of fire inlow-elevation yellow pine habitat, andmaintenance of shrub-scrub conditions.

PIF physiographic areas: southernRidge and Valley—This physiographicarea includes the southern end of theRidge and Valley and the tablelandsof the southern Cumberland Plateau.It is in eastern Tennessee, northwestGeorgia, and northeast Alabama.The upland forest is predominantlyin oak-hickory and pine (shortleafor loblolly) types.

The southern Ridge and Valleysupports 131 bird species; 21(16 percent) are of concern. Earlysuccessional scrub-shrub habitat isoccupied by the Bewick’s wren, blue-winged warbler, and orchard oriole.The hardwood forest componentsupports the Acadian flycatcher, yellow-throated warbler, prothonotary warbler,worm-eating warbler, and Swainson’swarbler among others. Red-cockadedwoodpeckers and brown-headednuthatches are found in southern pines.

Conservation issues focus onthe conversion of hardwood forestto monocultures of loblolly pine. Alarge percentage of natural vegetationhas been cleared for agriculture andurban development. Birds dependenton mature forest may be at risk because

the amount of public land may not besufficient to support viable populationsof sensitive species (Hunter and others2001). Enhancement of habitat forthese species will require the use oflong-rotation harvests. Specificrecommendations include expansionof longleaf habitat using prescribed fireand the enhancement of scrub habitat.

PIF physiographic areas: southernBlue Ridge—The Southern BlueRidge runs along the border betweenTennessee and North Carolina, extend-ing into South Carolina, Georgia, andVirginia. The area is comprised ofrugged mountains, broad ridges, steepslopes, and deep ravines. Spruce-firforests at the highest elevationtransition into northern hardwoods,hemlock-white pine, and Appalachianoaks at lower elevations. Cove forestsoccur on mesic sites, while fire-associated yellow pines occur ondry ridges. Disturbances from fire,grazing, and storms are primary factorsin determining forest compositionand structure.

The southern Blue Ridge supports156 bird species; 20 (13 percent) areof concern. Among species of concern,the northern saw-whet owl, black-capped chickadee, red-breastednuthatch, golden-crowned kinglet, redcrossbill, and yellow-bellied sapsuckerare distinct subspecies whose rangesare centered within the southern BlueRidge. With the exception of thesapsucker, each species occupies highforested peaks. The yellow-belliedsapsucker, as well as the golden-wingedwarbler, inhabits disturbed forest areas.Among species of concern requiringmature forest in the southern BlueRidge are Acadian flycatchers, yellow-throated vireos, wood thrushes,blackburnian warblers, Swainson’swarblers, Kentucky warblers, andCanada warblers.

Conservation issues includepopulation declines of both migrantand resident birds. The rapid con-struction of new homes and associateddevelopments along roads contribute tohabitat loss and fragmentation. Anotherconcern is the decline of high-elevationspruce-fir forests resulting fromexotic pests and reduced air quality.Atmospheric pollution is reducing treegrowth, insectivore food availability,and supplies of important mineralsnecessary for successful bird repro-duction (Hunter and others 2001).

Many species in this habitat are inisolated endemic populations that maybe genetically distinct from populationselsewhere. Populations of priority birds,such as the Appalachian subspecies ofBewick’s wren, have declined in recentyears. Maintenance of early successionalhabitat is a conservation need. Otherrecommendations include managementof riparian zones and the provisionof old-growth forest.

PIF physiographic areas: southernPiedmont—This physiographicarea extends through central NorthCarolina, South Carolina, and Georgiainto eastern Alabama. Plains, hills,tablelands, and numerous riverscharacterize the Piedmont. The area liesbetween the Appalachian Mountainsand the Coastal Plain. The dominantvegetation includes oak-hickory andmixed hardwood forests. Shortleaf,loblolly, and scattered longleaf pinesare prevalent on disturbed sites.

The southern Piedmont supports125 bird species; 14 (11 percent) are ofconcern. Among species of concern, theprairie warbler, Bachman’s sparrow, andHenslow’s sparrow are supported bygrassland and shrub habitat. Southernpine forests support the red-cockadedwoodpecker and brown-headednuthatch, while prothonotary andSwainson’s warblers use the bottomlandhardwoods. Upland hardwood habitatsupports the whippoorwill, woodthrush, and cerulean warbler.

Conservation challenges focus onhuman population growth, urbansprawl, and the intensification ofagriculture and timber harvesting.Several bird populations have declinedin patches of protected mature forestsembedded in suburban settings. Inaddition, changing land use hasresulted in a loss of early successionalhabitat. Public lands provide core areasin the Piedmont on which to managehabitat. The maintenance of birdcommunities requires coordinationamong public agencies, forest industry,and private landowners.

PIF physiographic areas:south Atlantic Coastal Plain—Thesouth Atlantic Coastal Plain coversnortheastern Florida, southern Georgia,the eastern Carolinas, and the GreatDismal Swamp in Virginia. Coastalareas contain barrier islands, maritimeforests, marshland, and estuaries.Inland areas support bottomlandhardwood forests, pocosins, and


Carolina bays. Fire-maintained forestsof longleaf, shortleaf, and loblollypine once dominated upland areas.

The south Atlantic Coastal Plainsupports 161 bird species; 26 (15percent) are of concern. Amongspecies of concern, the Americankestrel, red-cockaded woodpecker,and brown-headed nuthatch requirepine forest, and Henslow’s sparrowrequires pocosin grasslands. Theswallow-tailed kite, northern parula,Swainson’s warbler, and hoodedwarbler occupy bottomland andupland hardwood forests. The prairiewarbler and painted bunting arefound in the scrub-shrub habitat.

Conservation concerns include firemanagement, land conversion, andshort-rotation pine plantations.Restoration of fire-maintainedpine savanna benefits pine-grasslandspecies, particularly the red-cockadedwoodpecker. Pine plantations are usedby other species, but the maintenanceof age class diversity is important.Other recommendations includemaintenance of large tracts of bottom-land forest in river systems to benefitblack-throated green warblers andbreeding swallow-tailed kites, andretention of coastal maritime forestand scrub-shrub habitats for thebunting and in-transit migrants.

PIF physiographic areas: peninsularFlorida—This physiographic areaextends from the northern edge of LakeOkeechobee in central Florida to theSuwanee River in northern Florida.Habitat includes sandhill, scrub,and xeric hammock communities.Longleaf pine, turkey oak, andwiregrass characterize the fire-dependent sandhill communities.Dominant scrub vegetation includessand pine and scrub oak. Xerichammocks support live oak, laureloak, and saw palmetto. Uplandhardwoods, wetlands, and man-groves are also locally common toabundant in the physiographic area.

Peninsular Florida supports 128 birdspecies; 21 (15 percent) are of concern.Among species of concern, crestedcaracara (threatened), burrowing owl,Florida scrub jays (endangered), andgrasshopper sparrows inhabit the scruband grassland habitat. Wetland andmangrove habitats support the swallow-tailed kite, snail kite (endangered), andshort-tailed hawk. The painted buntingoccurs in maritime scrub, while the

American kestrel, red-cockadedwoodpecker, and Bachman’s sparrowuse pine forests.

Conservation actions are directedat fostering cooperative relations withprivate landowners, and encouragingproper habitat management througheducation, tax breaks, and conservationeasements. Conservation goalsalso include the public acquisitionof acreage in sandhills, oak scrub,upland forest, and floodplainswamp communities.

PIF physiographic areas:subtropical Florida—Thisphysiographic area extends south fromLake Okeechobee in central Floridato the Florida Keys. The tropicalecosystem contains the Evergladesand the Big Cypress Reserve. Fireis an important feature in the pine,marsh, and prairie communities.Hurricane disturbances create earlysuccessional habitat. Distinct dryand wet seasons influence the nestingcycles of many birds.

Subtropical Florida supports 103 birdspecies; 14 (13 percent) are of concern.Pine rocklands, flatwoods, and sandscrub habitats are used by the Floridascrub jay, sedge wren, and palmwarbler. Grassland and dry prairiecommunities support the sandhillcrane and grasshopper sparrow. Theshort-tailed hawk, white-crownedpigeon, and gray kingbird inhabitsubtropical deciduous forests. Thereddish egret, white ibis, wood stork,seaside sparrow, and several speciesof rails use the brackish saltwaterand freshwater marsh habitats of theEverglades. Mangrove swamps supportthe mangrove cuckoo, the black-whiskered vireo, and the Cubansubspecies of the yellow warbler.

Conservation concerns are directedtowards the rapidly growing humanpopulation in the region. Habitats havebeen lost by converting land to urbanand agricultural uses, such as sugarcaneand citrus production. Other problemsinclude pollution and alteration of thehydroperiod and natural water cycles.Recommendations include aggressiveacquisition programs and the main-tenance of pine-dominated stands andprairies through prescribed burning.Programs for bird conservationwere created by the Surface WaterImprovement and Management Act,Florida’s Everglades Forever Act, and

the South Florida EcosystemRestoration Task Force.

PIF physiographic areas: InteriorLow Plateaus—The Interior LowPlateaus extend from Alabamanorthward across central Tennesseeand Kentucky into southern Illinois,Indiana, and Ohio, encompassing thecentral basin and Tennessee Valley.Oak-hickory and beech-maple forestswere historically the most abundantcover types. There were also tallgrassprairies and oak savannas in thenorthern section. Barrens and glades arerare in the central regions, and forestedwetlands occur along major waterways.

The Interior Low Plateaus support159 bird species; 15 (9 percent) areof concern. Priority species inhabitinghardwood forest include thewhippoorwill, cerulean warbler, andLouisiana waterthrush. The grassland,savanna, and old-field habitats supportthe Bewick’s wren, blue-wingedwarbler, and dickcissel.

Conservation issues center on habitatloss from land conversion, habitatdeterioration, and fragmentation.Pastureland has replaced grasslandand savanna, while glades and barrenshave become urban areas. Fire suppres-sion has allowed woody vegetationto encroach into open areas. Flood-plain forests have been convertedto reservoirs or row crops. Previousforest management and chipping ofall woody vegetation have influencedcanopy characteristics, understorydevelopment, and age structureof upland forests.

Specific recommendations includethe reestablishment of greater prairiechicken and swallow-tailed kitepopulations, maintenance of existingforested acreage, and the restorationof forested wetlands, warm seasongrasses, and oak savannas.

PIF physiographic areas: Ozarksand Ouachitas—The Ozark Mountainsextend from southern Missouri intonorthern Arkansas and consist ofdissected plateaus covered by oakforest with glade and savannainclusions. The ridge and valleysystem of the Ouachitas coverscentral Arkansas, reaching intoeastern Oklahoma. Vegetation includesshortleaf pine and deciduous forests.The vegetation changes to prairie inthe northern reaches.


TERRESTRIAL

The Ozarks and Ouachitas support151 bird species; 17 (11 percent) are ofconcern. Deciduous and mixed foresthabitat supports the whippoorwill,worm-eating warbler, and Kentuckywarbler. The red-cockaded woodpeckerand Bachman’s sparrow occur in pinesavanna; populations of both specieshave declined dramatically due to fireexclusion and forestry practices. TheBewick’s wren and the field sparrow useearly successional habitat; both speciesare undergoing significant declines.

Conservation actions include theimprovement of shortleaf pine, glade,and savanna communities throughthe use of thinning, overstory removal,and dormant season burns. Otheractivities include the prevention offorest fragmentation stemming fromurbanization and the management ofhabitat required by early succes-sional species.

PIF physiographic areas: East GulfCoastal Plain—The East Gulf CoastalPlain extends from Louisiana andwestern Florida northwards throughMississippi and Alabama into Tennesseeand Kentucky. Numerous streams andrivers break the rolling topography.Uplands are dominated by shortleafpine and mixed hardwoods. Loblollypine and bottomland hardwood forestsoccur in the lowland areas.

The East Gulf Coastal Plain supports161 bird species; 20 (12 percent)are of concern. Swallow-tailed kites,prothonotary warblers, and Kentuckywarblers occur in the forested wetlandsand other habitat. The northernbobwhite, Mississippi sandhillcrane (endangered), red-cockadedwoodpecker, and sedge wren occupythe pine and savanna habitats.Chuck-will’s-widow occurs in uplandhardwoods, while the LeConte’ssparrow and orchard oriole are presentin the scrub habitat. Numerous springmigrants use the maritime forests.Emergent wetlands support the reddishegret, yellow and black rails, andNelson’s sharp-tailed sparrow. Snowy,piping, and Wilson’s plovers inhabitthe beach dunes community.

Conservation issues include theconversion of longleaf pine and uplandhardwoods to other species, hydro-logical alteration, land use changesincluding coastal development, andthe changes in species compositionand structure resulting from firesuppression. Specific recommendations

include maintenance of large tracts oflongleaf pine and upland hardwoodsfor red-cockaded woodpeckers,swallow-tailed kite, cerulean warbler,Swainson’s warblers, and associatedspecies. Other actions include thecontrol of exotic plants and therestoration of maritime forest, emergentwetlands, and beach dunes thatare important to priority breedingand wintering birds.

PIF physiographic areas:Mississippi Alluvial Valley—Encompassing the floodplain of theMississippi River, the valley includeseastern Louisiana, eastern Arkansas,northwestern Mississippi, and portionsof Tennessee, Kentucky, and Missouri.The South’s biggest concentration ofbottomland hardwoods is in theMississippi River Valley, whereagricultural conversion has resulted inforest fragmentation. The MississippiRiver and its flood regimes, whichinfluence vegetation communitiesand bird habitat conditions, shapethis physiographic area.

The Mississippi Alluvial Valleysupports 143 bird species; 17 (12percent) are of concern. Among speciesof concern, the swallow-tailed kite,northern parula, and painted buntingare supported by bottomland hardwoodforests. Marsh, wetland, and open landsupport several species of shorebirdsand waterfowl and provide importantwintering areas for mallards, woodducks, and other birds.

Conservation recommendationstarget the restoration of bottomlandhardwood forest to support healthypopulations of a suite of birds. Sincesettlement, over 80 percent of the foresthas been cleared for agriculture andother uses. The hydrology has beendramatically altered, inhibitingecosystem functions. The resultingforest fragmentation has reduced theability of the area to support many birdpopulations. The Lower MississippiValley Joint Venture leads restorationefforts (Pashley and others 2000).

PIF physiographic areas: WestGulf Coastal Plain—The West GulfCoastal Plain is located in northwesternLouisiana, southwestern Arkansas,eastern Texas, and southern Oklahoma.The physiographic area is characterizedby loblolly pine, shortleaf pine, andlongleaf pine forests on the uplands,hardwood forests in the bottomlands,

and grasslands in the southern-most areas.

The West Gulf Coastal Plain supports130 bird species; 18 (14 percent)are of concern. Among such species,the American kestrel, chuck-will’s-widow, scissor-tailed flycatcher,brown-headed nuthatch, Bewick’swren, prairie warbler, and Bachman’ssparrow are supported by pineforests and associated grasslands.The swallow-tailed kite, white-eyedvireo, worm-eating warbler, Swainson’swarbler, and hooded warbler occupyhardwood forests and other supportedhabitats. The bottomland forests andriparian habitats are important forstopover migrants.

Conservation issues include firesuppression and regeneration practicesthat have replaced native species withloblolly or slash pine. Althoughmany bird species occur in youngpine plantations, others such as thered-cockaded woodpecker requirenative pine savanna conditions ormature longleaf pine stands. Specificrecommendations include themaintenance of mature longleafpine stands with fire, preventionof additional forest conversion toagricultural uses, and deterrenceof bottomland hardwood loss dueto inundation by reservoirs. Theimportance of these hardwoods forarea-sensitive species and springmigrants extends beyond the WestGulf Coastal Plain.

PIF physiographic areas: oaksand prairies—This physiographicarea extends from the Red River ofOklahoma southward into Texas.Tallgrass prairie, post-oak savanna,bottomland hardwood forests, riparianforests, and upland hardwood forestsassociated with dense scrub layerscharacterize the area. Wetlands andfreshwater marshes are associatedwith streams, rivers, and reservoirs.

The oaks and prairies support147 bird species; 13 (9 percent) areof concern. Among such species,the greater prairie chicken, northernbobwhite, scissor-tailed flycatcher,Bell’s vireo, and painted buntingare supported by grassland andscrub habitats.

Conservation issues focus on theloss of prairie habitat. Areas of tallgrassprairie have been converted to cropproduction; less than 10 percent of


Salamanders107 (63%)

Frogs and toads63 (37%)

Table 5.4—Amphibian species in the South that are listed as threatened or endangered


Frogs and toadsBufo houstonensis Houston toad (E) TX

SalamandersAmbystoma cingulatum Flatwoods salamander (T) AL, FL, GA, SCEurycea nana San Marcos salamander (T) TXEurycea sosorum Barton Springs salamander (E) TXPhaeognathus hubrichti Red Hills salamander (T) ALPlethodon shenandoah Shenandoah Mountain salamander (E) VATyhplomolge rathbuni Texas blind salamander (E) TX

Source: U.S. Department of the Interior (2000).

original prairie exists. The continuedloss of tallgrass habitat inhibitsrestoration efforts by reducing geneticdiversity; preservation of remaininghabitat is critical. Encroachmentby heavy woody growth and exoticspecies also causes loss of grasslandhabitat. Prescribed fire and grazingmanagement through incentiveprograms are beneficial.

PIF physiographic areas: coastalprairies—This physiographic area isfound along the Gulf Coast shorelinein Louisiana and Texas. The areasupports a complex of marshes, uplandgrassland, and forested habitat. Marshcommunities include salt, brackish,and freshwater marsh. The majority ofgrassland habitats have been convertedto pasture and rice farms. Forestedareas occur along major rivers, beach-front ridges, salt domes, and manmadelevees. These woodlands are comprisedof hackberry and live oak, while thebottomland hardwood forests containthe cypress-tupelo, hackberry-ash-elm,and oak-willow forest types.

The coastal prairies support 168 birdspecies; 20 (11 percent) are of concern.Priority grassland birds include thegreater prairie chicken, short-earedowl, sedge wren, and Sprague’s pipit.The bottomland hardwood forestsupports the swallow-tailed kite,American woodcock, prothonotarywarbler, and Swainson’s warbler. Bell’svireo and painted bunting occupyscrub-shrub habitat. In addition,many passerine species use the coastalhabitat during spring migration.

Conservation concerns focus onthe alteration of natural communitiesin the coastal prairies. Oil and gasdevelopment, dredging, and impound-

ments have degraded marsh habitat.Grazing animals have degraded grass-land and woodland areas. Specificrecommendations include cooperativemanagement with private landowners,incentive programs, and identificationof potential habitat for priority birds.Other actions include marshrestoration, retention of forestedwetlands, exotic species control(especially Chinese tallow), andmonitoring the influence of rice cropconversion on waterbird species.

Additional information on the habitatassociations of bird species in the Southcan be found in Hunter and others(2001) and Hamel (1992). Thephysiographic associations for nonbirdtaxa are not as well developed as thosepresented above for birds. Therefore,the habitat needs of mammals, reptiles,and amphibians will be discussed bybroad taxonomic grouping.

AmphibiansTwo orders of amphibians are present

in the Southern United States: Caudata(salamanders) and Anurans (frogs andtoads). The South supports the highestdensity of amphibian species in NorthAmerica (Echternacht and Harris1993). The total includes 107 sala-manders and 63 species of frogs andtoads (fig. 5.4). In individual States,the number of amphibian speciesranges from 80 in North Carolinato 49 in Arkansas (NatureServe 2000).Numbers in other States are 77 inGeorgia, 75 in Texas, 73 in Virginia,and 70 in Tennessee.

The Southern Appalachians have anunusually large number of salamanderspecies, because many plethodontidspecies evolved there. These lungless

animals are believed to have evolvedin fast-flowing, oxygenated streams.The numbers of salamanders inhabitingNorth Carolina (50), Virginia(48), Tennessee (48), and Georgia(44) reflect the importance of theAppalachian Mountains. The numberof salamanders occurring in theCoastal Plain is lower because habitatand temperature are less suitableand because densities of terrestrialand aquatic predators are higher(Echternacht and Harris 1993).

Numbers of frogs and toads arehighest in the southernmost CoastalStates. Numbers of species are 43 inTexas, 33 in Georgia, 32 in Florida, 31in Louisiana, 31 in South Carolina, and30 in Alabama (NatureServe 2000). Themajority of southern species are in fivefamilies: true frogs; tree, chorus, andcricket frogs; true toads; narrowmouthtoads; and spadefoot toads. Elevenspecies are endemic to the South(Echternacht and Harris 1993).

Figure 5.4—Species richness by majorsubgroups of amphibian taxa occurringwithin the South (NatureServe 2000).


TERRESTRIAL

Table 5.5—The relationships of amphibians to physiographic features andother habitat elements a

Forest cover types Salamanders Frogs and toads

No. % No. %Physiographic feature

Sandhills 3 3 8 20Flatwoods 12 13 12 30Narrow stream margins 29 32 3 8Broad stream margins 22 24 20 50Swamps 16 17 17 43Cypress strands 10 11 5 13Cypress ponds 14 15 12 30Cypress drains 13 14 9 23Willow heads 11 12 9 23Bays and pocosins 15 16 14 35Rivers and streams 34 37 9 23Permanent ponds 8 9 20 50Vernal ponds 16 17 27 68Lakes 9 15 13 33

Specific requirementClosed canopy 76 83 11 28Open canopy 12 13 32 80Shrub thickets 0 0 8 20Moist soil 69 75 25 63Sandy or friable soils 5 5 18 45Leaf litter 75 82 22 55Snags 0 0 2 5Fallen logs 70 76 5 13Rock outcrops 8 9 1 3Crevices and/or caves 11 12 0 0Seepages 23 25 12 30Potholes 12 13 22 55Aquatic rocks and/or logs 30 33 0 0Aquatic vegetation 10 11 26 65

a Data are summarized from species accounts presented in Wilson (1995).

Seven species of amphibians arelisted as threatened or endangered bythe U.S. Fish and Wildlife Service (table5.4). In addition, several amphibiansare classified as imperiled or vulnerableby the Natural Heritage agencies(chapter 1).

Amphibians have complex life cyclesand inhabit a variety of environments.Habitats include ephemeral pools,caves, forests, wetlands, savannas, andseveral aquatic habitats. The longleafpine-wiregrass community, cypress-gum swamps, isolated wetlands, andmixed hardwood-pine habitats supporta diversity of species. The federallylisted flatwoods salamander is found inthe longleaf pine-wiregrass ecosystem.Coastal Plain forests provide habitatfor ambystomatid species. In evengreater abundance in the South arethe many species of tree frogs, toads,and other frogs. Pine barrens treefrogs occur in Coastal Plain forestsfrom Alabama northwards (Gibbonsand Buhlmann 2001).

Amphibians are very differentphysiologically from reptiles, but thetwo groups are classified together asherpetofauna. Amphibians are morerestricted by environmental moisturethan other terrestrial vertebrates.They depend on areas where thereis sufficient moisture for reproductionand survival. Since the glandularthin skin of amphibians is permeableto water, evaporative water lossis a serious problem. In addition,drought affects egg laying and larvaesurvival. The demands of water balanceand thermoregulation may restrictmovement, which occurs in a narrowrange of environmental conditions.

Many amphibian species havegeographic ranges that are restrictedto particular physiographic regions.Some salamander species are consid-ered glacial relicts that were isolatedon mountaintops that retained northernclimates (Gibbons and Buhlmann2001). Similarly, frog species such asthe pine barrens tree frog, Houstontoad, and Florida bog frog occur insmall, isolated populations throughouttheir ranges. The distances betweensuch disjunct populations makerecolonization difficult.

Salamanders—The majorityof southern salamanders are insix families: mole salamanders,amphiumas, hellbenders, lunglesssalamanders, waterdogs or mud-

puppies, and sirens. Salamandersare inconspicuous species that areimportant components of the forestecosystem. They are small, secretive,and primarily nocturnal. They rangefrom 5 cm to over 1 m in length.Limited data suggest that generationtimes are relatively long. For example,the generation times for several speciesof salamanders range between 4.4and 9.5 years.

The rate of reproduction inamphibians is highly variable, butmany species exhibit low frequenciesof reproduction. Often salamandersbreed only in alternate years, whenthey lay a single clutch of eggs.

Moisture is a limiting factor for allsalamander species. Some species are

totally aquatic, but even the terrestrialspecies can only survive in moistmicrohabitats. Ambystoma andHemidactylus salamanders requiremoist, friable soils for burrowing.Several terrestrial species migrate toaquatic habitats for egg deposition,while others require damp micro-habitats. In addition, some aquaticspecies use terrestrial habitat fordispersal and other seasonal activity.

Salamanders inhabit areas with avariety of physiographic features, butrivers, streams, and stream marginsfigure prominently in their occurrence(table 5.5). Coastal bayous, ponds,and slow-moving rivers support sirensand amphiumas, while the hellbenderoccurs in cooler, fast-flowingupland rivers.


Table 5.6—The relationship between forest cover type and amphibianoccurrence in the South a

Forest cover types Salamanders Frogs and toads

No. % No. %

Everglades 4 4 9 23Tropical hardwoods 0 0 14 10Longleaf-slash pine 6 7 14 35Pine-flatwoods 9 10 19 48Virginia-pitch pine

(xeric upland pines) 19 21 10 25Longleaf pine 7 8 22 55Loblolly-shortleaf pine 28 30 15 38White pine-hemlock 43 47 10 25Pond pine 5 5 14 35Longleaf-scrub oak 2 2 8 20Mixed, pine-hardwood (mesic) 66 72 33 83Mixed, pine-hardwood (xeric) 0 0 22 55Spruce fir 10 11 0 0Upland hardwoods (mesic)-

white oak-red oak 59 64 19 48Cypress tupelo 26 28 22 55Bottomland hardwoods-

(sweetgum-willow oak) 43 47 31 78Sweetgum–yellow-poplar 30 33 20 50Bay-pocosin 20 22 22 55Live oak (maritime) 6 7 12 30Maple-beech 24 26 9 23Cove hardwoods 29 32 7 18Spartina 0 0 4 10Elm-ash 1 1 4 10Oak-hickory (xeric hardwoods) 38 41 15 38Cave dwelling 6 7 0 0Aquatic dependent 19 21 0 0

a Data summarized from species accounts presented in Wilson (1995).

Leaf litter, fallen logs, moist soils,and other surface debris serve asrefuges from drying conditions. Theringed and streamside salamandersuse moist soil, while the flatwoodsand Jefferson salamanders use leaflitter. Fallen logs provide an importanthabitat component for the marbledand mole salamanders. Several species,including the spotted and Mabee’ssalamanders, also prefer closed-canopyconditions adjacent to water sources.

Table 5.6 shows the associationsbetween 23 vegetative cover types(following Hamel 1992) and salaman-ders in the South. Mesic, mixed pine,and hardwood forests support 72percent of species, including ringed,marbled, and mole salamanders.Sixty-four percent of the salamandersoccupy mesic, upland hardwoods.These species include streamside,smallmouth, seepage, and duskysalamanders. White pine-hemlockand bottomland forests are used byslightly less than half of the southernsalamanders. Jefferson, spotted,and green salamanders occupy whitepine-hemlock forests, while severalamphiuma species are found inbottomland hardwood forests. Xericoak-hickory forests also support avariety of salamanders.

Salamander diversity appears to beless on the Coastal Plain than in theAppalachian Mountains. The formerhas much sandy, well-drained soil,high summer temperatures, and higherdensities of predators (Echternachtand Harris 1993).

Connectivity between preferred foresthabitats reduces population isolationand promotes dispersal (Wilson 1995),a management concern for manyamphibian species. Many salamandersare adapted to travel only shortdistances in response to habitatalteration, while others with restrictedgeographic ranges become imperiled ifhabitat modification is rapid enough topreclude dispersal to similar habitats(Gibbons and Buhlmann 2001).

Table 5.7 illustrates relationshipsbetween salamander occurrence andforest successional stage. The seralstages follow those used by Hamel(1992): grass-forb, seedling-sapling,poletimber, and sawtimber. Note thatnot all cover types contain each seralstage. The Everglades type, for example,only exists in the grass-forb stages.

Most salamander species find optimumhabitat conditions in sawtimber stands.

Frogs and toads—The South isinhabited by numerous species of frogsand toads, each with its own particularrequirements. The region supportssuch diversity due to its warm, humidclimate, diversity of vegetative com-munities, and abundance of aquaticenvironments, particularly wetlands.

Wilson (1995) places these speciesinto: (1) terrestrial species that migrateto standing water for egg deposition,(2) semiaquatic species requiringterrestrial habitat for dispersal, and (3)aquatic species that may use terrestrialhabitat during rainy conditions. Eachspecies requires standing water for eggdeposition and larval development.

Several species exhibit two distinctstages: an aquatic larval stage (tadpole)and an adult stage. The eggs developinto tadpoles, which then undergoa complex metamorphosis intoadults. The two stages have differenthabitat requirements that influencedistributions and habitat associations.

Tadpoles consume algae and bacteria,while adult frogs and toads rely uponinvertebrates. Some species, such asthe pig frog, remain semiaquatic asadults, while others become terrestrial.Frogs and toads are important preyfor wading birds, raptors, foxes,raccoons, and snakes.

Moisture also is a limiting factorfor most frog and toad species; eventerrestrial species require moistmicrohabitat (table 5.5). In addition


TERRESTRIAL

Table 5.7—The relationship between forest successional stage andamphibian occurrence in the South a

Taxa subgroup/ Grass/ Seedling/ Pole- Saw-habitat conditions forb sapling timber timber

- - - - - - - - - - - Number of species - - - - - - - - - - -Salamandersb

Optimalc 0 0 0 76Suitabled 10 15 25 5Marginald 6 50 54 0

Frogs and toadse

Optimal 15 0 0 7Suitable 16 34 31 14Marginal 8 4 5 13

a Summarized from Wilson (1995).b Based on habitat relationships information from 92 species.c Habitats in which the species occurs with highest frequency.d Habitats in which the species occurs with successively lower frequency.e Based on habitat relationships information from 40 species.

The change in successional stage fromsawtimber to grass-forb that resultsfrom timber harvest may enhancehabitat suitability for one species, yetcreate marginal habitat for another.

Amphibian declines—Reporteddeclines of amphibian populations havedrawn considerable attention over thepast two decades. Many are associatedwith high elevation, pristine areasthat are remote from surroundinglandscape modification. Amphibiansare particularly sensitive to theirenvironment. Their permeable skinand the lack of protective eggshellsmake them vulnerable to toxicantspresent in soil and water.

Southern species showing evidenceof declines include the flatwoodssalamander, Red Hills salamander,Texas blind salamander, wood frog,southern dusky salamander, andgreen salamander. Numerous othersare categorized as imperiled andvulnerable (chapter 1). Endemicspecies are of particular concernin the Edwards Plateau, OzarkHighlands, Atlantic Coastal Plain,and Appalachian Mountains.

Amphibian declines have beenattributed to several factors. Theseinclude habitat loss, wetland alteration,climate changes leading to droughts,diseases, exotic species, and agriculturalchemicals. Other factors include acidprecipitation and ultraviolet radiation.These are briefly reviewed later.

to broad stream margins, frequentlyused habitats include both permanentand seasonal swamps and ponds.Many species, including the Americantoad and southern cricket frog, requiremoist soils for burrowing.

Leaf litter, potholes, and aquaticvegetation often provide moisture(table 5.5). The oak toad and pinebarrens tree frog use leaf litter, whilethe southern chorus frog and the bird-voiced tree frog use aquatic vegetation.Potholes provide an important habitatcomponent for Brimley’s chorus andsouthern leopard frogs (Wilson 1995).Species that prefer open-canopyconditions include the Houstontoad and the northern cricket frog.

Although wetlands are importantbreeding habitats, many frog andtoad species spend part or all of theirnonbreeding season in trees andshrubs. Forest structure creates diversehabitats with many niches. Forests alsocontribute organic matter and moderatethe temperature and evaporation rateof adjacent aquatic habitats.

Southern frogs and toads inhabit awide variety of forest cover types (table5.6). Mesic, mixed pine and hardwoodforests support 83 percent of species,including the American toad, Cope’sgray tree frog, and northern cricketfrog. Seventy-eight percent of thespecies inhabit bottomland hardwoodforests, including Woodhouse’s toads,pine woods tree frogs, squirrel treefrogs, and gray tree frogs. Longleafpine, cypress-tupelo, and bay-pocosinhabitats are used by over half of the frogspecies in the region. Oak and southerntoads and southern cricket frogs occupylongleaf pine forests, while several treefrogs are characteristic of cypress-tupeloassociations. It appears that a majorityof species finds optimum and suitablehabitat conditions in the grass, sapling,and poletimber stages (table 5.7).

Habitat management foramphibians—The complex life cycleof frogs and toads requires managementof both terrestrial and aquatic habitats.Tiger salamanders and other ambys-tomas breed in the water but remainterrestrial during nonbreeding season.Thus, providing only one habitatcomponent would fail to maintainviable populations of these species.Some terrestrial species require pondsor other standing water during thebreeding season. Consequently, theremoval of barriers such as roads

between terrestrial habitat andaquatic habitat is important.

The semiaquatic species requireaquatic areas that have rocks, woodydebris, or other similar shelter in thewater. Emergent and floating vegetationis important for breeding of somespecies. The adjacent terrestrial habitatalso is important because many species,such as the Eurycea and Desmognathusgenera, spend significant portions oftheir lives foraging and occupyingterrestrial areas. Buffers adjacent tostreams provide access to uplandforested habitats. Aquatic habitatsshould be protected against thermalchanges, water pollution, and excessivesiltation (Wilson 1995). Habitatalteration due to dredging, channel-ization, and impoundment canbe detrimental to many species.

Forest management alters thevegetative composition, seral stage,and structure of amphibian habitat.For example, prescribed burningtemporarily removes leaf litter, herba-ceous cover, and woody understoryvegetation. Vegetative structure, snags,loose bark, and surface debris areimportant factors in managingamphibian habitat. Disking, wind-rowing, and furrowing during someforestry operations (Gibbons andBuhlmann 2001) may negatively impactspecies dependent upon the understory.Conversion from one forest type toanother may be beneficial to somespecies and detrimental to others.


Wetlands and vernal pools areimportant for several amphibians.There have been significant lossesof wetlands in the last two centuries(chapter 1). Declines in wetland qualitythrough eutrophication, pollution, andfish stocking also impact amphibianpopulations. The hellbender is affectedby stream degradation, while thegopher frog is influenced by theconversion of pine and hardwoodforests to tree plantations, agriculture,and urban uses. In addition, habitatfragmentation by roads contributes tothe mortality of breeding adults anddispersing juveniles (Wilson 1995).

Ozone depletion in the upperatmosphere increases the amountof ultraviolet radiation on the Earth’ssurface, particularly at high elevations.Ambient radiation damages cellularDNA (Reaser and Johnson 1997);amphibians with low levels ofphotolyase enzyme have embryosthat are susceptible to ultravioletradiation, which causes mortalityand abnormal development, includingskeletal, eye, and skin deformities.

Their porous skin makes amphibianssusceptible to herbicides, pesticides,heavy metals, and petroleum productsin aquatic systems. Pollutants such asgasoline, oil, and antifreeze sometimesoccur in runoff into amphibian habitat.Relatively high nitrate levels causephysical and behavioral abnormalitiesin a number of species; syntheticchemicals interfere with hormonalprocesses, inhibiting amphibiandevelopment (Reaser 1996). Theapplication of fertilizers and pesticides,particularly by aerial spraying, oftenimpacts amphibians far from the pointof application in nontarget areas.

The introduction of exotic species,such as fish, crayfish, and bullfrogs,into lakes and wetlands also influencesamphibian populations. Fish intro-duced into wetlands for mosquitocontrol prey upon amphibian eggsand larvae. Chytrid fungi, trematodeparasites, and viruses carried byexotic fish may also contribute topopulation declines.

Several of the factors discussedabove have been implicated as causesof amphibian abnormalities. Theseinclude parasite infestation, toxincontamination, radiation, radioactivesalts, ground-level ozone, excessiveheating of eggs, and reformulated

gasoline (Reaser and Johnson 1997).Of these, only the parasite, toxin,radiation, and predation hypotheseshave supportive evidence. Thefrequency of malformations ishighest in frogs that have recentlymetamorphosed from tadpoles.

Concern about the status ofamphibian populations is clearlywarranted. Physiological constraints,limited mobility, and changes in sitecharacteristics hinder recolonizationof sites of local extinction. Thetemporal and spatial populationdynamics of many amphibians arenot well understood; it is unknownwhether observed declines exceednatural population fluctuation.

There are other concerns facingindividual amphibian species. Manyof these are discussed in the section onreptiles, as these concerns are shared byherpetofauna as a group. In addition,some of these concerns are mentionedin Species accounts presented next.

Species accounts—The followingare accounts for selected amphibianspecies that are of concern in the South.Several are federally listed as threatenedor endangered. Others are classified asimperiled or vulnerable by NaturalHeritage agencies. The species accountsand management recommendationsfollow Wilson (1995) unless other-wise noted.

Species accounts: flatwoodssalamander—The population of thisthreatened species has declined duringthe past 10 to 15 years (Wilson 1995).The cause of the decline is uncertain,and the salamander is uncommonthroughout its range from SouthCarolina, southern Georgia, andFlorida westward to Mississippi.

The salamander inhabits pine flat-woods dominated by longleaf and slashpines and wiregrass, which is importantfor egg disposition. It is often foundin association with cypress ponds,swamps, and pitcher plant bogs thatare used for reproduction.

Management activities focus onavoidance of intensive site preparationbefore harvest, avoidance of prescribedburning during peak surface activityand breeding periods (Novemberthrough April), and protection ofbreeding ponds. Fish stocking shouldbe avoided (Bury and others 1980).

Species accounts: Florida bogfrog—This species is classified asimperiled by Natural Heritage and isa species of special concern in Florida(NatureServe 2000). The frog iscurrently known to exist in 23 localitiesin the Panhandle (Moler 1992b). Manyare found on the Eglin Air Force Base.

This frog species inhabits nonstagnantacidic seeps and the shallow backwatersof larger streams. It is frequently foundin association with sphagnum mossand early seral stages of Atlantic white-cedar. Shrubby streamside habitats thatdo not have developing hardwoodforests are preferred. The frog’sdiet consists of insects and othersmall arthropods.

Stream contamination andimpoundment, and forest successionthreaten the survival of this species(Moler 1992b). Conservation actionscenter on the protection of suitablehabitat. Management of streamsidevegetation to maintain the shrub-bogcommunity is advised.

Species accounts: gopher frog—This uncommon species is classifiedas vulnerable by Natural Heritage(NatureServe 2000). Alabama, Florida,Georgia, Mississippi, North Carolina,and South Carolina list the frog as ofspecial concern. The gopher froghistorically was distributed along theGulf and Atlantic Coastal Plain, withisolated populations in the Valleyand Ridge Province of Alabama. Itwas last documented from Louisianain the 1960s. The frog has declinedthroughout its range with the lossof longleaf pine habitat (Martof andothers 1980).

The gopher frog is associated withsandy pine flatwoods, turkey oak-pinesandhills, and xeric hammocks. Itbreeds in shallow, temporary pondswith open canopies and emergentherbaceous vegetation. Ditches andborrow pits are occasionally used.Adults seek refuge in the burrows ofgopher tortoises, mice, and crayfish.Stump holes, root mounds, densegrass clumps, and thick mats of leaflitter may also be used.

The frog is an opportunistic feederwith a diet of arthropods, small frogs,and toads. Predators include watersnakes, turtles, bluegills, andmosquitofish.

Management centers on protection ofthe sandhills and scrub-oak ecosystems


TERRESTRIAL

and halting the losses of this habitatto circular irrigation farming andindustrial development. Prescribedburning and other managementpractices that retain the open scrubnature of this habitat benefit thisspecies (Wilson 1995), while practicesthat drain or alter breeding pondsare detrimental.

Species accounts: greensalamander—This species is classifiedas vulnerable by Natural Heritage and isa species of special concern in Alabama,Georgia, Mississippi, and NorthCarolina (NatureServe 2000).Impoundment of several rivers inthe Carolinas has extirpated severalknown populations.

The unique habitat of this speciesis limited and localized. The greensalamander lives in damp crevices inshaded rock outcrops and under thebark of cove hardwood trees. It also isfound in upland areas of Virginia pineand white pine-hemlock with mountainlaurel understories. The salamander’sdiet consists of small insects, spiders,and earthworms.

Conservation efforts focus onprotection of rock outcrops and theestablishment of buffer zones in areasof timber harvest.

Species accounts: Houston toad—This endangered species is restrictedto southeastern Texas, where itspopulation is very small and frag-mented. Human alteration of naturalwatersheds has eliminated many ofits natural breeding pools, resultingin hybridization with the Gulf Coasttoad and the Woodhouse’s toad(Wilson 1995).

This toad inhabits areas with sandy,friable soils and is found most oftenin loblolly pine or mixed deciduoushabitats interspersed with grassy areasunder a range of conditions. Breedinghabitats include roadside ditches,temporary ponds, and other seasonallyflooded low spots. The toad’s dietconsists primarily of insects.

The recovery plan requires protec-tion of critical habitat for this species.Habitat is maintained in a pristine state,and several breeding projects have beenattempted. Development projects havebeen regulated in areas designated ascritical habitat (Brown 1975).

Species accounts: one-toedamphiuma—This species is classifiedas vulnerable by Natural Heritage and

listed as rare in Georgia (NatureServe2000). It occurs in restricted geographicareas in northern Florida, MobileBay in Alabama, and the OchlockneeRiver drainage in Georgia.

This semiaquatic salamander requiresmucky habitats in association withpermanent streams (Means 1992).Management actions center onprotection of muck areas, whichare threatened by sand and siltsedimentation during constructionactivities. Other actions include theregulation of amphiuma collection.

Species accounts: Red Hillssalamander—This species is listed asthreatened at both the Federal and Statelevel. It is confined to a narrow beltwithin the Tallahatta and Hatchetigbeegeological formations in the Red Hillsof Alabama (Wilson 1995).

This nocturnal salamander lives inburrows on high, steep, uncut slopeswith high soil moisture content andfull tree canopy (Dodd 1991). Theburrows are often near the base of atree or under siltstone outcroppings.The salamander feeds on spiders andinsects (U.S. Fish and Wildlife Service1983, 1990b).

The majority of land in its rangeis privately owned. Habitat has beenreduced by timber harvest, conversionof forest to agriculture, and ridgetopclearing. Overcollecting may havecaused a decline in some areas. Feralhogs are a threat in localized areas(NatureServe 2001).

Conservation actions includecooperation with private and corporatelandholders to restrict clearcutting andheavy site disturbance. Under publicownership, two areas have been setaside to support limited populations(NatureServe 2001). These includeAlabama Forestry Commission andU.S. Army Corps of Engineers lands.In 1991, International Paper Companyinitiated work on a Habitat Conser-vation Plan (HCP) for this species.Other companies subsequentlydeveloped HCPs (Bailey 1995).Research needs include determinationof the microhabitat effects of timbermanagement practices and thecollection of data on reproductiveviability and recruitment withinexisting populations.

Species accounts: Shenandoahsalamander—This species isendangered due to restricted range,

habitat modification, and competitiveinteractions with the redbacksalamander. Inhabiting the high-elevation mountains of Virginia, thespecies requires talus slopes with deepsoil pockets in mixed coniferous anddeciduous forests. Its diet consists ofsmall arthropods and earthworms.

Conservation efforts includerestriction of construction activities thatcould disturb the limited talus habitatsof this salamander (Martof and others1980). Any construction of trails, roads,or overlooks in the ShenandoahNational Park should be carefullymonitored so as not to impact thissalamander’s limited habitat.

Species accounts: Tennessee cavesalamander—This species is classifiedas imperiled by Natural Heritage andis listed as endangered in the States ofAlabama and Tennessee (NatureServe2000). The salamander is found inpermanent streams and pools inlimestone caves of central and south-west Tennessee, northern Alabama,and extreme northwest Georgia. Itis believed to occur in approximately1 percent of the caves in its range.

The Tennessee cave salamander feedson arthropods, other small aquaticinsects, and earthworms. Managementcenters on restricting human access andprotecting limestone cave habitat. Thespecies is very sensitive to pollutantsand disturbances within its habitat(Wilson 1995).

ReptilesThe reptiles of the South belong to

three orders: Crocodilia (alligators andcrocodiles), Squamata (lizards, amphis-baenians, and snakes), and Testudinata(turtles). The South supports a diversityof reptiles (fig. 5.5), including 89snakes (11 endemic), 75 lizards (6endemic), 29 turtles (13 endemic),and 4 other reptiles (including 2crocodilians). The number of reptilespecies ranges from 155 in Texas to54 in Kentucky (NatureServe 2000).Species richness is impressivein Florida (94), Alabama (87),Georgia (87), and Mississippi (86).

The South’s Coastal Plain possessesNorth America’s highest diversity ofreptiles (Gibbons and others 1997,White and others 1998). Twenty-ninepercent of southern reptiles areclassified as endemic (Dodd 1995a).


Snakes89 (45%)

Lizards75 (38%)

Turtles29 (15%)

Other reptiles4 (2%)

Table 5.8—Reptile species within the South that are listed as threatened or endangered


TurtlesCaretta caretta Loggerhead (T) AL, FL, GA, LA, MS, NC, SC, TX, VAChelonia mydas Green turtle (E) AL, FL, GA, LA, MS, NC, SC, TX, VAClemmys muhlenbergii Bog turtle (T) GA, NC, SC, VADermochelys coriacea Leatherback; tinglar (E) AL, FL, GA, LA, MS, NC, SC, TX, VAEretmochelys imbricata Hawksbill (E) AL, FL, GA, LA, MS, NC, SC, TX, VAGopherus polyphemus Gopher tortoise (T) AL, LA, MSGraptemys flavimaculata Yellow-blotched map turtle (T) MSLepidochelys kempii Kemp’s or Atlantic ridley (E) AL, FL, GA, LA, MS, NC, SC TX, VAPseudemys alabamensis Alabama red-belly turtle (E) ALSternothesis depressus Flattened musk turtle (T) AL

LizardsEumeces egregius lividus Bluetail mole skink (T) FL

SnakesDrymarchon corais couperi Eastern indigo snake (T) AL, FL, GA, MS, SCNerodia clarkii taeniata Atlantic salt marsh snake (T) ALNerodia paucimaculata Concho water snake (T) TX

Other reptilesAlligator mississippiensis American alligator (T) AL, AR, FL, GA, LA, MS, NC, OK, SC,

TXCrocodylus acutus American crocodile (E) FL

T = Threatened; E = endangered.Source: U.S. Department of the Interior (2000).

Reptiles occupy forest, freshwater,marine, and urban habitats. Most usethe same habitat for breeding andnonbreeding activities, but aquaticand marine species require adjacentterrestrial habitats in order tosuccessfully reproduce.

The forested mountain regions alsosupport an abundance of reptiles,including worm snakes, copperheads,

ringneck snakes, bog turtles, and coalskinks. The longleaf pine-wiregrasscommunity is vital habitat for thegopher tortoise and important habitatfor mole skinks, glass lizards, scarletsnakes, pine snakes, and coachwhipsnakes. Cypress-gum swamps arehome to rainbow snakes, mudsnakes, western green watersnakes,and striped crawfish snakes (Gibbonsand Buhlmann 2001). Some reptilesplay important roles in southerncommunities in nutrient cycling.Their burrows provide refugesfor other species during extremeclimatic conditions.

The numbers of turtles in Mississippi(31), Texas (30), Alabama (30), Georgia(27), Louisiana (26), and Florida (26)reflect the abundance of coastal andfreshwater habitats. Numbers of lizardspecies in Texas (51) and Florida (38)far surpass the richness in otherSouthern States (NatureServe 2000).Both States are relatively large andhave a wide variety of habitats in them.The number of lizards residing in theremaining States ranges from 17 speciesin Oklahoma to 8 species in Kentucky.

The number of snakes tends to behighest in the southernmost CoastalStates. There are 73 in Texas, 46in Florida, 42 in Alabama, 42 inMississippi, 41 in Georgia, and 41in South Carolina. Species richness inthe mountain States is slightly lower.Virginia supports 30 snake species.Snakes reach their highest diversitiesin southern forests and their peripheralhabitats, such as rivers, streams, andisolated wetlands.

Seventeen species of reptiles are listedas threatened or endangered by theU.S. Fish and Wildlife Service (table5.8). In addition, numerous reptilesare classified as imperiled or vulnerableby the Natural Heritage agencies(chapter 1). Many of these speciesoccur on the Coastal Plain; severalare narrowly restricted endemics.

Although the variation in life-historytraits is quite marked, many speciesof reptiles have long lives, variablereproductive rates, and high mortalityamong eggs and neonates. Suchcombinations of life-history charac-teristics are particularly commonamong turtles, crocodilians, and snakes.

Figure 5.5—Species richness by majorsubgroups of reptilian taxa occurringwithin the South (NatureServe 2000).


TERRESTRIAL

Table 5.9—The relationships of reptiles to physiographic features and other habitat elements a

Alligators/ Habitat element Turtles Lizards Snakes crocodiles

No. % No. % No. % No. %Physiographic feature

Sandhills 1 2 8 40 14 25Flatwoods 1 2 4 20 12 21Narrow stream margins 2 5 1 5 7 12Broad stream margins 11 27 1 5 18 32 1 50Swamps 14 34 0 0 13 23 1 50Cypress strands 2 5 0 0 8 14Cypress ponds 3 7 0 0 6 11Cypress drains 1 2 0 0 4 7Willow heads 1 2 0 0 3 5Bays and pocosins 3 7 1 5 9 16Rivers and streams 21 51 0 0 10 18 1 50Permanent ponds 12 29 0 0 9 16 1 50Vernal ponds 3 7 0 0 2 4Lakes 15 37 0 0 8 14 2 100Marshes 13 32 0 0 16 28 1 50

Specific requirementClosed canopy 0 0 0 0 5 9Open canopy 29 71 14 70 43 75 2 100Forest openings 3 7 10 50 8 14Shrub thickets 1 2 2 10 4 7Moist soil 5 12 3 15 11 19Sandy or friable soils 36 88 12 60 18 32Leaf litter 3 7 11 55 34 60Snags 0 0 4 70 3 5Fallen logs 0 0 11 55 35 61Rock outcrops 0 0 4 70 4 7Crevices and/or caves 0 0 1 5 5 9Seepages 4 10 0 0 1 2Potholes 1 2 0 0 3 5Aquatic rocks and/or logs 23 56 0 0 12 21 1 50Aquatic vegetation 17 42 0 0 13 23 2 100


Due to their ectothermic physiologyand seasonal inactivity, reptiles haverelatively slow growth rates, advancedages at maturity, and advancedgeneration times. Lizards have theyoungest ages at maturity (1.5 years),while turtles and crocodilians have theoldest age at maturity (20 to 50 years).Age at maturity is estimated at over 30years for some marine turtles.

Rates of reproduction are variable.Clutch frequency in sea turtles variesfrom one to four clutches every 3to 4 years, whereas annual multipleclutches are common for somefreshwater turtles. Reproductionoccurs in alternate years (or less often)for terrestrial tortoises. Most lizards

produce at least one clutch each year,and multiple clutches per year arecommon. In contrast, biennialreproduction is typical in snakes.

Turtles—Six turtle families are foundin the South. These include the seaturtles, snapping turtles, water andbox turtles, mud and musk turtles,tortoises, and softshell turtles. Thegreatest diversity occurs in the CoastalPlain, which supports a variety offreshwater and coastal marsh speciesand several species of sea turtles.

The gopher tortoise is a keystonespecies in the communities where itoccurs. Its burrows provide refuges fora variety of species, including indigosnakes and diamondback rattlesnakes.

The tortoise is threatened throughoutits range as a result of habitatdestruction associated with landdevelopment (Echternacht andHarris 1993).

Turtles are scavengers, herbivores,and carnivores and often contributesignificant biomass to variousecosystems. They provide dispersalmechanisms for plants, contributeto environmental diversity, and fostersymbiotic associations with a diversearray of organisms.

Many species have experiencedsignificant declines in abundance anddistribution during the last century.Among such species are the bog turtle,spotted turtle, common box turtle,


gopher tortoise, common slider, andalligator snapping turtle. Some species,such as map turtles, have limitedranges, placing them at risk fromhabitat alteration or illegal collectionfor the pet trade. Disease also appearsto contribute to population declinesin some turtles. The diamondbackterrapin was exploited heavily forfood during the 19th century. Althoughthe species recovered, the terrapinis again imperiled due to regionalharvesting, habitat destruction,vehicular mortality, and drowningin crab traps (Lovich 1995).

Although the habitat requirementsof marine turtles are beyond the scopeof this terrestrial assessment, concernsover the future of these species warrantsmention. Five species of marine turtlesfrequent the beaches, bays, estuaries,and lagoons of the South: loggerhead,green, Kemp’s ridley, leatherback, andhawksbill turtles. These species havehad dramatic declines attributable tocommercial turtle fishing, exploitationof the juvenile populations, beachdevelopment, polluted water,incidental take, and diseases such asfibropapillomas. Monitoring is difficultdue to their longevity. They continue tobe threatened and their conservationinvolves international efforts.

Forest conditions influence bothaquatic and terrestrial turtles. Mapturtles, cooter turtles, and musk turtlesinhabit streams and rivers that areinfluenced by adjacent riparian forests.Forest cover reduces sedimentationrates, affects water temperature, andinfluences availability of basking sites(Gibbons and Buhlmann 2001). Inaddition, many species such as mudturtles use terrestrial habitat for nestingand winter dormancy, spending thesummer in wetland areas. Riparianforests are also quite importantfor map turtles.

Turtles inhabit areas with a varietyof physiographic features (table 5.9).Not surprisingly, rivers, streams,swamps, lakes, and marshes figureprominently in their occurrence.Ninety percent of the species dependon aquatic environments. Commonand alligator snapping turtles are foundin swamps, deep rivers, and canalswhile marshes support bog and paintedturtles. River and stream habitatssupport several species of map turtles.

Turtles are also associated with sandysoils, logs, and rocks that serve as

shelter and as basking surfaces. Theornate box turtle and gopher tortoiserequire sandy or friable soils in whichto burrow or deposit eggs (Wilson1995). A majority of species (71percent) prefers open-canopyconditions that aid in thermoregulation.Such species include the painted turtle,spotted turtle, Alabama map turtle,and striped mud turtle.

Tables 5.10 and 5.11 list thevegetative cover types and successionalstages that are associated with turtlesin the South. Bottomland hardwoodforests support 81 percent of species,including the wood turtle, the commonmap turtle, and the Pascagoula mapturtle. Sixty-three percent of the turtlesoccupy cypress-tupelo forests. Thesespecies include the Barbour’s, theEscambia, and the yellow-blotchedmap turtle. Mesic, mixed pine-hardwoods stands are used by slightlyover half of the southern species,including the painted and spottedturtles. Approximately one-third ofthe species find optimum or suitablehabitat in grass-forb cover; mostof these species presumably areassociated with aquatic conditions.

Lizards and snakes—Four familiesof lizards inhabit the South. Thesespecies include anole, fence, collared,and horned lizards; whiptails; skinks;and glass lizards. All lizards areterrestrial; most species have smallhome ranges.

Sandhills and flatwoods are importanthabitats for lizards (table 5.9). TheFlorida scrub lizard, the island glasslizard, and the coal skink inhabit theseareas. Leaf litter, fallen logs, and snagsprovide shelter as well as places to huntfor prey. The fence lizard and five-linedskink are associated with snags, whilethe slender glass lizard and the broad-head skink use fallen logs. Friable soilsare an important habitat component for60 percent of the species. Mimic andeastern glass lizards deposit eggs andburrow in these soils. The majority ofspecies (70 percent) require an openforest canopy, a forest opening, ora rocky outcrop as basking sites forthermoregulation. Such species includethe slender and island glass lizards,the collared lizard, and the GreatPlains skink.

Twenty-one forest cover types areassociated with lizards in the South(table 5.10). Although lizards in theregion use a variety of forest cover, over

half of the species inhabit longleaf pine-scrub oak, xeric mixed pine-hardwood,and live oak stands. Longleaf pine andscrub oak forests support the fencelizard, island glass lizard, and mimicglass lizard. The sand skink, groundskink, and six-lined racerunner occurin mixed pine and hardwoods. Liveoak forests are used by 55 percent ofthe southern lizards, including themole skink and broadhead skink.Mesic, mixed pine-hardwood standsalso support a variety of lizards.Approximately half of the speciesfind optimum conditions amonggrasses and forbs (table 5.11).

Three families of snakes occur inthe South: nonvenomous snakes,coral snakes, and pit vipers. Speciesthat inhabit the water are especiallyprevalent. Three of the largest snakesin North America occur in the region:the indigo snake, eastern diamondbackrattlesnake, and timber rattlesnake.

In the absence of a large assemblageof mammalian predators, snakesassume special importance as toppredators in some communities, andtheir low metabolic rates allow themto occur at impressive densities inundisturbed habitat (Echternachtand Harris 1993).

Since the larger species of snakeshave fairly large home ranges—125to 250 acres—fragmentation of existinghabitat poses a significant threat(Wilson 1995). Several snakes, suchas the brown snake and the commongarter snake, have significantpopulations in suburban areas.

The majority of snake and lizardspecies have become imperiled dueto insular populations, restrictedranges, habitat degradation, or theloss of suitable habitat. Maliciouskilling, biocides, exotic species, andillegal trade have also contributedto their decline.

Many snakes require shelter in theform of friable soil, fallen logs, leaflitter, rocks, or similar surface debris(table 5.9). As with lizards, snakesrequire open-canopy forest conditionsto aid thermoregulation. Habitatmanagement that leaves surfacedebris and tree stumps can benefittheir habitat. Leaf litter and fallen logsprovide refuges for snakes as well astheir prey, which include invertebrates,small mammals, and amphibians. Racerand ringneck snakes are found in leaf


TERRESTRIAL

Table 5.10—The relationship between forest cover type and reptile occurrence in the South a

Alligators/ Forest cover types Turtles Lizards Snakes crocodiles

No. % No. % No. % No. %

Everglades 8 20 2 10 13 23 1 50Tropical hardwoods 5 13 4 20 13 23 1 50Mangroves 5 13 2 10 3 5 2 100Longleaf-slash 2 5 4 20 11 19Pine-flatwoods 3 7 8 40 23 40Virginia-pitch pine

(xeric upland pines) 1 2 5 25 14 25Longleaf pine 4 10 15 19 21 37Loblolly-shortleaf pine 5 12 7 35 20 35White pine-hemlock 4 10 2 10 16 28Pond pine 2 5 2 10 4 7Longleaf-scrub oak 2 5 10 50 13 23Mixed, pine-hardwood

(mesic) 22 54 8 40 37 65Mixed, pine-Hardwood

(xeric) 3 7 11 55 15 26Spruce fir 0 0 0 0 1 2Upland hardwoods (mesic) -

white oak-red oak 9 22 3 15 25 44Cypress tupelo 26 63 3 15 12 21 1 50Bottomland hardwoods-

(sweetgum-willow oak) 33 81 5 25 29 51 1 50Sweetgum-yellow-poplar 16 39 1 5 9 16Bay-pocosin 9 22 3 15 10 18Live Oak (maritime) 1 2 11 55 22 39Maple-beech 4 10 1 5 16 28Cove hardwoods 0 0 0 0 3 5Spartina 7 17 0 0 3 5 2 100Elm-ash 0 0 0 0 5 9Oak-hickory (xeric

hardwoods) 4 10 3 15 24 42Aquatic dependent 37 90 0 0 15 26


litter, while fallen logs are importanthabitat components for indigo andcorn snakes. Seventy-five percentof snake species are associated withopen canopy forest; these includescarlet and Kirtland’s snakes.

Forests provide essential habitatcomponents for terrestrial speciesas well as those that live in aquatichabitats. Table 5.10 presents the26 vegetative cover types that areassociated with snakes in the South.Snakes use a diversity of forest cover;there are, however, a group of specifictypes that are used most often. Mesic,mixed pine and hardwood forestssupport 65 percent of species,

including the western worm snake,corn snake, and rat snake. Fifty-onepercent of the snakes occupybottomland hardwoods. These speciesinclude the mud, rainbow, and scarletking snake. Mesic upland hardwoodsand xeric oak-hickory hardwoods areused by over 40 percent of the southernspecies. The prairie king snake, milksnake, and the common water snakeoccupy mesic hardwood types, whileeastern and southern hognose snakesare characteristic of xeric hardwoods.The pine flatwoods forests also supporta variety of snakes.

Table 5.11 illustrates the relation-ships between snake occurrence

and forest successional stage. Approx-imately half of the species find suitablehabitat in seedling-sapling andpoletimber conditions. Slightly morespecies find optimum and suitableconditions in grass-forb stages thanin sawtimber stands.

Crocodilians—Two native speciesof crocodilians, the America alligator(family Alligatoridae) and the Americancrocodile (family Crocodylidae), occurin the South. A large breedingpopulation of the introduced spectacledcaiman, native to the American tropicsfrom southern Mexico to Argentina,occurs in Dade County, FL (Echternacht


Table 5.11—The relationship between forest successional stage and reptileoccurrence in the South a

Successional stage

Taxa subgroup/ Pole- Saw-habitat condition Grass Sapling timber timber

Turtlesb

Optimalc 13 0 0 0Suitabled 3 8 8 4Marginald 0 7 5 8

Lizardse


Snakesf


Alligators and crocodilesg


a Summarized from Wilson (1995).b Based on habitat relationships information from 41 species.c Habitats in which the species occurs with highest frequency.d Habitats in which the species occurs with successively lower frequency.e Based on habitat relationships information from 20 species.f Based on habitat relationships information from 57 species.g Based on habitat relationships information from 2 species.

and Harris 1993). This exotic speciesis discussed further in chapter 3.

The alligator is a wide-ranging animalthat occurs from coastal North Carolinasouth to Florida and westward toeastern Texas. It has recovered fromprevious declines and now has peststatus in Louisiana and Florida. Thealligator creates marsh pools thatprovide habitat for many other species.Its larger and more secretive relative,the American crocodile, is restricted inits North American range to extremesouth Florida.

During the last century, wetlanddrainage for agriculture and develop-ment activities permanently reducedalligator populations in freshwatermarshes. Recent environmentalcontamination has been associatedwith declines in alligator populations(Woodward 1995). Widespreadpollution of wetlands by toxic petro-chemicals and metals may continue tothreaten population viability. Although

the status of the Florida alligatorpopulation appears secure, continuedhabitat loss and toxic contaminationmay compromise its conservation.

The crocodile remains endangered,while the alligator is federally listedas threatened due to “similarity ofappearance.” This designation reflectsthe special instance when a species soclosely resembles a listed species thatit is difficult in the wild to differentiatebetween the two. The effect of thisdifficulty is an additional threat tothe listed species.

The alligator is doing well insuitable habitat, while the crocodileis struggling to survive in its limitedrange in southern Florida (Wilson1995). Management plans for bothspecies protect aquatic and terrestrialhabitat, particularly for nesting andbasking. Management includes captiveprograms to manage the species formeat and hide production, as well aseffective protection from poaching.

Not surprisingly, both species occurin areas limited in the number ofphysiographic features and vegetativecover types (tables 5.9, 5.10, and5.11). Lakes, marshes, rivers, streams,permanent ponds, and swamps figureprominently in their occurrence.Aquatic vegetation is important toboth species; rocks and logs in thewater serve as useful basking areas.Alligators and crocodiles requireopen canopy forest conditions toaid thermoregulation. This needmay explain their use of stands ofgrasses and forbs.

Six vegetative cover types areassociated with these species in theSouth. Mangrove and spartina habitatsupports both species. Tropicalhardwoods, cypress tupelo, andbottomland hardwood forests arealso occasionally used. Additionaldetails on each species are presentedin the Species accounts at the endof the reptile section.

Habitat management for reptiles—The general problems faced by reptilesin southern forests center on theenvironmental impacts resultingfrom human activities. Difficultiesin assessing problems and monitoringpopulations hinder managementof these vertebrates.

The life history and ecology of reptilesdiffer markedly from those of othertaxa. Many reptile species take longerto mature and have long lifespans.For example, the forest-inhabiting boxturtle and snapping turtle take over 10years to reach sexual maturity (Gibbonsand Buhlmann 2001). Managing forsustainable populations of long-lived,late-maturing species requires differentstrategies than for short-lived, rapidturnover species (Congdon and others1994, Ernst and others 1994).

The primary threats to reptiles inthe South stem from habitat destructionand alteration, including changes inwater quality. The drainage of wetlandsand temporary ponds has reducedthe population of striped newts(Dodd 1995a) and extirpated theflatwoods salamander from a portionof its range (White and others 1998).Destruction of wetlands has reducedspotted turtle populations, and otheraquatic habitats do not meet theturtle’s specialized needs.

Impoundments have affected severalspecies of map turtles native to large


TERRESTRIAL

southern rivers. The damming ofstreams to form reservoirs hascontributed strongly to the eliminatingseveral species (Mitchell 1994). Inaddition, the removal of dead trees andthe dredging of river bottoms, whichharbor mollusks that the turtles eat,have negatively affected these species.

The gopher tortoise and other reptileshave become threatened in part becauseof the loss of longleaf pine habitat(Dodd 1995b, Guyer and Bailey 1993).Many species of snakes and box turtlesare also declining in numbers due toloss of suitable habitat. Accidentaldeath by vehicles and intentional killingare other factors contributing to snakedecline. Several of these reptiles, suchas short-tailed snakes and flattenedmusk turtles, have relatively smallgeographic ranges. Others, such asthe pinewoods snake, coal skink, andWebster’s salamander, have disjunctpopulations that make them quitevulnerable to habitat loss. Effects ofhabitat alteration can be far-reaching.

Management of sea turtles hasemphasized the acquisition andprotection of nesting habitat. Otherconcerns include ocean pollution,fishing and shrimping nets, beachdevelopment, and enforcement ofinternational regulations. Theidentification of migration routes andother life history information also willbenefit future management strategies.

Degradation of aquatic habitat isthe primary management concernfor freshwater turtles. Conservationactions are directed at monitoring theextent of thermal pollution, dredging,channelization, and incidental takesby commercial fishing. Protectionof nesting beaches and adjacentnest areas, and the prevention ofdeliberate killing are also importantmanagement priorities.

In addition to intentional killing,which affects snakes as well as turtles,several reptiles suffer direct losses dueto exploitation. Unregulated harvestaffects a number of the listed turtleand tortoise populations, as well asthe majority of sea turtles. Collectionfor the pet trade is another seriousmanagement problem. Some species,particularly the genera Clemmys andGraptemys, require strict regulation dueto rising demands in domestic andforeign pet markets. Commercialcollectors also threaten the spottedturtle and box turtle.

The invasion of introduced exoticscan also be detrimental to nativereptiles. Fire ants, in particular,have been implicated in the reductionof terrestrial egg-laying reptiles(Mount 1986).

Management can enhance reptilehabitat in many ways. One way isthrough the retention of microhabitatfeatures that provide refuges. Forexample, the disruption of under-ground root systems in managed pineplantations may displace species suchas the eastern diamondback rattlesnake.The importance of leaving terrestrialbuffer zones around forest wetlands iswell documented (Burke and Gibbons1995, Semlitsch and Bodie 1998). Theretention of habitat elements such asleaf litter, snags, coarse woody debris,and fallen logs benefits the habitat ofmany reptiles (refer to the individualsubtaxa sections mentioned earlierand Species accounts that follow).

Delayed sexual maturity andindividual longevity contribute to thevulnerability of reptiles and inhibit therecovery of several threatened species.Several reptiles have existed virtuallyunchanged for centuries. Unfortunately,some of the same traits that allowedthem to survive the ages predisposethem to endangerment. Conservationactions should be directed towardsareas of high species diversity, specieswith limited distributions, and locationssuch as shallow wetlands and coastalzones where reptiles are at risk.Insufficient knowledge of the distri-bution and ecology of native reptilesis a major shortcoming in any regionaleffort to detect change and avoid lossin these taxa.

Species accounts—The followingare the species accounts for selectedreptiles that are of concern in theSouth. Several are federally listed asthreatened or endangered. Others areclassified as imperiled or vulnerable byNatural Heritage agencies. Managementrecommendations follow Wilson(1995) unless otherwise cited.

Species accounts: Alabama redbellyturtle—This endangered species isrestricted to Mobile Bay in southernAlabama. It has declined due tohabitat modification and becauseit was trapped and netted for food(Dobie and Bagley 1988). Habitatdisturbance has altered the turtles’nesting and feeding habitat.

Primary habitat areas are the upper,freshwater portions of Mobile Bay,where there are abundant suppliesof submerged plants and algae,which are preferred foods.

Conservation actions emphasizeprotection of the primary nestingsite on Gravine Island, restrictionof herbicide use, and limitationof dredging activity on the lowerTensaw River.

Species accounts: alligatorsnapping turtle—This species isclassified as vulnerable by NaturalHeritage and is listed in the Statesof Alabama, Georgia, and Texas(NatureServe 2000). The turtlehas declined due to habitat lossand commercial exploitation forfood and the pet trade.

The species is typically found indeep rivers and canals, but may alsooccur in lakes, swamps, and smallstreams. Although it nests on land,the turtle is primarily aquatic andfeeds on fish, mollusks, and crayfish.

Conservation measures includeregulation of collection and theprotection of suitable habitatwith adequate prey populations(Wilson 1995).

Species accounts: Americanalligator—This species is federallylisted as threatened due to similarity ofappearance to the American crocodile.The alligator ranges from coastal NorthCarolina to extreme southern Florida,west to east Texas, and north to centralArkansas. Current threats include theconversion of habitat for recreationaluse and urban development.

Alligators prefer large, shallow lakes,fresh or brackish marshes, and savannasthat border aquatic habitat. Alligatorsare strictly carnivorous and will eat anyanimal they can subdue and swallow.

Conservation actions for the Americanalligator focus on habitat protectionand control of human disturbance.

Species accounts: Americancrocodile—The crocodile is federallylisted as endangered. The speciesoccurs in south Florida and the FloridaKeys. It inhabits the Caribbean, CentralAmerica, and South America. Habitatloss is the primary reason for thisspecies’ imperilment in the South(Moler 1992a).

The crocodile is found in brackishor salt water in coastal canals,


mangrove thickets, or tidal creeks.The crocodile is carnivorous.Conservation actions center onprotection of the remaining habitatin southern Florida (Wilson 1995).

Species accounts: Atlantic saltmarsh snake—This threatened speciesis restricted to a small coastal strip inFlorida. It is imperiled by wetlandhabitat alteration stemming fromdrainage and impoundment.

This snake preys on fish andis typically found in salt marshes,tidal creeks, and mangrove swamps.Conservation action for this speciesis concerned with protection ofthe remaining unaltered habitat(Conant and Collins 1991).

Species accounts: bog turtle—This threatened species occurs insouthwestern Virginia, easternTennessee, northern Georgia, andthe Carolinas. The bog turtle is injeopardy due to collection for the pettrade and habitat loss. The drainageof grassy and marshy wetlands hasresulted in the destruction of therequired habitat for this species.

The bog turtle feeds on a variety ofanimals including tadpoles, frogs,various invertebrates, and baby rodents.The species does not tolerate closed-canopy forests. Management actionsfocus on the maintenance of early seral(grassy) habitat and halting the illegalpet trade (Ernst and others 1994).Drainage of wetlands is detrimentalto this species (Wilson 1995).

Species accounts: Florida scrublizard—This species is classifiedas vulnerable by Natural Heritage(NatureServe 2000). Disjunct popula-tions occur along the east coast ofFlorida, in central Florida, and inisolated areas on the west coast ofFlorida. The species is threatened byconversion of habitat to other uses.

The lizard prefers open sandy edgesin xeric sand pine scrub and longleafpine habitat (Conant and Collins1991). It feeds on ants, beetles, spiders,and other small arthropods.

Conservation strategies focus on themanagement of sand pine scrub andlongleaf pine-turkey oak habitats toretain the open character that the lizardrequires. The Ocala National Forestmanages large areas of this habitat(Wilson 1995). Habitat maintenanceoften requires prescribed burning.

Species accounts: gopher tortoise—This threatened species occurs inFlorida, Georgia, South Carolina,Mississippi, Alabama, and Louisiana.Habitat loss and the pet trade arethe primary factors contributing tothe decline of the tortoise (U.S. Fishand Wildlife Service 1990c).

Well-drained sandy soils supportingpine and scrub oaks in the sandhillsare preferred habitat. The tortoise feedson grasses, forbs, and other vegetation(Ernst and others 1994).

Habitat management for the gophertortoise includes selective harvestand prescribed burning to maintainthe open, grassy nature of sand ridges.Ground disturbance such as heavysite preparation and root raking canbe detrimental to young tortoises(Wilson 1995).

Species accounts: indigo snake—The population of this threatenedspecies has declined rapidly in recentyears. Primary threats appear to behabitat loss and exploitation for thepet trade (Speake and others 1982).The indigo snake is currently foundin southeastern Georgia and Florida.

The species coexists with gophertortoises throughout much of its rangeand frequently uses tortoise burrows.Preferred habitat is pine-scrub oakwoodlands and palmetto-covered hillswith well-drained sandy soils. Indigosnakes may also be found in mesichabitats bordering swamps, streams,or canals. The snake feeds on frogs,toads, birds, small mammals, andother reptiles.

Conservation actions necessaryto protect indigo snake populationsinclude the retention of existinghabitat, maintenance of pine-scruboak woodlands in a subclimaxcondition, and protection of gophertortoise burrows (Moler 1992c).

Species accounts: Louisiana pinesnake—This species is endemic toeastern Texas and western Louisiana,primarily in areas currently or oncedominated by longleaf pine. Thespecies is associated with fire-main-tained pine forests on well-drainedsandy soils with well-developedherbaceous vegetation (Rudolph andBurgdorf 1997). Pocket gophers are theprimary prey of Louisiana pine snakes,and pocket gopher burrows are used forescape from predators, avoidance ofhigh temperatures, and hibernation.

The species has apparently declinedin recent decades, and existingpopulations are thought to be smalland isolated (Reichling 1995, Rudolphand Burgdorf 1997).

Loss of habitat due to conversionto intensive silviculture and changesin the fire regime are the primary causesof population decline. Fire suppressionand inadequate prescribed fire haveresulted in widespread successionalchanges in pine forests throughout therange of Louisiana pine snakes, leadingto loss of herbaceous vegetation andpocket gophers. Habitat loss anddegradation has been more extensiveon private than on public land. Roadsand associated vehicle traffic are verylikely impacting populations in muchof the remaining habitat.

Conservation action centers onthe management of fire-maintainedpine habitat on a scale sufficient tosupport viable populations of thisspecies. Prescribed burning sufficientto maintain abundant herbaceousvegetation and support of pocketgopher populations are required.

Species accounts: mimic glasslizard—This species is classifiedas vulnerable by Natural Heritage(NatureServe 2000). The lizardoccurs on the Atlantic Coastal Plainfrom North Carolina to Florida andwestward as far as the Pearl River inMississippi. The species is imperileddue to excessive development andhabitat modification in its range.

The lizard inhabits open-canopiedpine forests with thick forest litter. Itfeeds on a variety of invertebrate preyas well as small lizards and snakes(Palmer and Braswell 1995).

Conservation actions to benefitthis species include maintaining anopen canopy through burning andthinning (Wilson 1995).

Species accounts: bluetail moleskink—This threatened subspeciesoccurs in Florida. Residentialdevelopment and agriculturalconversion have altered its habitat.The mole skink prefers open, sandyedges in sand pine scrub and sandhillhabitats. The species consumes avariety of invertebrate prey, includingants, beetles, and spiders.

Conservation actions for this speciesfocus on protection of essential habitatareas from conversion to other uses(Conant and Collins 1991).


TERRESTRIALOther mammals

41 (17%)

Whales and dolphins30 (12%)

Rodents86 (35%)

Bats37 (17%)

Carnivores52 (24%)

Species accounts: rim rockcrowned snake—This species isclassified as critically imperiled byNatural Heritage and occurs solelyin Florida (NatureServe 2000).Development and the resultanthabitat loss threaten the snake.

This snake is found in flatwoods,tropical hardwood hammocks, andpastures and on fossil coral reefs(Porras and Wilson 1979). Thissnake consumes insects and othersmall arthropods.

Because of the intense developmentoccurring in the habitat of thisspecies, conservation action centerson the protection of suitable habitat(Wilson 1995).

Species accounts: ringed mapturtle—This threatened turtle inhabitsthe Pearl River drainage of southernMississippi and southeastern Louisiana.Primary threats are illegal collection forthe pet trade and habitat degradation(U.S. Fish and Wildlife Service 1988).

This turtle leaves the river to baskand to lay eggs. Preferred habitatincludes river stretches with moderatecurrent, sandbars, and debris forbasking sites. The diet is comprisedof insects and mollusks. Because thisturtle is restricted to the Pearl River,modifications of natural conditionsthere could prove detrimental.

Species accounts: sand skink—This species is classified as imperiledby Natural Heritage (NatureServe2000). It is restricted to well-drainedsandy soils in the interior centralFlorida highlands. Much of this habitathas been converted to citrus grovesand residential areas.

The sand skink lives in loose,dry sandy areas with sparse grasscover. It subsists on a diet of ants,spiders, termites, beetle larvae, andother invertebrates.

Protection of the remaining habitatand acquisition of additional areasare the primary conservation actionsrequired to preserve this species(Christman 1992). In addition, theuse of prescribed fire is importantfor maintaining the open nature ofsand skink habitat.

Species accounts: short-tailedsnake—This species is classified asvulnerable by Natural Heritage andis endemic to Florida (NatureServe2000). Habitat destruction is the

primary threat, particularly in centralFlorida, where land is in demandfor agricultural, residential, and otheruses (Wilson 1995).

The primary habitats of this snakeare longleaf pine-turkey oak and sandpine scrub communities with loosesandy soils. It feeds on small snakesand lizards.

Management actions center on theprotection of remaining occupiedhabitat from development and theretention of prey populations(Campbell and Moler 1992).

Species accounts: southern hognosesnake—This species is classifiedas imperiled by Natural Heritage(NatureServe 2000). The snake occursprimarily on the Coastal Plain fromNorth Carolina westward into southernMississippi. There is one disjunctpopulation in central Alabama.Development of preferred habitatis the primary cause for imperilment,but fire ants may also be impactingpopulations (Wilson 1995).

This snake is found in sandhills,pine-scrub oak woodlands, pine andwiregrass flatwoods, and other openxeric communities with loose, sandysoils (Martof and others 1980). It feedsprimarily on toads, frogs, and lizards.

Conservation actions for this speciesinclude the protection and restorationof remaining habitat, restriction ofadditional development, and fireant control.

MammalsTerrestrial, marine, and freshwater

habitats in the South are home to 246mammalian species (NatureServe2000). The number of mammals rangesfrom 176 species in Texas to 62 speciesin Mississippi. There are 102 speciesin Georgia, 101 in South Carolina,96 in Oklahoma, and 95 in Florida.The total includes rodents, carnivores,bats, whales, dolphins, and othermammals (fig. 5.6).

This vertebrate group comprises11 major orders and 26 families(Echternacht and Harris 1993). All butfive families have one or more sensitivespecies (Laerm and others 2000).These families include Didelphidae(opossum), Dasypodidae (armadillo),Castoridae (beaver), Myocastoridae(nutria), and Suidae (wild boar). Theorder Rodentia dominates the region’smammalian fauna in the number of

different species. This order includeschipmunks, squirrels, pocket gophers,mice, rats, voles, muskrats, nutria, andbeavers. Examples of carnivores includethe Florida panther, red fox, bobcat,river otter, and mink. The category of“other mammals” in figure 5.6 includesthe Florida manatee, white-tailed deer,eastern cottontail rabbit, opossum,armadillo, shrews, moles, and severalother species.

Five mammal species are known orpresumed to be extinct or extirpatedfrom the region. These are the jaguar,ocelot, gray wolf, elk, and bison(Echternacht and Harris 1993). Beaverswere once extirpated in the Southbut were reestablished over the pasttwo decades.

Endemic species represent a relativelysmall percentage of the mammals inthe region. Eight rodent species areendemic to the Coastal Plain: thesoutheastern pocket gopher, colonialpocket gopher, Sherman’s pocketgopher, Cumberland Island pocketgopher, oldfield mouse, Florida mouse,Perdido Key beach mouse, and round-tailed muskrat (White and others1998). The region also has eight speciesof introduced mammals, includingthe coyote, wild boar, and nutria.

Thirty-three species of mammalsare listed as threatened or endangered(table 5.12). These include the Keydeer, red wolf, Louisiana black bear,Indiana bat, gray myotis, Virginianorthern flying squirrel, andsoutheastern beach mouse. Ten ofthe listed rodent species inhabit theCoastal Plain of Florida or Alabama.

In addition, 12 species are classifiedas imperiled or vulnerable under theNatural Heritage system (chapter 1).

Figure 5.6—Species richness by majorsubgroups of mammalian taxa occurringwithin the South (NatureServe 2000).


Table 5.12—Mammal species within the South that are listed as threatened or endangered


BatsCorynorhirus townsendii ingens Ozark big-eared bat (E) AR, OKCorynorhinus townsendii

virginianus Virginia big-eared bat (E) NC, VAMyotis grisescens Gray bat (E) AL, AR, FL, GA, KY, OK,

TN, VAMyotis sodalis Indiana bat (E) AL, AR, GA, KY, MS, NC,

OK, SC, TN, VA

RodentsGlaucomys sabrinus coloratus Carolina northern flying squirrel (E) NC, TNGlaucomys sabrinus fuscus Virginia northern flying squirrel (E) KY, NC, VAMicrotus pennsylvanicus

dukecampbelli Florida salt marsh vole (E) FLNeotoma floridana smalli Key Largo woodrat (E) FLOryzomys palustris natator Rice rat (E) FLPeromyscus gossypinus

allapaticola Key Largo cotton mouse (E) FLPeromyscus polionotus allophrys Chocawhatcher beach mouse (E) FLPeromyscus polionotus ammobates Alabama beach mouse (E) ALPeromyscus polionotus niveiventris Southeastern beach mouse (T) FLPeromyscus polionotus peninsularis St. Andrew beach mouse (E) FLPeromyscus polionotus phasma Anastasia Island beach mouse (E) FLPeromyscus polionotus trissyllepsis Perdido key beach mouse (E) AL, FLSciurus niger cinereus Delmarva Peninsula fox squirrel (E) VA

CarnivoresCanus rufus Red wolf (E) NC, TN, FLHerpailurus yogouaroundi

cacomitli Gulf Coast jaguarundi (E) TXLeopardus pardalis Ocelet (E) TXPanthera onca Jaguar; Otorongo (E) TXPuma concolor Puma (T) FLPuma concolor coryi Florida panther (E) FLPuma concolor cougar Eastern puma (E) KY, NC, SC, TN, VAUrsus americanus luteolus Louisiana black bear (T) LA, MS, TX

Whales and dolphinsBalaenoptera physalus Fin whale (E) AL, FL, GA, LA, MS, NC,

SC, TX, VAEubalaena glacialis Black right whale (E) FL, GA, NC, SC, VAMegaptera novaeangliae Humpback whale (E) AL, FL, GA, LA, MS, NC,

SC, TX, VAPhyseter catodon Sperm whale (E) NC

Other mammalsMonachus tropicalis Caribbean monk seal (E) FLOdocoileus virginianus clavium Key deer (E) FLTrichecchus manatus Manatee (E) AL, FL, GA, LA, MS, NC,

SC, TXSylviagus palustris hefneri Lower Keys marsh rabbit (E) FL

T = threatened; E = endangered.Source: U.S. Department of the Interior (2000).


TERRESTRIAL

Table 5.13—Bat species occurring within the South

Scientific name Common name Status

Artibeus jamaicensis Jamaican fruit-eating bat Limited numbersCorynorhinus townsendii Townsend’s big-eared bat Endangereda

Corynorhinus rafinesquii Rafinesque’s big-eared bat Special concernEptesicus fuscus Big brown bat CommonEumops glaucinus Wagner’s mastiff bat RareLasionycteris noctivagans Silver-haired bat Relatively uncommonLasiurus borealis Eastern red bat CommonLasiurus cinereus Hoary bat Relatively commonLasiurus intermedius Northern yellow bat Relatively commonLasiurus seminolus Seminole bat CommonMolossus molossus Pallas mastiff bat Limited numbersMyotis austroriparius Southeastern bat Special concernMyotis grisescens Gray bat EndangeredMyotis leibii Eastern small-footed bat Special concernMyotis lucifugus Little brown bat Scarce or locally commonMyotis septentrionalis Northern long-eared bat CommonMyotis sodalis Indiana bat EndangeredNycticeius humeralis Evening bat CommonPipistrelle subflavus Eastern pipistrelle CommonTadarida brasiliensis Brazilian free-tailed bat Locally common/abundantb

a Two subspecies: Ozark big-eared bat and the Virginia big-eared bat.b Two subspecies: LeConte’s free-tailed bat and the Mexican free-tailed bat.Source: Adapted from Harvey and Saugey (2001).

These include the Rafinesque’s big-eared bat, gray-footed chipmunk,round-tailed muskrat, Alleghenywoodrat, and swift fox. These speciesare in jeopardy due to habitat loss,land use change, human disturbance,and coastal development.

The white-tailed deer is the mostwidespread browsing species repre-sented in the region today. Elk haverecently been reintroduced intoselected locations. The absence oflarge carnivores (wolves, jaguar)reflects history since Europeansettlement (chapter 1). The blackbear is the largest carnivore now inthe South. Four wild canids occur inthe region. The coyote has expandedits range, while the red wolf is criticallyimperiled due to habitat loss andhybridization with other canids.Red and gray foxes remain relativelycommon. The Florida panther is injeopardy, while the bobcat remainswidespread throughout the region.

The absence of large predatorshas encouraged the proliferation ofraccoons, opossums, and skunks. Thesespecies demonstrate broad ecologicaltolerance, inhabiting virtually everytype of habitat available. They consume

a variety of foods: frogs, turtles, snakes,mice, berries, and other vegetation.These mammals are rapidly becomingurban wildlife in many communitiesof the South.

Rodents are a diverse group thatpersists in abundance in many areas.They tend to have high birth rates thatpermit the maintenance of stablepopulations despite predation pressureand control measures. The rodentspecies that are most at risk in theSouth have narrow distributions. Inbeach habitats, feral cats represent asignificant threat. Pesticide residuesaffect shrews and other insectivores.The fox squirrel that inhabits longleafpine savannas is threatened by firesuppression and land use conversion(White and others 1998).

The absence of mountain barriersand other opportunities for isolationand speciation contribute to the lackof species richness among squirrelsand burrowing mammals (Echternachtand Harris 1993). The eight species ofsciurid rodents in the region includethe 13-lined ground squirrel, graysquirrel, fox squirrel, and two flyingsquirrels. The region’s 10 burrowingrodents include the hairy-tailed mole,

eastern mole, and star-nosed mole;woodchuck; eastern chipmunk; and5 species of pocket gophers. Soil typeis the primary factor determiningthe ranges of pocket gophers.

The following sections discuss thehabitat needs for two of the highestprofile groups of mammals: bats andcarnivores. Additional species arealso profiled in the Species accountsection that concludes the segmenton mammals.

Bats—The 20 species of bats in theSouth are key components of forestedecosystems. Four bats are listed asendangered: the gray bat, Indiana bat,and Ozark and Virginia big-eared bats(table 5.13). The southeastern bat, theeastern small-footed bat, Rafinesque’sbig-eared bat, and Wagner’s mastiffbat are of special concern.

Forest bats depend on forests forshelter, roosting sites, and foragingareas. Bats are in two major classes:cave bats and tree bats. Cave batsinhabit caves during all or part of theyear, while noncave species seldomenter caves. Some of their ranges arelimited to relatively small geographicareas. Insectivorous bats have tiny


eyes and are capable of sight, but mostspecies locate prey by echolocation.

Bats hibernate in a variety of locationsincluding leaf litter, woody debris,caves, hollow trees, and rock crevices.Many species hibernate underexfoliating bark and in tree cavities,mines, and buildings. Roosting sitesrange from solitary sites to cavescontaining thousands of individuals.Sites selected for roosting andhibernation meet precise environmentalconditions, such as stable temperaturesand high relative humidity. Disturbanceoften results in the abandonment ofthe site.

Bats have evolved to fill a varietyof food niches. These mammals beginforaging at dusk. The diet consistsof insects and other arthropods andvaries by species.

Widespread pesticide use causedsignificant declines in bat populationsduring the past several decades (Harveyand others 1999). This threat hasdiminished somewhat with pesticideuse regulations. The current threat tobats stems from habitat destructionand cave disturbance. Few caves meetthe narrow temperature and humidityrequirements for hibernation. The largenumbers of bats occupying specificcaves make these species vulnerableto disturbance of an individual cave.

Various locations are used asmaternity roost sites. Snags are usedby Indiana, northern, and evening bats,while hollow trees are important forRafinesque’s and southeastern bats. Aparticular threat is human disturbanceto hibernation and maternity colonies.Hibernating bats wake when disturbedand expend critical winter stores offat. Summer maternity colonies havelow tolerance of disturbance; disturbedparents will often abandon theiroffspring. Bats produce an averageof one offspring per year, but somespecies give birth to three or fourbabies at a time. The low rate ofreproduction results in populationsthat can be quickly destroyed with littleopportunity for recovery. Other adverseimpacts include habitat destruction;direct killing; vandalism; and predationby raptors, raccoons, skunks, andsnakes (Tuttle 1995).

A number of forest managementactions can enhance bat habitat. Seed-tree and shelterwood harvests open upforest canopies, creating foraging

opportunities by reducing branchobstructions (Krusic and others 1996).Retention of cavity trees and snags,creation of large snags, and designationof streamside zones also are beneficial(Harvey and Saugey 2001, Kulhavy andConner 1986). The creation of pondscan also enhance habitat by providingwater, breeding sites, and a source ofinsect prey (Wilhide and others 1998).

Even-aged poletimber stands oftenare unsuitable for bole and cavity usersand do not provide the cavities andbark characteristics preferred bybats (Pierson 1998). Clearcuttingeliminates roosting opportunitiesuntil replacement trees of suitablesize become available (Harvey andSaugey 2001). However, the resultingavailability of herbaceous growthresults in increased insect populations(Barclay and Brigham 1998). Standrotations long enough to allow forcavity development are importantfor species that require cavities.

Prescribed burning can enhanceinvertebrate biomass by reducingmidstory trees and shrubs, allowingthe regeneration of herbaceous plants.The resulting canopy gaps provideadditional foraging opportunities.However, fire may jeopardize batshibernating on the ground duringwinter when they are torpid and slowto arouse (Harvey and Saugey 2001).The impact of dormant-season burningon species that roost in ground litteris unclear. Snags used by bats maybe felled by fire if their bases burnthrough, resulting in the loss of cavitiesor roosting sites under exfoliating bark.

Finally, recreational caving should beminimized to prevent disturbance tomaternity and hibernating colonies.Properly designed gates on caveentrances afford the best protection.Other protective measures includelimiting the use of pesticides andpreventing destruction of habitat.

Carnivores—Carnivores are a viablecomponent of the southern landscape,whose management has changed sig-nificantly over the last several decades.The perception that carnivores mustbe eliminated is no longer widelyheld. These mammals contribute toecosystem stability by controllingrodent populations.

There are few reliable densityestimates for furbearers because theyare secretive and difficult to census.

Most are territorial. Population densityis relatively low, reflecting their positionat the top of the food chain. Twocarnivores (the bobcat and river otter)are protected under the Convention forthe International Trade of EndangeredSpecies of Fauna and Flora (CITES)and are monitored closely by Statesthat allow harvest of these species(Leopold and Chamberlain 2001).

The diet of carnivores is primarilycomposed of other animals. Bobcats,river otters, weasels, and minkcharacteristically have diets in whichanimal material exceeds 95 percent.The amounts of fruits, berries, andseeds vary with seasonal availability.For example, gray and red fox foodschange from animal foods in the falland winter to invertebrates and fruitsduring spring and summer.

Each species is associated withspecific habitats that provide requiredfood, water, and cover. Often, areas thatare diverse in vegetative composition,structure, and seral stage are inhabitedby a diversity of these mammals. Asubstantial number of carnivoresdepend on forested ecosystems toprovide one or more habitat require-ments. Mosaics of cover types and theecotones between successional stagesenhance prey and other food diversity.The structural components importantto many mammals include maturetrees, standing dead trees, woodydebris, and patchy understories.Structural diversity and decayingtrees provide suitable cover andforaging habitat.

Habitat quality determines thestability of these populations, whilehabitat loss is the primary threat tothese species. Habitat modificationinfluences species distribution andabundance. Forest clearing, grasslandconversion, irrigation, and wetlanddrainage have improved habitat forsome species and damaged habitatfor others. The expanded range ofthe coyote throughout the Southresulted from urbanization and theremoval of large predators such asred wolves and Florida panthers.

Species with restrictive habitatrequirements are vulnerable to losses ofhabitat. The swift fox depends on nativeshortgrass prairie communities; itsrange has become restricted due to theconversion of prairies into cultivatedfields. Mammals associated withwetland habitats are not very resilient


TERRESTRIAL

Table 5.14—Important habitat components and associated management guidelines for selected mammalsin the South (continued)

Species Key components of habitat Management guidelines

Raccoon Wetlands, riparian habitats, suitable den sitesand winter food.

Red fox High degree of habitat edge; interspersion ofmosaic of woodland, shrubland, cropland,and grassland habitat.

Red squirrel Dense or clumped stands of mature forest;multi-storied stands; suitable nest sites;sufficient shade for cone storage.

Ringtail cat Rocky, brushy areas, talus slopes or woodedhabitats in close proximity to water.

River otter Water quality; permanent surface water,vegetative cover adjacent to wetlands;structural cover to provide foraging andden sites.

Spotted/striped Ecotones between forest/shrubland andskunks agricultural lands; riparian areas in arid

regions.

Swift fox Mid to shortgrass prairie habitats suitableto support an adequate prey base.

Source: Adapted from Allen (1988).

Maintain woodlots in agricultural areas toenhance vegetation diversity; maintainfencerows for travel corridors; encouragesoftmast production.

Maintain large deciduous trees with cavities;preserve densely branched trees; provideclumped stands near mature conifers withinterlocking crowns.

Maintain riparian vegetation in associationwith draws and ridgelines as travelcorridors.

Maintain vegetative cover adjacent to wet-lands; increase pool to riffle ratios; ensurewater permanence; encourage beaver estab-lishment.

Maintain woodlots in agricultural areas toenhance vegetation diversity; maintainfencerows for travel corridors; encouragesoftmast production.

Establish vegetative communities to supportprey base; maintain interspersion of grasslandcommunities with agricultural lands.

Preserve wetlands and riparian areas; maintainsnags or diseased trees for den sites; encour-age mast species; maintain fencerows.

to habitat modification. For example,river channelization reduces habitatsuitability for river otters (Allen 1988).

Large mammals such as the redwolf, Florida panther, and black bearhave extensive home ranges. Themaintenance of a mosaic of vegetationtypes and multiple seral stages supportsprey populations and the food-producing plants that comprise thediet of these species. In contrast, themajority of carnivores depend on muchsmaller geographic areas. These speciesrely on a diversity of cover types inrelatively close proximity to provideseasonal cover and food. Red foxes,gray foxes, and weasels are associatedwith early to mid-successionalvegetation and the ecotones betweenthese communities. Management thatmaintains fencerows, shelterbelts, and

riparian vegetation will benefit thesespecies and enhance their distribution.

The elimination of woody debrisinfluences small mammal populationsand makes them easier prey forassociated predators. Timber harvestand prescribed burning changevegetation composition and enhancesunderstory growth. However, timberremoval may harm other mammals thatrequire mature forest. In some cases,the protection of critical habitat maybe the preferred management strategy.

Conservation of wetland carnivorescenters on prevention of wetlanddegradation. Vegetative structure,surrounding land use, water quality,and cover diversity influence habitatquality for these mammals. Forexample, the manipulation of water

levels and the planting of desiredvegetation can enhance habitat. Themaintenance of water availabilityand prey species also improves habitatpotential. Debris and structuraldiversity along shorelines enhance preyavailability for river otters. The removalof aquatic shoreline vegetation reducesavailability of prey for mink.

Important habitat features forcarnivores as well as other mammalsoccurring in the South are summarizedin table 5.14. Detailed information forselected species is presented in thefollowing section.

Species accounts: beaver—Thisspecies was extirpated from most ofits southern range by the 1950s dueto extensive trapping that began inthe 18th century. Restocking programs


in Louisiana, Mississippi, Alabama,Florida, Georgia, Oklahoma, Virginia,Arkansas, and North and SouthCarolina have led to viable populationsacross most of the South (Jones andLeopold 2001).

Beavers use freshwater habitats suchas ponds, small lakes, and streams.Slow-moving streams and creeks withproximity to trees and shrubs thatprovide a food source are important.Beaver damming can flood forests,causing substantial economic impactfrom prolonged flooding. However,beavers create a complex successionalmosaic of aquatic and terrestrialhabitats that enrich landscape diversity.The creation of wetlands positivelyinfluences ground water, water quality,structural diversity, and erosionresistance. Beaver impoundmentscreate favorable conditions for fish,birds, and amphibians. Beaver pondson intermittent streams provide aquatichabitat conducive to the river otter.

River channelization significantlyaffects habitat quality by reducingamounts of riparian vegetation,macroinvertebrates, and fish biomass.The modification of river flow rates alsoreduces the number of islandsoccurring in the channel, impactingpotential den habitat.

Species accounts: black bear—Black bears historically ranged overmost of the South. Habitat loss, frag-mentation, and unrestricted harvesthave significantly changed theirdistribution and abundance.

Their current distribution is restrictedto relatively undisturbed forests in theAppalachian Mountains and the InteriorHighlands of Arkansas and in scatteredcoastal areas from Virginia to Louisiana(Vaughn and Pelton 1995). Populationsappear to be secure and increasing inparts of Virginia, Tennessee, NorthCarolina, northern Georgia, andnorthern South Carolina, where theysupport regulated hunting seasons.In Tennessee, the species is knownonly from the mountains in the easternpart of the State (Chapman and Laerm,in press). In Kentucky, the black bearis designated as a species of specialconcern. Texas biologists indicatethere is no resident breeding popu-lation there.

Two subspecies are of special concern.The Louisiana black bear is designatedas threatened on the Federal species

list and as endangered by the Statesof Mississippi, Louisiana, and Texas.The Florida subspecies is listed asthreatened by the Florida Game andFresh Water Fish Commission. Untilrecently, this subspecies was consideredfor protection under the FederalEndangered Species Act. Both sub-species populations are restricted toislands of public land and inaccessibleareas of bottomland forest.

Black bears inhabit diverse foresthabitats and are often found in oak-hickory and mixed mesophytic forests.Forested areas of 150 to 300 squaremiles with limited human intrusion areneeded to sustain viable populations.In coastal areas, the species occupiespocosins, hardwood bottomlands,Carolina bays, mixed hardwoodhammocks, cypress swamps, pineflatwoods, and sand pine scrub.Black bears need dense understorycover, such as laurel thickets andgreenbriar, to provide refuge coverin the Coastal Plain.

Adequate denning cover is a necessarycomponent of black bear habitat inthe South. Such cover includes cavitiesin large trees, logs, stumps, rockoutcroppings, and impenetrablethickets. Females and cubs are verysusceptible to disturbance. Black bearsneed secure corridors to make seasonalmovements for food, for dispersal ofyounger animals, and for movementby males during the breeding season(Pelton 2001).

The diet of black bears is primarilyhard and soft mast, including berries,nuts, acorns, wild cherries, and grapes,as well as invertebrates. In some areas,bears feed on agricultural crops such ascorn, wheat, or soybeans. Black bearswill occasionally eat opossums,armadillos, feral pigs, raccoons, andyoung white-tailed deer.

The seasonal variations in availabilityof soft and hard mast influence shiftsin home range to locate these foods.State biologists indicate that duringperiods of drought and food scarcity,bears further disperse and becomevictims of vehicular accidents,nuisance control, and illegal hunting.

Bear populations in the SouthernAppalachians have been monitoredsince the 1960s. Although bear popu-lations have increased during thisperiod, the illegal trade in bear gallbladders has raised concerns about

the effect of poaching. Because bearshave low reproductive rates, theirpopulations recover slowly from losses.

Habitat degradation continues tothreaten black bears in the South.Forest fragmentation and the conver-sion of forests to agriculture, urbandevelopment, and pine monoculturesrestricts available habitat (Pelton 2001).The fragmented nature of black bearpopulations in the Coastal Plain maycontribute to a loss of genetic diversity.As the human population in the Southcontinues to expand into bear habitat,increased incidents of road kills arebeing reported in North Carolina,Tennessee, and Florida. As peoplesettle into established bear ranges,increased human-bear interactionsare inevitable. Poaching and increasedaccess capabilities can resultin overexploitation.

Components of black bear manage-ment include hunting access, habitat,protection, nuisance control, education,and research (Pelton 2001). Accesscan be restricted through road gating,designation of no-hunting zones, andprovision of escape cover. Habitatmanagement includes oak enhance-ment, protection of old growth (forden trees), and management of forestopenings for soft mast production. Theestablishment of black bear sanctuariesand viable corridors on public land hasprotected bears in the region (Vaughnand Pelton 1995). Texas has proposedthe establishment of bear “recoveryzones” through a partnership amongFederal and State agencies, forestindustry, and other owners of largeparcels of timberland. Stringent lawenforcement also is required to reduceillegal hunting. Finally, State biologistssuggest that education of the generalpublic is critical to increase awarenessand acceptance of regulations such asthose that discourage feeding of bears.

Species accounts: bobcat—Bobcats are found throughout theSouth with the exception of north-central Kentucky, coastal Louisiana,and eastern Virginia (Leopold andChamberlain 2001). Populationdensity varies according to habitattype and prey density.

Bobcats use several habitats,preferring areas with dense under-story vegetation that supports preypopulations. A mixture of matureand early successional forest habitats isbest. Other habitats include agricultural


TERRESTRIAL

fields and pastures. Home ranges ofbobcats throughout the SoutheasternUnited States range from less than740 acres to 17,830 acres. Home rangesmay reflect road avoidance. Importantprey species include rabbits and variousrodents, opossum, game birds, andsnakes (Chapman and Laerm, in press).

There are no major threats to bob-cats in the South due to their widedistribution and ecological tolerance.Potential risks include overharvestby trapping, forest fragmentation,and road construction.

Species accounts: Carolinanorthern flying squirrel and Virginianorthern flying squirrel—Thesetwo endangered subspecies inhabithigh-elevation sites in the SouthernAppalachians. The Carolina squirreloccurs in isolated locations in NorthCarolina and Tennessee, while theVirginia subspecies is in Virginia andWest Virginia. The disjunct distributionof these subspecies in the SouthernAppalachians suggests they are relictsthat have become isolated in smallpatches of suitable habitat by changingclimatic and vegetation conditions sincethe last Ice Age (U.S. Fish and WildlifeService 1990a).

Flying squirrels are associatedwith high-elevation boreal habitats,especially spruce-fir and northernhardwood forests (Fridell and Litvaitis1991). They occur in conifer-hardwoodecotones consisting of red spruce andfir associated with mature beech, yellowbirch, maple, and several other species.Widely spaced, mature trees and snagsprovide cavities for nesting. Understorycomponents do not appear to beimportant components of northernflying squirrel habitat (U.S. Fishand Wildlife Service 1990a).

Their diet consists of lichens, fungi,seeds, fruit, staminate cones, andinsects. Periodic dependence on certainspecies of fungi may be a factor inrestricting the species to high-elevation,mesic habitats (U.S. Fish and WildlifeService 1990a).

The limited range of this speciesmakes it vulnerable to natural andhuman-related impacts. Isolatedpopulations suffer from insufficientgene pools. Other concerns includehabitat destruction, insect pests suchas the balsam woolly adelgid and thegypsy moth, recreational use, acid rain(which contaminates their mycorrhizal

food source), and heavy metals (lead,copper, nickel, zinc, and manganese)in forest litter and soil (U.S. Fish andWildlife Service 1990a).

Conservation strategies includedetermination of species distributions,protection of occupied sites fromhuman-related disturbance, andimplementation of habitat manage-ment guidelines on national forestsand parks.

Species accounts: coyote—Thedistribution of coyotes has extendedinto the South during the past fewdecades in response to the eliminationof gray and red wolves from theirformer ranges. Prior to 1970, redwolves were common throughoutthe South, but trapping and poisoningeliminated free-ranging populations.Gray wolves also once inhabitedKentucky, Tennessee, Virginia, andNorth Carolina. Removal of thesetwo species contributed to coyoteexpansion. Leopold and Chamberlain(2001) indicate that coyote popula-tions have expanded throughoutthe South, with the exception ofsouthern peninsular Florida. Thecurrent population density ofcoyotes is unknown.

Coyotes occupy a broad range ofhabitats and occur in grassland, forest,agricultural fields, and urban areas.In the South, this species has beenobserved in open fields, brushlands,thickets, young forest, and forest-edgehabitats. Habitat use by coyotes inthe South is diverse and reflectstheir opportunistic feeding habits.

Their diet includes rabbits, smallmammals, ground-nesting birds andtheir eggs, amphibians, lizards, fish,snails, crustaceans, insects, carrion,fruits, and plant roots (Chapmanand Laerm, in press).

There are no known threats to coyotesurvival in the region. Animal damagecontrol programs in the Western UnitedStates have been unsuccessful.

Species accounts: Florida panther—The Florida panther, one of 30subspecies presently recognized, isthe only subspecies of mountain lionremaining in the South. The speciesoriginally ranged from eastern Texaseastward through Arkansas, Louisiana,Mississippi, Alabama, Florida, Georgia,and portions of Tennessee and SouthCarolina. Due to large-scale habitatdestruction and indiscriminant

shooting, panthers were extirpatedthroughout most of their range bythe early 1900s. Although periodicsightings are reported in remote areas ofselected States, it is unlikely that viablepopulations exist outside of Florida.Currently, the population is estimatedat between 20 and 50 animals.

Panthers prefer large remote tractsthat are typically heavily vegetatedand have minimal human disturbance.These animals use highly diversehabitats including hardwoodhammocks, saw-palmetto woodlands,sawgrass prairies, cypress strands,and oak-pine woodlands. Homeranges average 200 square miles formales and 75 square miles for females.

Panthers subsist on a variety ofmammalian prey, particularly white-tailed deer and feral hogs. In thenorthern portion of its range, feralhogs constitute the bulk of the diet,whereas white-tailed deer are moreimportant in the southern portion.Panthers also readily take raccoons,armadillos, rabbits, and other smallanimals (Clark 2001).

Loss of habitat is the greatest threatto viable panther populations, butillegal shooting and highway collisionsalso are major problems. Off-roadvehicle traffic has increased, makingaccessible large areas that formerlyhad been isolated wilderness. Intolerantof human disturbance, panthers aresensitive to habitat fragmentationstemming from road construction,agricultural development, and urbanexpansion. Other threats includeparasites, diseases such as felinedistemper and upper respiratoryinfections, and inbreeding depression.Panther populations are losinggenetic diversity by 3 to 7 percentper generation; at this rate, extinctionis probable in the next few decades(Clark 2001). Reduced prey base alsois a concern. Panthers consume up toone deer or hog weekly. Due to habitatalteration, these prey animals may notbe sufficiently abundant in Floridato meet this need.

Since panther habitat includes publicand private land, management effortsmust be coordinated. The key topanther conservation is habitatprotection and acquisition of large,interconnected blocks of woodland.The recovery plan recommends:(1) enhancing existing populations


through genetic management includingcaptive breeding programs and geneticrestoration; (2) protecting andmanaging existing habitat, includingprescribed burning and exotic plantcontrol; (3) establishing public supportby educating private landowners; and(4) reintroducing panthers into areas ofsuitable habitat (U.S. Fish and WildlifeService 1995). Potential release sitesinclude the lower Coastal Plain ofAlabama, Mississippi, Arkansas, andLouisiana and the lower ApalachicolaRiver in Florida.

Species accounts: gray fox andred fox—Foxes occur throughoutVirginia, North Carolina, SouthCarolina, Georgia, Alabama,Mississippi, Louisiana, and easternTexas. The gray fox does not occurin coastal Louisiana or the FloridaKeys, while the red fox does not inhabitthe southern Florida peninsula. Thepopulation density of red and grayfoxes in the South is not known.

Foxes occur in a variety of habitats.The red fox prefers open habitatsincluding old fields, shrublands,pastures, and mixed hardwood forests;the gray fox is more of a woodland-edge species. Both prefer areassupporting an interspersion of differentvegetative communities. Hollow logs,trees, brush piles, and rock outcropsare often used as dens. Patterns ofhabitat use change seasonally withfood availability.

Foxes are opportunistic feeders.During the fall and winter, smallanimals comprise the bulk of their diet.Common prey includes rabbits, voles,mice, wood rats, and various birds(Fritzell 1987). Fruits, berries,arthropods, and amphibians are addedto the diet during the summer and fall.

The planting of blackberry,honeysuckle, and other soft mastenhances fox habitat. Prescribedburning maintains old fields and forestsin desirable condition. Cultivationof trees that produce hard mast alsois important.

Trapping, hunting, road kills, andrabies are the major causes of foxmortality. The decline in red foxpopulations in some areas of the Southhas been attributed to interspecificinteraction with coyotes.

Species accounts: gray bat—TheU.S. Fish and Wildlife Service lists thisspecies as endangered. The species

distribution in the South includes thecave regions of Alabama, Arkansas,Kentucky, and Tennessee, but a fewoccur in Florida, Georgia, northeasternOklahoma, Mississippi, Virginia, andNorth Carolina. Bat populations havebecome fragmented during the pastfew decades (Harvey and Saugey2001). Ninety-five percent of graybats hibernate in 10 caves.

Gray bats are year-round caveresidents but usually occupy differentcaves in summer and winter. During thewinter, they hibernate primarily in deepvertical caves with large rooms acting ascold-air traps (42 to 52 o F). Maternityroosts are established in warm, humidcaves that provide domed ceilingscapable of trapping body heat from batclusters. Less than 5 percent of availablecaves in the South have the rightproperties of temperature, humidity,and structure to make them suitablefor gray bat occupation (Harveyand Saugey 2001).

Like many bats, this species huntsfor insects above forested riversand streams. Moths, beetles, flies,mosquitoes, mayflies, and otherinsects are important in the diet.

The primary reasons for populationdeclines include disturbance,vandalism, cave destruction, andpollution. Disturbance duringhibernation depletes energy reservesand increases mortality. Conservationactions focus on the protection ofoccupied caves and appropriatemanagement of the surrounding forestand aquatic foraging sites. Cave gatesand fences must be properly designedto allow bat movement. This speciesis recovering due to the protectionof four critical caves (Harvey andSaugey 2001).

Species accounts: Indiana bat—The Indiana bat is listed as endangeredby the U.S. Fish and Wildlife Service.This species is known to occurthroughout much of the Midwesternand Eastern United States; however,it has been virtually eliminated frommuch of its former range. The batoccurs in the northern portions of theSouth, including Tennessee, Virginia,and Kentucky. Isolated sightings havebeen made in the Carolinas, Alabama,and Mississippi. The current populationof the species nationwide is estimatedat 400,000 individuals; approximately

85 percent of the population is limitedto 7 caves (Harvey and Saugey 2001).

During the summer, maternity roostsare established between exfoliating barkand the bole of snags, in hollow trees,or in live trees. Male bats often usepitch pine and shortleaf pines. Thesebats need winter caves or minesretaining stable temperatures of 39 to46 oF and standing water that maintainsrelative humidity. The bats forage abovestreams, water bodies, and open areas.Riparian, upland, and floodplain forestsare also used.

During hibernation, the Indiana batis extremely vulnerable to any typeof disturbance. Factors contributingto its decline include cave disturbance,improperly designed cave gates, andintentional killing. Habitat lossstemming from deforestation andstream channelization is anotherconcern. Natural elements that imperilthe species include flooding ofoccupied caves, exposure to freezingtemperatures, and cave ceiling collapse.Forest management centers on theprovision of summer roost sites andforaging habitat.

Species accounts: mink—Minkoccur throughout the South, withthe exception of central Florida andwestern Texas. They are common inthe marshes along the Atlantic and GulfCoasts and are widespread in Virginia,North Carolina, and South Carolina(Chapman and Laerm, in press).Population densities vary with thetype and permanence of aquatic habitatand are influenced by climate, trapping,and intraspecific interaction.

Mink require wetland habitats, suchas marshes, swamps, riverbanks, andstreams. Habitat use varies bygeographic area and season. Thereare no published data on mink homeranges or habitat use patterns in theSouth. Muskrats, mice, and lagomorphsare the preferred prey; mink diets alsoinclude birds, amphibians, crawfish,and fish.

Habitat degradation as a resultof wetland alteration is a concernin the South. Mink are vulnerableto environmental contaminants,particularly mercury and pesticideresidues, concentrated in prey foods.The prevention of high levels ofenvironmental contaminants isneeded to ensure habitat qualityfor this species.


TERRESTRIAL

Species accounts: Ozark big-earedbat and Virginia big-eared bat—Thesetwo subspecies are endangered and arefederally protected throughout theirrespective ranges. Only a few cavesin eastern Oklahoma, Arkansas, andMissouri are known habitats for theOzark subspecies. The Virginia batinhabits eastern Kentucky, Virginia,North Carolina, and West Virginia,but fewer than five caves are knownto contain nursery colonies of thissubspecies (Harvey and Saugey 2001).

The bats inhabit caves in limestoneand schist formations throughout theyear. Adjacent land use does not appearto influence cave selection. Roostingsites are often near mature bottomlandand upland hardwood forests adjacentto water. Important habitat featuresinclude hollow trees, loose bark, androck shelters. The bats prefer relativelycold, well-ventilated locations and areoften found near cave entrances whenhibernating. Big-eared bats forage inforested areas among the canopies oflarge trees, consuming beetles, flies,mosquitoes, gnats, moths, and manyother insects.

The species is vulnerable to pesticidesand human disturbance of their caves.They are easily disturbed and quick totake flight. Conservation actions centeron the protection of roosting sites andthe retention of hollow trees.

Species accounts: red wolf—Thered wolf is an endangered species.The original distribution of the wolfincluded southern Illinois, Indiana,and Pennsylvania south to Florida andwest to southern Texas. Indiscriminatetrapping, hunting, and poisoning, lossof habitat, and expansion of urban andagricultural areas contributed to thedemise of this species. The last remnantpopulations in the wild were verifiedin southern Louisiana and Texas inthe 1970s.

In the late 1980s, efforts were madeto translocate wolves to five locations:Alligator River National WildlifeRefuge, North Carolina; Bull’s Island,South Carolina; St. Vincent Island,Florida; Horn Island, Mississippi; andthe Great Smoky Mountains NationalPark. Recent threats center on geneticdilution due to hybridization withwild dogs and coyotes.

Historically, the wolf was foundin old-growth forests, pine forests,bottomland hardwood forests, coastal

prairies, and marshes. Currentinformation on wolf ecology is limitedto studies in the coastal marshes ofTexas and Louisiana during the 1960sand 1970s and to observations atrestoration sites (Crawford and others2001). Heavy vegetative cover alongbayous and fallow fields is ideal habitat.Home ranges vary from 17 to 38 squaremiles, depending upon habitat andprey density. Red wolves requirelarge tracts of land relatively freeof human development, paved roads,and livestock.

Red wolves are opportunisticpredators, preying upon feral pigs,white-tailed deer, nutria, easterncottontails, swamp rabbits, marshrice rats, and fox squirrels. They willalso eat birds, rodents, frogs, andturtles. A diversity of prey is necessaryfor sustaining population levels.

The recovery plan objectives centeron the achievement of populationlevels large enough to ensure geneticintegrity (U.S. Fish and Wildlife Service1989). Potential reintroduction sitesare examined for biological factors(prey abundance, habitat types) andsocioeconomic factors (agriculturalpractices, land ownership patterns,proximity of towns). Areas of at least170,000 acres are required by thisspecies. The absence of coyotes ispreferable to avoid hybridization.Site considerations include the potentialfor wolf-livestock interaction andhuman disturbance. Public attitudesabout wolves are significant factorsin their recovery.

Species accounts: river otter—The river otter is listed as a threatenedspecies in Tennessee and as a speciesof concern in Oklahoma and Virginia.Otters occur regionally in many habitatsassociated with waterways, and theirnumbers are increasing in some partsof the region. The species is increasingin abundance throughout Virginia,where it is most common in the CoastalPlain and Piedmont. It also is relativelycommon in western Tennessee. Reliablecensus procedures for the river otterhave not been developed, and fewresearchers have attempted to estimatepopulation levels.

River otters use a variety of aquatichabitats including coastal estuaries,marshes, and streams. Riparian andshoreline vegetation bordering water-ways is an important component ofriver otter habitat. Beaver impound-

ments, submerged trees, and logjamsprovide shelter and foraging areasfor otters. Otters feed primarily onfish; other foods include aquaticinsects, birds, small mammals,snakes, and amphibians.

Threats to otter populations includethe clearing of bottomland forests,wetland modification, and pollutionof aquatic environments. Otters arefrequently caught in traps intended forbeaver; the low reproductive potentialof the otter, and the restricted nature ofits habitat make the species susceptibleto overharvest. As a result of trappingpressure, the otter was given protectionunder the Convention on InternationalTrade in Wild Species of EndangeredFlora and Fauna.

Strict population monitoring isneeded. Continued managementincludes the restoration of otterpopulations in Kentucky, Oklahoma,Tennessee, and North Carolina.Reintroduction in the Great SmokyMountains National Park began inthe 1980s, where otter populationswere once extirpated.

Species accounts: white-taileddeer—Deer are widespread andrelatively abundant throughout theSouthern United States. Populationson some islands have declined.Deer populations have fluctuateddramatically since European settlementof the South. Populations in the pastdeclined to critical levels becauseof intensive hunting, widespreadagricultural clearing, and other habitatalteration. Populations have reboundedduring the last several decades due tofarm abandonment, lower huntingpressure, and the extirpation of largepredators. In some locations,populations are increasing to levelsthat make the species a pest.

The endangered Key deer is restrictedto the lower Florida Keys. Four othersubspecies of concern occur on Sapeloand Blackbeard Islands in Georgia andon Hilton Head Island, Bull’s Island,and Hunting Island in South Carolina.

White-tailed deer use a wide range ofhabitat types and benefit from a mosaicof wetlands, forests, farmland, and earlysuccessional habitats. Preferred foodsare acorns, blueberries, sumac, grapes,hawthorns, common persimmons,dwarf palmettos, and blackberries.

There are no threats to the survivalof the white-tailed deer in the region.


8

23

5

10

11

7

6

13 11

11

26

15

15

3

10

2

9

11

1

2

10 9

9

16

5

7

However, coastal development hascontributed to the decline of the islandsubspecies. Key deer are threatenedby habitat loss, poaching, vehicularaccidents, and attacks by feral dogs(White and others 1998).

Discussion andConclusions

Based on listings from the U.S.Department of Interior (2000), everySouthern State contains species thatare under Federal protection (figs. 5.7and 5.8). The endangered categoryrefers to species that are in dangerof extinction in the foreseeable futurethroughout significant portions of theirrange. The threatened designation isassigned to species likely to becomeendangered in the future. Statusdeterminations are based onmodification or restriction of habitat,commercial overutilization, diseaseor predation pressure, the inadequacyof existing regulations, and other factorsaffecting continued existence.

There are a number of differentexplanations for the number of listedspecies in a State. A State may supportmany unique habitats with high speciesrichness. Texas is the largest State in theSouth in both area and species richness.The wide range of environmentalconditions and diverse habitats thatoccur in Texas also support the secondhighest level of protected species.Larger areas on average support agreater diversity of habitats and a widervariety of species, listed or otherwise.

A species that has been extirpatedfrom adjacent States may persist inareas that support the last remnantsof suitable habitat. For example, the redwolf formerly ranged from Texas to theAtlantic Coast. It presently occurs inNorth Carolina, Tennessee, and Florida,where it has been reintroduced. TheFlorida panther, another far-rangingmammal, once occurred throughout theregion. This species presently is foundsolely in isolated areas in Florida.

A high number of listed species mayalso reflect an inherently fragile fauna,such as that in the high-elevationhabitats of the Southern Appalachians.It also may reflect a high level ofendemic species, such as thoseassociated with scrub habitats of centralFlorida. Finally, the number of listedspecies in a State may reflect

deteriorating environmental conditionsand modification of natural ecosystems,such as longleaf pine forests. Each ofthese factors contributes to the numberof federally protected species in aState. Each reason has bearing on howhabitat is managed and protected.

Various natural and human-causedfactors contribute to a speciesimperilment. Some species occur in avery localized geographic area or in afew isolated areas of suitable habitat.These narrowly restricted species tendto be vulnerable to local disturbancesthat would have little effect on specieswith wide ranges. The summits and thebogs of the Southern Appalachian

Mountains support some highlyvulnerable species, such as the northernflying squirrel and the water shrew.

Scattered populations in fragmentedhabitat can be at risk. They becomedemographically isolated because theyhave little or no interaction with otherpopulations. These isolated populationsare prone to inbreeding depressionand genetic drift, which inhibit viability.Localized populations are alsovulnerable to catastrophic eventssuch as floods, droughts, and fires.

Many species have declined becauseof habitat alteration stemming fromhuman activities. These species are

Figure 5.8—The distribution of terrestrial vertebrateslisted as threatened throughout the South (U. S.Department of the Interior 2000).

Figure 5.7—The distribution of terrestrial vertebrateslisted as endangered throughout the South (U. S.Department of the Interior 2000).


TERRESTRIAL

unable to adapt due to changes inhabitat features such as vegetativecomposition and structure andwater quality.

Several factors repeatedly surfaceas threats to terrestrial vertebrates.The most prevalent factor is humandevelopment for urban, industrial, andagricultural land uses. Environmentalcontamination is a second prominentthreat, especially in the SouthernAppalachians and along the gulf coast.Coastal development contributes toendangerment on both the gulf andAtlantic Coasts. Exploitation occursprimarily on shorelines and in coastalwetlands. Other factors contributingto species endangerment include firesuppression, introduction of exoticspecies, and the loss of aquatic andwetland habitats.

Habitat loss affects all species,including migrating birds, wide-rangingmammals, and species like the gophertortoise, which cannot disperse overlong distances. Imperiling factorsinfluence species unequally. Turtlesare especially vulnerable to humanexploitation for food and pets. Batsand snakes are heavily impacted byhuman disturbance. Beavers and riverotters are imperiled by channelmodification and impoundments.Environmental contaminants impactthe spruce-fir forests used by thenorthern flying squirrel and the high-elevation mountain streams occupiedby a diversity of salamanders. Theuse of agricultural pesticides affectsgamebirds, bats, and amphibians.Wetland alteration affects theMississippi sandhill crane, mink,and several species of frogs and toads.Lastly, coastal development negativelyinfluences the habitat of thesoutheastern beach mice, woodstorks, marine turtles, and Key deer.

Often, it is difficult to identify aspecific factor responsible for thechanges observed in a species popu-lation. For example, many migratorybirds that breed in the South are alsodependent on wintering habitatsoutside of the country. Neotropicalmigrants are influenced by the lossof wintering habitat in the tropics,while wintering mallard populationsare affected by breeding habitat inthe prairie pothole region. Therefore,it is vital to understand the temporaland spatial context in which a speciesoccurs. Local changes in the population

of species may be a result of dramaticchanges in habitat occurring elsewhere.

Maintaining viable populations ofsouthern vertebrate species requiresthe protection of critical habitat as wellas the proactive management of otherhabitat. Public lands have a key rolein species conservation (chapter 1).In some instances, protecting sensitivehabitats from further alteration is thebest management action. In otherinstances, active enhancement maybe the most appropriate action. Forexample, treatments may be neededto increase understory growth, createmultiple seral stages, restore uniquehabitats, and control exotic species.Professional foresters, resourcemanagers, and conservationists playan important role in this regard.

There have been notable successstories in managing southernvertebrates. Restrictions on pesticideshave improved the status of baldeagles. Red-cockaded woodpeckershave benefited from the managementof mature pine forests, provision ofartificial cavities, and translocationefforts. River otters and beavers havebeen restored to areas they formerlyinhabited. Alligator populations haverebounded because of managementof harvest levels and the protectionof wetlands. Many of these specieshave proven far more resilient andadaptable than once thought.

However, additional efforts arenecessary to restore and enhanceecosystem integrity and resiliency onthe southern landscape. Managementplans should consider the assemblageof reptiles, amphibians, birds, andmammals. Herpetofauna havetraditionally received less managementattention than other vertebrates.Wetland buffers, travel corridors, andforest composition are important fortheir viability. Many species are long-lived and late maturing, and haverestricted geographic ranges; theirmanagement requires differentstrategies than those used for birdsand mammals. Management remainssomewhat hindered, however, bythe limited knowledge about thestatus of terrestrial vertebrates andtheir habitat relationships.

Land ownership patterns associatedwith the occurrence of southernspecies have management implications.Approximately 90 percent of the land

in the South is privately owned. Theprotection and management of specieshabitats can no longer be relegatedsolely to public land. To be successful,comprehensive conservation strategiesrequire the cooperation of privatelandowners. Cooperative forestryprograms and county extensionservices are two sources of expertisethat contribute to the managementof private lands.

In the past few decades, residentialand industrial areas have grownrapidly to serve an expanding southernpopulation. Although the extent ofsouthern forests has remained relativelystable in recent years, human andwildlife interactions have increased,and they will continue to do so. Publicperceptions about particular speciescan hinder or foster conservationefforts, highlighting the role ofenvironmental education.

One role for wildlife professionalsin the South is to identity the speciesthat face imperilment, determine theactions necessary to eliminate thosethreats, and then take the necessaryactions. Another role is to provideand manage habitat for several gamespecies. The many species inhabitingthe southern landscape have a widevariety of habitat requirements; anunderstanding of these requirementscan lead to management plans thatpromote viable populations andhabitat enhancement.

Needs for AdditionalResearch

Further research is needed on thestatus, distribution, population trends,and habitat requirements of manysouthern species. Although there arestandardized inventories for birdand game species across the region,there is a lack of comparablemonitoring protocols for manyother species. The importance ofregional monitoring and long-termresearch cannot be overstated.

Additional data are necessary toexamine the attributes that make somespecies associations resistant or resilientto disturbance. We need to understandwhy some associations are more fragilethan others. We also need to know howto mitigate negative disturbance factors.

Habitat relationships of listed andimperiled species need further study.


Examination of the connectionsbetween landscape patterns, land uses,and the presence or absence of concernspecies also would prove beneficial. Theestablishment of regional databases andstandardized sampling protocols formonitoring trends of terrestrial speciesacross all public lands also is needed.

A profound need exists for thecoordination of regional inventorieson public lands to monitor the statusand trends of reptile and amphibianpopulations. Assemblages associatedwith specific habitats need tobe identified.

Further research is necessary on thedistribution, ecology, and life historyof herpetofaunal species and commu-nities. In particular, additional data areneeded on species such as the flatwoodssalamander, gopher frog, southernhognose snake, and pine snake.This basic information is essentialto developing land managementprograms for these species.

Additional research is neededto determine the impact of naturaland human-caused factors on thedevelopment and environment ofamphibians. Additional informationneeds include the identificationof critical habitats and migrationroutes. The concern over amphibiandeclines highlights the lack of basicinformation about these species.

The ecology of furbearers, such asmink and weasel, is poorly understood,as are the potential impacts on othercarnivores resulting from coyoteexpansion throughout the South.Basic ecological data are neededon free-ranging red wolves to addressthe challenges of restoration. Thedegradation of river otter habitatsuggests the need for continuedmonitoring to ensure populationviability. Careful monitoring of blackbear populations also is essentialto ensure their continued existenceover the long term.

Finally, there is a paucity of infor-mation about specific habitat needsfor several bat species and the influenceof different silvicultural treatments ontheir populations.

Acknowledgments

Special thanks to John Greis andDavid Wear for their encouragementand support during the assessment

process. John Pye graciously handledthe Internet conversion ofthe document.

Appreciation is extended to TomDarden and his staff in the SouthernRegional Office for their ongoingsupport and assistance. An earlier draftbenefited from insightful comments byDennis Krusac and George Bukenhofer,USDA Forest Service; and fouranonymous scientific reviewers.Their comments strengthened andimproved the final manuscript.

My appreciation also is extended tothe regional coordinators of Partners inFlight for supplying conservation plansand other bird information: RobertFord, Dean Demarest, Jane Fitzgerald,and Ken Rosenburg. Recognition also isoffered to Chuck Hunter, U.S. Fish andWildlife Service, for supplying the birdphysiographic region map and draftsof his upcoming publications.

Kenneth Graham and Joe Johnston,U.S. Fish and Wildlife Service, suppliedrecovery plans and other informationon listed species. Craig Rudolph, USDAForest Service, provided the Louisianapine snake account. John Jensen,Georgia Department of NaturalResources, supplied publicationson herpetofauna. Robert Curry, U.S.Fish and Wildlife Service, providedinformation on bats. Special thanksto James Dickson, Louisiana StateUniversity, for supplying chapters ofhis upcoming textbook on southernwildlife prior to publication.

Paris Griep, USDA Forest Service,surveyed State agency biologists forinformation on the current status andhabitat concerns for black bear. Thesebiologists included Joe Hemphill(Oklahoma), Jason Plaxico (Kentucky),Denny Martin (Virginia), GordonWarburton (North Carolina), TomEason (Texas), Keith Guyse (Alabama),Rick Eastridge (Arkansas), CathyShropshire (Mississippi), andNathan Garner (Texas).

I also thank the other questionmanagers and participants (generalpublic, industry, agencies, andorganizations) for sharing theirideas and concerns during Assess-ment meetings.

Finally, my appreciation is offered toseveral USDA Forest Service individualswho assisted with the Internet searchesand the production of tables andfigures. The manuscript would not

have been possible without their help.These individuals include Pal Mattox,Mike Donahue, Christine Gabbard,Brenda Ayers, Sara Patterson, KaySpangler, and Ron Underwood.

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The USDA prohibits discrimination in all its programsand activities on the basis of race, color, national origin,sex, religion, political beliefs, sexual orientation, ormarital or familial status (Not all prohibited bases applyto all programs). Persons with disabilities who requirealternative means for communication of programinformation (Braille, large print, audiotape, etc.) shouldcontact the USDA's TARGET Center at 202-720-2600(voice and TDD).

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In: Wear, David N.; Greis, John G., eds. 2002. Southern forest resourceassessment. Gen. Tech. Rep. SRS-53. Asheville, NC: U.S. Departmentof Agriculture, Forest Service, Southern Research Station. 635 p.

The southern forest resource assessment provides a comprehensiveanalysis of the history, status, and likely future of forests in the SouthernUnited States. Twenty-three chapters address questions regarding social/economic systems, terrestrial ecosystems, water and aquatic ecosystems,forest health, and timber management; 2 additional chapters provide abackground on history and fire. Each chapter surveys pertinent literatureand data, assesses conditions, identifies research needs, and examinesthe implications for southern forests and the benefits that they provide.

Keywords: Conservation, forest sustainability, integrated assessment.

Chapter 5 (TERRA-5): Maintaining Species in the South170 amphibians, 197 reptiles, 595 birds, and...

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