Chapter 1 (TERRA-1): Terrestrial Ecosystems › sustain › report › pdf › chapter_01e.pdf · Chapter 1: Terrestrial Ecosystems 3 TERRESTRIAL Since presettlement, there have been

Chapter 1: Terrestrial Ecosystems 3TERRESTRIAL

■ Since presettlement, there havebeen significant losses of communitybiodiversity in the South (Noss andothers 1995). Fourteen communitiesare critically endangered (greaterthan 98-percent decline), 25 areendangered (85- to 98-percentdecline), and 11 are threatened(70- to 84-percent decline). Commonfactors contributing to the loss ofthese communities include urbandevelopment, fire suppression,exotic species invasion, andrecreational activity.

■ The term “fragmentation”references the insularization ofhabitat on a landscape. The changein arrangement of remaining habitatscan be accompanied by a loss ofhabitat area. Habitat fragmentationcan result in the decline of interior-dwelling birds; the decline of somelarge, wide-ranging species; andthe loss of other specialized species.Habitat fragmentation affects thepatch, connectivity, and edgecharacteristics of a landscape.

■ Connectivity within a landscapemay facilitate movement andfecundity for some species, whilethe size and shape of landscapepatches influences the integrityof both biotic and abiotic processes.Edge characteristics also haveimportant implications forthe persistence of an arrayof terrestrial species with verydifferent habitat requirements.

■ The availability of hardand soft mast can influencesome terrestrial vertebratespecies. Mast is an essentialcomponent in the diet of manybirds and mammals. Disease,

insect infestation, advanced age,climatic processes, and distur-bance influence mast yields.

■ The ranges of many speciescross both public and privateland ownerships. The numbers ofimperiled and endangered speciesinhabiting private land indicate itscritical importance for conservation.

■ The significance of land owner-ship in the South for the provisionof species habitat cannot beoverstated. Each major landownerhas an important role to play inthe conservation of species andtheir habitats.

Introduction

The South has an impressivediversity of terrestrial communitiesand species associations. Thesecommunities range from mountainspruce-fir forests to tropical hardwoods,and from coastal dunes to prairies.Centuries of settlement and land usechange have brought a number ofthreats and pressures. The majorityof the landscape has been modifiedconsiderably, resulting in thedisappearance, degradation, andendangerment of native communities.

This chapter assesses the historicaland present status of terrestrial speciesacross the South. It is organized intosix major sections:

1. An overview of southern historicalconditions affecting terrestrialvertebrate species.

2. A review of populations, harvests,and the conservation status of speciesoccurring in the South.

Key Findings

■ There are 132 terrestrial vertebratespecies that are considered to beof conservation concern in the Southby State Natural Heritage agencies.Of the species that warrant conser-vation focus, 3 percent are classedas critically imperiled, 3 percentas imperiled, and 6 percent asvulnerable. Eighty-six percentof terrestrial vertebrate speciesare designated as relatively secure.The remaining 2 percent are eitherknown or presumed to be extinct,or have questionable status.

■ Species of conservation concernare dominated by amphibians andreptiles. Fifty-four amphibians,40 reptiles, 20 birds, and 18mammals are classed as imperiled.

■ Increasing population trends arereported for wild turkey, white-taileddeer, and black bear. Populationsof northern bobwhite quail, gray fox,and red squirrels have declined forseveral years. There have alsobeen declines in mourning doveand American woodcock populations.Cottontail rabbit and ruffed grousepopulations have demonstratedcyclical patterns. Among themigratory game birds, recordharvests of ducks and geese haveoccurred in recent years.

■ Groups of nongame birds withmore than 50 percent of their speciesshowing significant declining trendsinclude grassland-nesting birds(70 percent), ground-nesting birds(57 percent), and shrubland-nestingbirds (53 percent).

Chapter 1:TerrestrialEcosystems

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

What are the history,status, and projectedfuture of terrestrial

wildlife habitat types andspecies in the South?

Southern Forest Resource Assessment4

TERRESTRIAL

3. A review of selected sensitivecommunities in the region and thecommon threats to these communities.

4. An overview of vertebrate speciesthat consume hard and soft mast. Thissection also lists several mast-producingspecies that occur in the South.

5. An evaluation of the significance ofpublic and other land for maintainingspecies and their habitats.

6. A review of the literature onfragmentation and its influence onlandscapes and the species supportedby those landscapes.

Several species are included that,at one stage or another of their lives,return to land to reproduce or spenda part of their lives there. The focusis on vertebrates because informationon the regional biogeography of manyterrestrial invertebrate groups islacking (Echternacht and Harris 1993).Scientific names are provided in thechapter tables; therefore, commonnames will be used in the text. (Note:Additional information on the statusand habitat relationships of vertebrateresources across the South is providedin chapters 5 and 23, which includediscussions of threatened andendangered species.)

Methods andData Sources

Data on the conservation statusof terrestrial vertebrate species werecompiled from State Natural Heritageagencies using NatureServe (2000).The Natural Heritage database isan inventory of known occurrencesfor species of conservation concern,including federally listed species. Steinand others (2000) list multiple criteriaused by Natural Heritage for assessingconservation status: occurrence(number of distinct populations orsubpopulations); condition (viabilityof extant populations); population size;area of occupied habitat; short- andlong-term population trends; knownor suspected threats; susceptibilityto intrinsic biological factors; andthe number of protected occurrences.This methodology provides the basisfor conservation status designationsthat indicate the degree of imperilment.

Species known to be extinct (GX),or possibly extinct (GH), are recordedindependently. For example, the

passenger pigeon is assigned the GXranking because there is no questionabout its extinction. For a considerablenumber of species that have not beenobserved in many years, however,there remains some hope of rediscovery.That, for example, is the case forBachman’s warbler. These specieswere assigned the status of GH.

Information on game and furbearerabundance was obtained from theRenewable Resources Planning Act(RPA) Wildlife Report (Flather andothers 1999). The RPA is a periodicassessment of natural resources onthe Nation’s forests and rangelands.The RPA data on game populationsoriginated from State agencies usingquestionnaires developed by theUSDA Forest Service and the NaturalResources Conservation Service.Data from the RPA assessments aretaken from various State and Federalagencies. Population projections ofharvested animals are based on surveysof experts from State wildlife agencies.

Information on rare and threatenedcommunities was based on thecomprehensive reviews conductedby Grossman and others (1994),Noss and others (1995), White andothers (1998), and Walker (2001).

Information on the acreage anddistribution of Federal land wasobtained from the National Parksindex (U.S. Department of the Interior2000a), the Lands Report from the Fishand Wildlife Service (U.S. Departmentof Interior 2000b), and the LandsArea Report of the USDA Forest Service(U. S. Department of Agriculture,Forest Service 2000c). Agency reportsalso were compiled for national parks(U.S. Department of the Interior, ParkService 2000) and national refuges(U.S. Department of the Interior, Fishand Wildlife Service 2000), providingproperty descriptions and species lists.

Statewide timberland ownershipdata were obtained from the ForestInventory and Analysis ResearchWork Unit (FIA) of the SouthernResearch Station (U.S. Departmentof Agriculture, Forest Service 2000a).For each State, the acres in bothpublic and private ownershipcategories were analyzed.

A literature search was conductedfor information on fragmentation,rare communities, historical conditions,and species habitat relationships.In addition, research stations and

universities throughout the Southwere contacted to obtain additionalinformation. The results from thiseffort were combined with additionalinformation obtained from severalplant and animal field guides. A list ofmast-producing species was compiledusing vegetation guides; terrestrialvertebrate species that include mastas a component of their diet wereextracted from wildlife field guides.

Results

Historical ConditionsThe presettlement landscape of

the South was quite diverse: forestsof different ages were interspersedwith expansive savannas, densecane thickets, barrens, and swamps.Disturbance was a major influence onthe composition of southern forests,creating forest openings and resettingsuccession (Lorimer 2001). Forestswere dynamic; natural successionprogressed with shade-tolerant plantsreplacing pioneer species. Periodicflooding and associated sedimentationinfluenced the distribution andcomposition of local areas.

Frequent thunderstorms provideda source of natural fires, resultingin a landscape of mixed speciescomposition. Lightning fires burnedunabated (Williams 1989). Firefrequency and intensity were dominantforces (refer to chapter 25). Fire wasimportant for the persistence of manycommunities including pine forests,oak-hickory forests, savannas, barrens,and prairies (Trani and others 2001).

Native Americans, through use offire and crop cultivation (Buckner1989, Delcourt and Delcourt 1987),further modified the composition andopen character of the forest. Fires werefrequently set to create openings forcrops and to drive game for harvest.The effects of native inhabitationon southern forests were extensive(refer to chapter 24).

Wildlife of the presettlement Southwas quite impressive. Dickson (2001)describes large herds of bison andelk roaming throughout the prairiesand savannas of the region. White-tailed deer and wild turkey also werenumerous. Large carnivores (blackbear, cougar, red wolf, and bobcat)were abundant, and a diversity ofsuccessional seres supported a variety


Table 1.1—Terrestrial vertebrate species classified as presumed or possibly extinct in the South

Scientific name Common name Former areas of occurrence

Presumed extinctConuropsis carolinensis Carolina parakeet AL, AR, FL, GA, LA, MS, NC, OK, SC, TN,

TX, VAEctopistes migratorius Passenger pigeon AL, AR, FL, GA, LA, MS, NC, OK, SC, TN,

TX, VAMonachus tropicalis West Indian monk seal FL

Possibly extinctCampephilus principalis Ivory-billed woodpecker AL, AR, FL, GA, LA, MS, NC, OK, SC, TN, TXEurycea troglodytes Valdina farms sinkhole salamander TXPlethodon ainsworthi A plethodontid salamander MSVermivora bachmanii Bachman’s warbler AL, MS, OK, SC, TN, VA

Source: NatureServe 2000.

of prey populations. Other mammalsincluded mink, muskrat, river otter,beaver, gray fox, red fox, spottedskunk, long-tailed weasel, bats,and numerous small mammals.

Birds present in today’s forests alsowere likely present during presettle-ment (Dickson 2001). Raptors suchas the Mississippi kite, bald eagle,osprey, red-shouldered hawk, andbarred owl were likely occupantsof historic bottomland forests. TheSwainson’s and Bachman’s warblersinhabited cane thickets, while theyellow-breasted chat and indigobunting populated young forests.Cavity-nesting birds such as red-headed woodpeckers, Americankestrels, and great crested flycatcherswere abundant in the old-growthforests of eastern Texas (Truettand Lay 1984). The ivory-billedwoodpecker thrived in oak-gumforests, foraging on snags for insects.

Early records of reptiles andamphibians are limited, but theserecords make frequent referenceto rattlesnakes and alligators (Dickson2001). Historic forest habitats appearto have supported viable, diversepopulations of herpetofauna (Gibbonsand Buhlman 2001).

Extensive inundated bottomlandforests supported habitat for millionsof wood ducks and mallards(Heitmeyer 2001). Wood duckscommonly nested in the cavitiesof abundant old-growth forests.Hooded mergansers, green-wingedteal, gadwall, and American widgeonalso frequented flooded bottoms.

The southern landscape changeddramatically with the advent ofEuropean settlers. Settlement resultedin the extensive clearing of forest andconversion of the land to pasture orcropland (DeGraaf and Miller 1996).These lands were often managed withfire, which was also used to maintainsavannas and other open areas in theEast (Williams 1989). In particular,fire was used to create favorablegrazing conditions for domesticanimals (Healy 1985).

By 1819, all land was claimed eastof the Mississippi River (Dickson2001). Natural resources were treatedas if they were inexhaustible. Forestswere cut with little thought for forestregeneration, and soils were seriouslydepleted through erosion and excessivecropping. Wildlife species and theirhabitats were likewise exploitedwithout concern for their persistence.The decline in abundance of wildlifethat occurred during the last half ofthe 19th century remains unparalleledin the history of the South.

Deer populations nationwideplummeted to fewer than a millionanimals by 1900 (Dickson 2001).Bison and elk disappeared from theregion. The wild turkey disappearedfrom several States within its range.The wood duck was drastically reducedby indiscriminate harvest. Populationsof large carnivores, regarded as threatsto livestock and people, were deci-mated, and viable populations of blackbear and cougar were relegatedto relatively remote areas.

The loss of bottomland forest inthe Mississippi Alluvial Valley affectedwaterfowl and other species thatwere displaced into adjacent areas.Harvests of the passenger pigeonand the Carolina parakeet for marketled to their demise in the early 1900s(table 1.1). Market hunting, thedomestication of land, and the harvestof mature forests without regenerationled to the extirpation of some speciesin various Southern States (table 1.2).(Note: It is possible that some specieswere extirpated because their range ison the periphery of the region. Theirloss may be related to random effectsassociated with low populations atthe edges of their ranges.)

During the 1930s and 1940s,the States recognized the dire statusof wildlife populations and initiatedefforts to address the problem.The Duck Stamp Act (1934), thePittman-Robertson Act (1937),and the Dingle-Johnson Act (1950)apportioned funds to States forwildlife restoration projects, habitatacquisition, and research.

These efforts came too late forsome species (table 1.1). The ivory-billed woodpecker foraged in maturebottomland hardwoods along theAtlantic and Gulf coasts. Its dietconsisted of wood-boring insectlarvae occurring in dead and dyingtrees. Overhunting and intensiveharvesting of virgin hardwood forestsbetween the 1880s and 1920s ledto the decline of this species (U.S.Department of the Interior 1973).


TERRESTRIAL

Table 1.2—Vertebrate species extirpated from selected States within the South

Scientific name Common name Former areas of occurrence

MammalsRodents

Erethizon dorsatum Common porcupine NC, VAMicrotus ochrogaster Prairie vole LA

CarnivoresCanis lupus Gray wolf AR, GA, KY, NC, OK, TN, TX, VACanus rufus Red wolf AL, AR, FL, GA, KY, LA, OK, TX, VALeopardus pardalis Ocelot AR, LALeopardus wiedii Margay TXMartes pennanti Fisher NC, TNMustela nigripes Black-footed ferret OKPanthera onca Jaguar; otorongo LAPuma concolor Mountain lion ALUrsus arctos Grizzly or brown bear OK, TX

Other mammalsBos bison American bison AL, AR, FL, GA, KY, LA, MS, NC, OK, SC,

TN, VACervus elaphus Wapiti or elk AL, AR, GA, KY, LA, NC, OK, SC, TN, VALepus americanus Snowshoe hare NC

BirdsWading birds

Grus americana Whooping crane AR, FL, KYWaterfowl

Cygnus buccinator Trumpeter swan KY, LAShorebirds

Bartramia longicauda Upland sandpiper TNNumenus borealis Eskimo curlew OK, SC

Perching birdsCorvus corax Common raven AL

Other birdsAnhinga anhinga Anhinga KYCentrocercus urophasianus Sage grouse KS, OKGeotrygon chrysia Key West quail-dove FLTympanuchus cupido Greater prairie chicken AR, KY, LA, TNTympanuchus phasianellus Sharp-tailed grouse OK, TXZenaida aurita Zenaida dove FL

ReptilesSnakes

Masticophis flagellum Coachwhip KY

Source: NatureServe 2000.

Bachman’s warbler, last observed inthe 1960s, once inhabited Arkansas,Kentucky, Alabama, South Carolina,Louisiana, and Missouri. The extensiveclearing of bamboo and canebrakehabitat for agriculture along theMississippi River and West Gulf CoastalPlains bottoms degraded the winteringand breeding habitat for this species(Ehrlich and others 1992). Excessive

collecting for the millinery trade mayalso have contributed to the decline.

The Valdina Farms salamander wasendemic to Texas. The amphibianoccurred in isolated, intermittent pools.It is now extinct due to flooding ofits only known habitat. Populationsof the West Indian monk seal, whichoriginally inhabited the Florida coast,were decimated during the 19th century.

The major factor in its exterminationwas over-hunting, principally forblubber (to make oil) and for meat.The seal’s inherent tameness increasedits vulnerability to slaughter.

The last four decades of the 20th

century brought legislation thatfurthered species conservation efforts,including the Wilderness Act (1964),the Endangered Species Act (1966,


Other 1%

Vulnerable 6%

Imperiled 3%

Criticallyimperiled 3%

Presumedpossibly

extinct 1%

Secure/apparentlysecure 86% Imperiled (G2)

Critically imperiled (G1)

0 10 20 30 40 50 60

Birds

Amphibians

Mammals

Reptiles

Number of species

Presumed/possibly extinct (GX/GH)

Vulnerable (G3)

1969, and 1973), the NationalEnvironmental Policy Act (1970), theMarine Mammal Protection Act (1971),and the National Forest ManagementAct (1976). Through these and severalother conservation efforts, conditionsfor many species have improved acrossthe South (Dickson 2001). However,the loss and modification of uniqueforest communities continues to affectpopulations of other species. Theremainder of this chapter examinesthese influences, presenting the trendsfor a diversity of southern species.

Status and Trends ofTerrestrial VertebrateSpecies

Conservation status ranks forsouthern species—The databasesof the State Natural Heritage agencieswere used to derive a regional specieslist of global (G) conservation ranks.The G ranks reflect a species’ raritythroughout its range. For example,a species holding the G conservationranking of G1 in Virginia also carriesthe same rank elsewhere in the Nation.

These ranks are: GX (presumedextinct: intensive search has not locatedadditional populations); GH (possiblyextinct: historically known and may be

rediscovered); G1 [critically imperiledglobally because of extreme rarity(observations include 5 or fewerlocations or fewer than 1,000 animals)]or because some factor of its biologymakes it vulnerable to extinction];G2 [imperiled globally because ofrarity (observations reflect 6 to 20locations or 1,000 to 3,000 animals)]or because of other factors makingit vulnerable to extinction]; G3[vulnerable globally because of raritythroughout its range (observationsinclude 21 to 100 locations or 3,000to 10,000 animals) or because it isfound locally in a restricted area];G4 (apparently secure globally,although the species may be rare inparts of its range, especially at theperiphery; usually more than 100occurrences and 10,000 individuals);and G5 (secure globally: observationsare common and widespread).

Figure 1.1 shows the proportionof vertebrate taxa in each of theconservation ranking categories.One hundred thirty-two species areconsidered to be of conservationconcern. Among terrestrial vertebrates,28 species are classified as criticallyimperiled, 37 species as imperiled,and 67 species as vulnerable. Eighty-six percent of southern terrestrial

vertebrate species are designated asrelatively secure by Natural Heritage.

Figure 1.2 shows species rankedas presumed or possibly extinct,critically imperiled, imperiled, orvulnerable among the four majorvertebrate taxa. Collectively, thesespecies represent animals withelevated risks of extinction orof conservation concern.

The proportion of species at riskvaries greatly among taxonomicgroups. Forty-one percent of imperiledspecies are amphibians, followedby reptiles (30 percent), birds (15percent), and mammals (14 percent).With the exception of mammals,the number of species at risk withineach taxonomic group is notproportionate with their respectiverichness in the region. For example,amphibian species comprise only14 percent of the terrestrial vertebratesoccurring in the South, yet theycomprise 41 percent of the imperiledspecies list. Conversely, bird speciescomprise 48 percent of southernterrestrial vertebrates, but only15 percent of the imperiled species.Refer to chapter 5 for additionaldata on regional species richness.

Figure 1.1—Proportion of southern terrestrial vertebrate species at risk.The Other category includes species that have not been ranked or havequestionable status (NatureServe 2000).

Figure 1.2—Number of terrestrial vertebrate species at riskdelineated by major taxa in the South (NatureServe 2000).


TERRESTRIAL

Table 1.3—Amphibian species within the South with global rankings of G1, G2, and G3

Scientific name Common name Areas of occurrence

Frogs and toadsG1

Bufo houstonensis Houston toad TXG2

Rana okaloosae Florida bog frog FLG3

Rana capito Gopher frog AL, FL, GA, LA, MS, NC, SC, TNSalamanders

G1Desmognathus sp.1 Waterrock knob salamander NCEurycea latitans Cascade caverns salamander TXEurycea nana San Marcos salamander TXEurycea neotenes Texas salamander TXEurycea rathbuni Texas blind salamander TXEurycea robusta Blanco blind salamander TXEurycea sosorum Barton Springs salamander TXEurycea sp. 1 Jollyville Plateau salamander TXEurycea sp. 2 Salado Springs salamander TXEurycea sp. 4 Buttercup Creek caves salamander TXEurycea sp. 5 Georgetown salamander TXEurycea sp. 6 Pedernales River spring salamander TXEurycea sp. 7 Edwards Plateau spring salamander TXEurycea sp. 8 Comal Springs salamander TXEurycea tridentifera Comal Blind salamander TXPlethodon petraeus Pigeon Mountain salamander GAPlethodon shenandoah Shenandoah salamander VANotophthalmus meridionalis Black-spotted newt TX

G2Ambystoma cingulatum Flatwoods salamander AL, FL, GA, SCDesmognathus carolinensis Carolina mountain dusky salamander NC, TNDesmognathus ocoee Ocoee salamander AL, GA, NC, SC, TNDesmognathus orestes Blue Ridge dusky salamander NC, VAEurycea pterophila Blanco River Springs salamander TXGyrinophilus palleucus Tennessee cave salamander AL, GA, TNHaideotriton wallacei Georgia blind salamander FL, GAPhaeognathus hubrichti Red hills salamander ALPlethodon aureolus Tellico salamander NC, TNPlethodon caddoensis Caddo Mountain salamander ARPlethodon fourchensis Fourche Mountain salamander ARPlethodon hubrichti Peaks of Otter salamander VAPlethodon ouachitae Rich Mountain salamander AR,OKPlethodon virginia Shenandoah mountain salamander VANecturus alabamensis Black warrior waterdog ALNotophthalmus perstriatus Striped newt FL, GASiren sp. 1 Lesser siren (Rio Grande population) TX

G3Amphiuma pholeter One-toed amphiuma AL, FL, GA, MSAneides aeneus Green salamander AL, GA, KY, MS, NC, SC, TN, VADesmognathus aeneus Seepage salamander AL, GA, NC, SC, TNDesmognathus apalachicolae Apalachicola dusky salamander AL, FL, GADesmognathus brimleyorum Ouachita dusky salamander AR, OKDesmognathus imitator Imitator salamander NC, TN

(continued)


Table 1.3—Amphibian species within the South with global rankings of G1, G2, and G3 (continued)


Salamanders (cont.)G3 (cont.)

Desmognathus santeetlah Santeetlah dusky salamander NC, TNDesmognathus wrighti Pigmy salamander NC, TN, VAEurycea junaluska Junaluska salamander NC, TNEurycea sp. 9 Sandhills salamander NCEurycea tynerensis Oklahoma salamander AR, OKPlethodon punctatus White-spotted salamander VAPlethodon teyahalee Southern Appalachian salamander GA, NC, TNPlethodon websteri Webster’s salamander AL, GA, LA, MS, SCPlethodon welleri Weller’s salamander NC, TN, VANecturus lewisi Neuse River waterdog NC

G1 = critically imperiled; G2 = imperiled; G3 = vulnerable.Source: NatureServe 2000.

The conservation status of individualspecies are presented in tables 1.3, 1.4,1.5, and 1.6. Several of these species arediscussed in further detail in chapters 5and 23, including the factorsinfluencing imperilment and specieshabitat relationships. Species that arefederally listed as threatened orendangered are discussed in chapter 5.

Fifty-four amphibian speciesare of conservation concern (table1.3). Salamanders dominate with 51listings; frogs and toads have 3 listings.Examples include the Houston toad,gopher frog, flatwoods salamander,Ocoee salamander, green salamander,and several species in the Plethodon,Desmognathus, and Eurycea genera.

Forty reptile species are imperiledor vulnerable (table 1.4). Reptilesubgroups with global rankings ofconcern include turtles (19), lizards(10), snakes (9), and others (2).Oceanic and map turtles dominatethis list. Other reptiles of conservationconcern include the alligator snappingturtle, bog turtle, gopher tortoise,glass lizard, southern hognose snake,and crocodile.

Twenty avian species are of concern(table 1.5). Subtaxa include 2 wadingbirds, 3 shorebirds, 6 perching birds,and 9 others. Several of these speciesinclude the whooping crane, pipingplover, Bachman’s sparrow, Floridascrub jay, red-cockaded woodpecker,and lesser prairie chicken.

Eighteen mammal species areimperiled or vulnerable (table 1.6).

Mammalian subtaxa with globalrankings of concern include 5 bats,8 rodents, 3 carnivores, and 2 others.Bats are represented by the Indianabat, Rafinesque’s big-eared bat,southeastern myotis, and severalother species. Additional mammalsinclude the Allegheny wood rat,red wolf, and swift fox.

Population and harvest trendsfor southern species—The regionalpopulation and harvest trendspresented in this section, unlessotherwise stated, originated fromthe RPA (Flather and others 1999).The RPA represented the best sourceof quantitative data on regional trendsfor multiple species at the time ofthis Assessment. Information wascollected from cooperating Statewildlife agencies. Population estimateswere summed across those Statesthat provided data. (The list of Statesthat provided population estimatesis available at the Rocky MountainResearch Station, Fort Collins, CO.)The absence of data from certainStates resulted from variation in thedistribution of species or the lack ofdata for certain years. The RPA includedonly States that provided estimatesfor 1975 to 1990 (in 5-year intervals)and 1993 in the trend analysis.

Projections were based on a weightedaverage percentage change from 1993to the year 2000 and 2045 for Statesthat provided projection estimates.The average percentage change wasthen applied to the 1993 populationestimate in order to extrapolate a total

projected population for Statesthat provided population estimates(Flather and others 1999).

Population and harvest trendsfor southern species: big gamespecies—Big game species areprimarily large mammals takenfor sport or subsistence. Because ofState agency convention, the wildturkey also is included. The speciescomprising big game were the firstto stimulate widespread publicinterest in wildlife conservation.For this reason, historical informationabout game species is extensive forseveral States.

Wild turkey populations haveconsistently increased since 1975(fig. 1.3). Five States project that turkeypopulations will decline over the nextfour decades (Flather and others 1999).

For States reporting on white-taileddeer, populations have increasedapproximately fourfold since 1975(fig. 1.4). There is concern amongState personnel that deer may becomea management problem during thenext decade. Seven States expect deernumbers to decline slightly over thenext 50 years (Flather and others1999). (Additional information on deeris provided in chapters 3, 4, and 5.)

The trend in black bear numbers ispositive for the four States reporting(fig. 1.5). Biologists from these Statesexpect bear populations to declinesomewhat over the next few decades(Flather and others 1999). (Note: TheFlorida and Louisiana subspecies of


TERRESTRIAL

Table 1.4—Reptile species within the South with global rankings of G1, G2, and G3


TurtlesG1

Lepidochelys kempii Kemp’s or Atlantic ridley AL, FL, GS, LA, MS, NC, TX, VAPseudemys alabamensis Alabama redbelly turtle AL, FL, MS

G2Sternotherus depressus Flattened musk turtle ALGraptemys barbouri Barbour’s map turtle AL, FL, GAGraptemys ernsti Escambia map turtle AL, FLGraptemys flavimaculata Yellow-blotched map turtle MSGraptemys oculifera Ringed map turtle LA, MS

G3Macroclemys temminckii Alligator snapping turtle AL, AR, FL, GA, KY, LA, MO, MS, OK, TN, TXCaretta caretta Loggerhead AL, FL, GA, LA, MS, NC, SC, TX, VAChelonia mydas Green turtle AL, FL, GA, LA, MS, SC, TX, VAEretmochelys imbricata Hawksbill AL, FL, GA, LA, MS, NC, SC, TXDermochelys coriacea Leatherback tinglar AL, FL, GA, LA, MS, NC, TX, VAKinosternon hirtipes Mexican mud turtle TXClemmys muhlenbergii Bog turtle GA, NC, SC, TN, VAGopherus polyphemus Gopher tortoise AL, FL, GA, LA, MS, SCGraptemys caglei Cagle’s map turtle TXGraptemys gibbonsi Pascagoula map turtle LA, MSGraptemys nigrinoda Black-knobbed map turtle AL, MSTrachemys gaigeae Big bend slider TX

LizardsG2

Sceloporus arenicolus Sand dune lizard TXNeoseps reynoldsi Sand skink FL

G3Crotaphytus reticulatus Reticulate collared lizard TXHolbrookia lacerata Spot-tailed earless lizard TXHolbrookia propinqua Keeled earless lizard TXSceloporus woodi Florida scrub lizard FLColeonyx reticulatus Reticulated gecko TXCnemidophorus dixoni Gray-checkered whiptail TXOphisaurus compressus Island glass lizard FL, GA, SCOphisaurus mimicus Mimic glass lizard AL, FL, GA, MS, NC, SC

SnakesG1

Tantilla oolitica Rim Rock crowned snake FLG2

Clonophis kirtlandii Kirtland’s snake KYHeterodon simus Southern hognose snake AL, FL, GA, MS, NC, SCNerodia harteri Brazos water snake TXNerodia paucimaculata Concho water snake TX

G3Pituophis ruthveni Louisiana pine snake LA, TXStilosoma exenuatum Short-tailed snake FLTantilla atriceps Mexican blackhead snake TXSistrurus catenatus Massasauga OK, TX

Other reptilesG2

Crocodylus acutus American crocodile FLG3

Caiman crocodilus Spectacled caiman FL, GA

G1 = critically imperiled; G2 = imperiled; G3 = vulnerable.Source: NatureServe 2000.


Table 1.5—Bird species within the South with global rankings of G1, G2, and G3


Wading birdsG1

Grus Americana Whooping crane AL, GA, LA, OK, TXG3

Phoenicopterus ruber Greater flamingo FLShorebirds

G1Numenus borealis Eskimo curlew AR, LA, NC, TX

G2Charadrius montanus Mountain plover OK, TX

G3Charadrius melodus Piping plover AL, AR, FL, GA, KY, LA, MS, NC, OK, TN,

TX, VAPerching birds

G2Dendroica chrysoparia Golden-cheeked warbler TXVireo atricapillus Black-capped vireo MS, OK, TX

G3Aimophila aestivalis Bachman’s sparrow AL, AR, FL, GA, KY, LA, MS, NC, OK, SC,

TN, TX, VAAphelocoma coerulescens Florida scrub jay FLPipilo alberti Albert’s towhee TXVermivora crissalis Colima warbler TX

Other birdsG1

Pterodroma feae Fea’s petrel NCPterodroma hasitata Black-capped petrel FL, GA, NC, VA

G2Amazona viridigenalis Red-crowned parrot FL, TXa

G3Columba leucocephala White-crowned pigeon FL, TXPelecanus erythrorhynchos American white pelican AL, AR, FL, GA, KY, LA, MS, NC, OK, TN,

TXPicoides borealis Red-cockaded woodpecker AL, AR, FL, GA, KY, LA, MS, NC, OK, SC,

TN, TX, VAStrix occidentalis Spotted owl TXa

Thalassarche chlororhynchos Yellow-nosed albatross FL, NCTympanuchus pallidicinctus Lesser prairie chicken OK, TX

G1 = critically imperiled; G2 = imperiled; G3 = vulnerable.a West Texas.Source: NatureServe 2000.

black bear, of conservation concernin the region, are discussed separatelyin chapter 5.)

Population and harvest trendsfor southern species: small gamespecies—Species classified as smallgame typically include residentgame birds and mammals that areassociated with upland (forest, range,or agricultural) habitats. There is somevariation among State wildlife agencies

as to which species are managedas small game. In this chapter, quail,grouse, rabbits, and squirrels areconsidered small game. Few Statewildlife agencies monitor smallgame populations; therefore, thetrends reviewed here should beinterpreted carefully.

The populations of gray, red, andfox squirrels have been declining inthe South since 1985 (fig. 1.6).

Cottontail rabbit populations declinedslightly between 1975 and 1980(fig. 1.7), but recovered by 1990.One State projects that cottontail rabbitpopulations may decline by 2045(Flather and others 1999).

Northern bobwhite quail populationshave declined from 1975 to the present(fig. 1.8). Among the States reportingtrends in bobwhite abundance,populations have declined by nearly


TERRESTRIAL

Table 1.6—Mammal species within the South with global rankings of G1, G2, and G3


BatsG2

Myotis sodalis Indiana or social myotis AL, AR, KY, NC, OK, SC, TN, VAG3

Corynorhinus rafinesquii Rafinesque’s big-eared bat AL, AR, FL, GA, KY, LA, MS, NC, OK, SC,TN, TX, VA

Myotis austroriparius Southeastern myotis AL, AR, FL, GA, KY, LA, MS, NC, OK, SC,TN, TX, VA

Myotis grisescens Gray myotis AL, AR, FL, GA, KY, OK, SC, TN, VAMyotis leibii Eastern small-footed myotis AL, AR, GA, KY, NC, OK, SC, TN, VA

RodentsG1

Dipodomys elator Texas kangaroo rat OK, TXG2

Geomys texensis Llano pocket gopher TXG3

Tamias canipes Gray-footed chipmunk TXGeomys arenarius Desert pocket gopher TXGeomys knoxjonesi Jones’ pocket gopher TXNeofiber alleni Round-tailed muskrat FL, GANeotoma magister Allegheny woodrat AL, KY, NC, TN, VAPodomys floridanus Florida mouse FL

CarnivoresG1

Canus rufus Red wolf NC, SC, TNG3

Vulpes velox Swift fox OK, TXPanthera onca Jaguar; otorongo TX

Other mammalsG2

Trichecchus manatus Manatee FL, GA, LA, MS, NC, SC, TX, VAG3

Antilope cervicapra Blackbuck TXa

G1 = critically imperiled; G2 = imperiled; G3 = vulnerable.a Exotic.Source: NatureServe 2000.

50 percent, from 23 million birdsin 1975 to 12 million birds in 1993(Flather and others 1999). Forest(ruffed) grouse populations show acyclical pattern, but appear to havedeclined since 1985 (fig. 1.9).

Bobwhite quail trends from theBreeding Bird Survey (BBS) areconsistent with State agency estimates(Flather and others 1999). BBS datasuggest that the abundance of thisspecies has declined significantly(P < 0.05) in the South. Bobwhitenumbers have declined by 2.6 percentper year from 1966 to 1996, and havedeclined at an even greater rate since1985 (-5.6 percent per year).

State agency projections for mostsmall game species suggest minorchanges in future population status.Forest grouse are expected to remainstable. State biologists forecastdeclines for bobwhite quail, squirrels,and cottontails.

Population and harvest trends forsouthern species: migratory gamebirds—Migratory game birds includewaterfowl, such as ducks and geese,and other migratory species, suchas mourning doves and woodcock.The long history of migratory birdmanagement in North America hasresulted in an impressive monitoringsystem. Population and harvest trends

originate from annual reports publishedby the U.S. Fish and Wildlife Serviceand the North American WaterfowlPlan (Flather and others 1999).

Waterfowl trends are traditionallytracked by major flyways, which arethe migration routes from breedingto wintering habitat. In the South,the major routes are the Atlanticand Mississippi flyways (fig. 1.10).National duck harvests have beenrecorded since the early 1960s.

Over the last 25 years, 41 percentof the national harvest was taken inthe Mississippi flyway and 15 percentfrom the Atlantic flyway. Both had largeharvests during the 1970s, followed by


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Figure 1.3—Population trends of wild turkey in Southern Statesthat provided estimates and long-term projections [based onState wildlife agency data (Flather and others 1999)].

Figure 1.4—Population trends of deer in Southern States thatprovided estimates and long-term projections [based on Statewildlife agency data (Flather and others 1999)].

Figure 1.5—Population trends of black bear in Southern Statesthat provided estimates and long-term projections [based onState wildlife agency data (Flather and others 1999)].

Figure 1.8—Population trends of northern bobwhite quailin Southern States that provided estimates and long-termprojections [based on State wildlife agency data (Flatherand others 1999)].

Figure 1.7—Population trends of cottontail rabbits in SouthernStates that provided estimates and long-term projections [basedon State wildlife agency data (Flather and others 1999)].


TERRESTRIAL

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Figure 1.9—Population trends offorest grouse in Southern Statesthat provided estimates and long-term projections [based on Statewildlife agency data (Flather andothers 1999)].

Figure 1.11—Trends in goose harvest from 1965to 1995 by administrative flyway encompassingthe South (Flather and others 1999).

Figure 1.10—Trends in duck harvest from 1965to 1995 by administrative flyway encompassingthe South (Flather and others 1999).

substantial declines through muchof the 1980s, and substantial harvestincreases during the 1990s. Duckharvests in the Mississippi flywayincreased by 260 percent from 1988to 1995, with a record 6.6 millionducks harvested in 1995 (Flather andothers 1999).

Trends in goose abundance werederived from surveys conductedin migration and wintering areas.Record numbers of geese wereharvested for three consecutive yearsstarting in 1993 along the Mississippiflyway (fig. 1.11). After reaching apeak harvest of about 550,000 birdsin 1983, the goose harvest in theAtlantic flyway declined to nearly180,000 birds in 1995.

Management units are traditionallyused by agencies to report populationtrends of mourning doves andAmerican woodcock. Both speciesare monitored using call-count surveys,which provide an index of populationsize. National trends in populationindices for both species show evidenceof declines, but the magnitude of thedecline is greater for woodcock thanfor mourning doves. This pattern isconfirmed by BBS data, which indicate


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that doves declined annually at a rate of0.3 percent compared to a 3.2 percentdecline for woodcock over the 30-yearperiod (Flather and others 1999).

Mourning dove calling countsindicate declining populations duringthe last 10 years in the eastern andcentral management units (fig. 1.12).Intensive agricultural practices may beinfluencing the breeding populationsthroughout much of the bird’s range(Brady and others 1998). The acreageof agricultural land in the easternmanagement unit is positively related todove populations because agriculturalfields provide the forest edge habitatpreferred by doves. Increased herbicideuse and crop rotation may havecontributed to observed declines(Martin and Sauer 1993). In the centralmanagement unit, the trend towardfewer and larger farms also mayhave influenced dove populations.

Call-count trends for woodcockshow similar declines in both theeastern and central management units(fig. 1.13). Trends since 1968 indicatethat the number of woodcock heardhave declined by 2.5 percent per yearin the eastern unit and 1.6 percentper year in the central unit (Flatherand others 1999). In the last decade,this rate of decline has accelerated.Woodcock select early successionalhardwood forests interspersed withfields and forest openings. As withthe mourning dove, the widespreaddecline in woodcock may be linkedwith habitat alteration due to forestsuccession and land use intensification(Straw and others 1994).

Population and harvest trendsfor southern species: furbearerspecies—There are few comprehensiveexaminations of trends in furbearerpopulations nationwide. Often, theonly available data are temporal harvesttrends that reflect fur prices ratherthan population status. The limitedinformation on population trendsmakes furbearer projections uncertain.

The RPA used a compilation offurbearer status reports completedfor the International Association ofFish and Wildlife Agencies during1993. A survey of State agencybiologists provided populationprojections to 2003 (SouthwickAssociates. 1993. 1993 State andprovincial survey of furbearerswith emphasis on nuisance animals.Unpublished report. On file with:

Figure 1.12—Population trends in mourning dove from 1966 to 1996by management unit (Flather and others 1999).

Figure 1.13—Population trends in woodcock from 1968 to 1996by management unit (Flather and others 1999).


TERRESTRIAL

DecreasingStableIncreasing



Figure 1.14—Projected trends of beaver populations in the South[based on State wildlife agency data (Flather and others 1999)]. Statesthat provided estimates are shaded.

Figure 1.16—Projected trends of muskrat populations in the South[based on State wildlife agency data (Flather and others 1999)].States that provided estimates are shaded.

Rocky Mountain Research Station,2150 Center Avenue, Fort Collins,CO 80526).

Population projections of southernfurbearers are shown in figs. 1.14, 1.15,1.16, 1.17, 1.18, and 1.19. Of the10 Southern States reporting beaverpopulation projections, 5 expectedpopulation increases through 2003(fig. 1.14). The beaver population isprojected to decline in North Carolina,and remain stable (or increase) inthe remainder of the South.

The majority of raccoon populationsare projected to increase or remainstable throughout the South (fig. 1.15).Exceptions occur in Alabama and NorthCarolina, where disease-caused declinesare projected (Flather and others 1999).

Of the four States reporting onmuskrat populations, two expectpopulation increases through 2003(fig. 1.16). The remaining States(Virginia and Tennessee) project stablepopulations. Projections on coyoteabundance are limited to Georgiaand Mississippi (fig. 1.17). BothStates report that coyote populationsare expected to remain stable.

Bobcat projections are reported onlyfor Florida and Oklahoma (fig. 1.18).Florida biologists report stable bobcatpopulations, while Oklahoma biologistsreport that bobcat populations areincreasing. Finally, the five States thatmade projections for red and gray foxes(Virginia, Kentucky, Tennessee, SouthCarolina, and Texas) predicted stablepopulations (fig. 1.19).

Population and harvest trends forsouthern species: nongame birds—In the United States, nongame birds arenot legally taken for sport, subsistence,or profit. Nongame species comprisethe majority of taxa that inhabit theSouth. There are few data sources onpopulations of nongame species.

Data from the BBS were used toprovide information on breeding birdtrends in the South for the RPA. Detailson the implementation of the BBS canbe found in Droege (1990); informationon statistical analyses can be found inSauer and others (1997). The relativeabundance trend for each bird specieswas summarized in two ways. First,the numbers of species with statisticallysignificant increasing, decreasing, orstable trends were estimated. Second,birds were grouped according to life-history characteristics including nest

Figure 1.15—Projected trends of raccoon populations in the South[based on State wildlife agency data (Flather and others 1999)].States that provided estimates are shaded.





Figure 1.17—Projected trends of coyote populations in the South[based on State wildlife agency data (Flather and others 1999)].States that provided estimates are shaded.

Figure 1.18—Projected trends of bobcat populations in the South[based on State wildlife agency data (Flather and others 1999)].States that provided estimates are shaded.

Figure 1.19—Projected trends of red and gray fox populationsin the South [based on State wildlife agency data (Flather andothers 1999)]. States that provided estimates are shaded.


TERRESTRIAL

Spe

cies

gro

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All birds

CavityOpen cup

GroundMidstory

NeotropicalShort distancePerm. resident

WoodlandShrublandGrassland

WetlandUrban

Proportion of species (percent)

0 20 40 60 80 100

Increasing Decreasing Stable

Table 1.7—Number of breeding bird species with increasing,decreasing, and stable trends from 1966 to 1996 by life historycharacteristics for the South

Life history Total Increasing Decreasing Stablecharacteristic species species species species

N - - - - - - - - - - - N (Percent) - - - - - - - - -

All species 210 47 (22.4) 74 (35.2) 89 (42.4)

Nest type/locationCavity 29 10 (34.5) 8 (27.6) 11 (37.9)Open cup 86 18 (20.9) 42 (48.8) 26 (30.2)Ground/low 54 7 (13.0) 31 (57.4) 16 (29.6)Midstory/canopy 65 20 (30.8) 20 (30.8) 25 (38.5)

Migration statusNeotropical 76 12 (15.8) 31 (40.8) 33 (43.4)Short distance 50 17 (34.0) 20 (40.0) 13 (26.0)Permanent resident 42 9 (21.4) 16 (38.1) 17 (40.5)

Breeding habitatWoodland 58 15 (25.9) 19 (32.8) 24 (41.4)Shrubland 43 8 (18.6) 13 (53.5) 12 (27.9)Grassland 10 7 (70.0) 3 (30.0)Wetland/open water 46 11 (23.9) 8 (17.4) 27 (58.7)Urban 13 2 (15.4) 6 (46.2) 5 (38.5)

Source: Flather and others 1999.

Number of species11 – 3435 – 4142 – 4849 – 5455 – 72

Figure 1.20—The proportion of southern bird species with increasing,decreasing, and stable trends from 1966 to 1996. Birds have been groupedby broad life-history characteristics, migration status, and breeding habitat(Flather and others 1999).

type (cavity or open cup), nest location(ground, low, midstory, or canopy),migration status (neotropical migrant,short-distance migrant, permanentresident), and breeding habitat(woodland, shrubland, grassland,wetland, urban). The resultingtrends are presented in figure 1.20.

Approximately 42.4 percent ofbreeding bird species appear stable,35.2 percent have declined, and22.4 percent have increased acrossthe South (table 1.7). It is worth notingthat Flather and others (1999) foundthat the percentage of declining specieswas greater in the South than inany other RPA region. Abundancetrends among species groupsvary considerably. Species withdeclining trends include 70 percentof grassland-nesting birds, 57 percentof ground-nesting birds, 53 percentof shrubland-nesting birds, 49 percentof open-cup nesting birds, 46 percentof urban-nesting birds, and 41 percentof neotropical migrants. Numbers ofthe majority of cavity-nesting speciesand wetland species have been stable.

Figure 1.21 suggests that bird speciesrichness is high along the SouthernAppalachians and along the AtlanticCoast from northeastern North Carolinato the Chesapeake Bay. Because somespecies are missed during bird countsurveys including nocturnal species,raptors, and absent migrants, itis important to note that the birdrichness estimates are biased low(Sauer and others 1997).

Raptors include hawks, falcons,eagles, vultures, and owls. In contrastto other bird species, raptors naturallyexist at relatively low populationdensities and are widely dispersedin their habitats. The natural scarcityof raptors, their ability to move quickly,and the difficulties of detection inhibitthe determination of population status(Fuller and others 1995).

As a group, raptors are poorlysurveyed, and quantitative data arelacking to determine their populationtrends. Table 1.8 presents a nationalsummary of the status and populationtrends of 33 species and subspeciesof southern raptors. Two species, theAmerican kestrel and burrowing owl,are declining across the United States.Mississippi kites, osprey, bald eagles,and peregrine falcons are increasing.Populations of 22 species areconsidered stable nationwide.

Figure 1.21—Patterns of birdrichness in the Southbased upon countsfrom the BreedingBird Survey (Flatherand others 1999).


Table 1.8—The national trends of raptors that occur in the South

Scientific name Common name Status/Trend/Comments

Accipiter cooperii Cooper’s hawk StableAccipiter gentilis Northern goshawk Unknown/C2a

Accipiter striatus Sharp-shinned hawk Stable/regional differencesAquila chrysaetos Golden eagle StableAsio acadicus Northern saw-whet owl StableAsio flammeus Short-eared owl Stable/local concernAsio otus Long-eared owl Stable/local concernAthene cunicularia Burrowing owl Declining/local concernBubo virginianus Great horned owl StableButeo brachyurus Short-tailed hawk Stable/northern range limit, about <500 birds in

U.S.Buteo lagopus Rough-legged hawk StableButeo lineatus Red-shouldered hawk Stable/local concernButeo jamaicensis Red-tailed hawk Stable/local increases; Breeding Bird Survey dataButeo platypterus Broad-winged hawk Stable/migration count decline in 1980sButeo regalis Ferruginous hawk Unknown/C2Buteo swainsoni Swainson’s hawk Unknown/C3;b local concernCaracara plancus Crested caracara Unknown/northern range limitCathartes aura Turkey vulture StableCircus cyaneus Northern harrier Stable/nomadic, no standard survey; local concernCoragyps atratus Black vulture Stable/population estimation difficultElanoides forficatus American swallow-tailed kite Stable/historical rangeFalco columbarius Merlin StableFalco peregrinus anatum American peregrine falcon Endangered; increasingFalco sparverius American kestrel Stable/Breeding Bird Survey DataFalco sparverius paulus American kestrel, Florida Declining/C2Haliaeetus leucocephalus Bald eagle Threatened or endangered in contiguous U.S.;

increasing/status reassessment underwayIctinia mississippiensis Mississippi kite Increasing/range expansionNyctea scandiaca Snowy owl StableOtus asio Eastern screech-owl StablePandion haliaetus Osprey Increasing/good informationRostrhamus sociabilis Snail kite Endangered, stable/northern range limitStrix varia Barred owl Stable/western range expansionTyto alba Common barn owl Stable/local concern

a Proposal to list; available data are not conclusive for threatened or endangered status.b Proven more widespread than previously believed or not subject to identifiable threat.Source: Fuller and others 1995.

The status of a raptor populationoften reflects changes in the availabilityof prey species. However, changes inraptor status also can indicate subtleenvironmental conditions, such aschemical contamination or disease.

Nesting ospreys are concentratedalong the Atlantic Coast. Most regionalpopulations declined through the early1970s. Following the nationwide banon DDT in 1972, osprey productivityimproved, and population numbersincreased in many areas. Ospreynumbers are stable, and in someareas they are increasing.

The endangered snail kite breedsin central and southern Floridawetlands, the northern extent ofthe range. The species declined from1900 to 1960. Populations remainrelatively stable today.

Bald eagle populations declineddramatically between 1950 and 1970.Illegal shooting, habitat alteration,and DDT adversely affected birdpopulations. The species was classifiedas endangered in 1978. Followingthe DDT ban, bald eagle reproductionimproved, and populations beganincreasing. The active protection

of nesting habitat and release ofhand-reared eagles aided this increase.Habitat loss remains a threat in manyareas (Fuller and others 1995).

Ferruginous hawk populations arestable in some areas, but decliningin others. Status determinationis complicated by the low densityof nesting birds and fluctuation inbreeding associated with cycles ofprey abundance.

The peregrine falcon also sufferedfrom contamination by DDT and otherorganochlorine pesticides. Peregrine


TERRESTRIAL

Table 1.9—Ecosystem communities that have declined by 70 percent ormore in the South since European settlement

Ecosystem type Geographic area

Critically endangered: >98 percent lossOld-growth deciduous forests SoutheastSouthern Appalachian spruce-fir Tennessee, North Carolina, VirginiaLongleaf pine forests and savannas Southeastern Coastal PlainSlash pine and rockland habitat Southern FloridaLoblolly-shortleaf pine forests West Gulf Coastal PlainCanebrakes SoutheastBluegrass savannah-woodland

and prairies KentuckyBlack Belt and Jackson prairies Alabama, MississippiUngrazed dry prairie FloridaWet and mesic coastal prairies LouisianaAtlantic white-cedar Virginia, North CarolinaNative prairies KentuckyBottomland forest West VirginiaHigh-quality oak-hickory Cumberland Plateau, Tennessee

Endangered: 85-98 percent lossRed spruce Central AppalachiansSpruce-fir forest West VirginiaUpland hardwoods Coastal Plain, TennesseeOld-growth oak-hickory TennesseeCedar glades TennesseeLongleaf pine Texas, LouisianaLongleaf pine forest, 1936-87 FloridaMississippi terrace prairie, calcareous

prairie, Fleming glades LouisianaLive oak, live oak-hickory LouisianaPrairie terrace-loess oak forest LouisianaMature forest, all types LouisianaShortleaf pine-oak-hickory LouisianaMixed hardwood-loblolly pine LouisianaXeric sandhill LouisianaStream terrace-sandy wooded-savannah LouisianaSlash pine FloridaGulf Coast pitcher-plant bogs Coastal PlainPocosins VirginiaMountain bogs North CarolinaAppalachian bogs Blue Ridge, TennesseeUpland wetlands Highland Rim, TennesseeUltramafic glades Virginia

Threatened: 70-84 percent lossBottomland and riparian forest SoutheastXeric scrub, scrubby flatwoods, sandhills Lake Wales Ridge, FloridaTropical hardwood hammock Florida KeysSaline prairie LouisianaUpland longleaf pine LouisianaLive oak-pine-magnolia LouisianaSpruce pine-hardwood flatwoods LouisianaXeric sandhill woodlands LouisianaFlatwood ponds LouisianaSlash pine-pondcypress-hardwood LouisianaWet hardwood-loblolly pine Louisiana

Source: Noss and others 1995. Based on the published literature, Natural Heritage programs, andexpert opinion.

recovery has been hastened in the Eastby the release of hundreds of birds bredin captivity; these birds survived andproduced young in the wild.

Sensitive and RareCommunities

Extent of threatenedcommunities—Several authors havedescribed and identified the threatenedand sensitive communities in theSouth (Boyce and Martin 1993,Grossman and others 1994, Nossand others 1995, White and others1998). The South supports a diversityof communities; a high proportionof them are considered imperiledto some degree (Walker 2001).

Noss and others (1995) listednumerous threatened and endangeredcommunities that have experiencedlosses in the South (table 1.9). Theamount of areal loss relative to theestimated presettlement area was usedas an indicator of vulnerability. The14 communities listed as criticallyendangered have estimated lossesof over 98 percent of their areasince European settlement. Theseinclude old-growth deciduous forest,spruce-fir forests, longleaf pinesavannas, bottomland forest,and several types of prairies. Twenty-five endangered communities haveexperienced losses between 85 and 98percent. These communities includeCoastal Plain hardwoods, pocosins,mountain bogs, ultramafic glades,and Louisiana prairies.

Having experienced over 70percent losses compared to estimatedpresettlement area, 11 communitiesare regarded as threatened. Theseinclude tropical hardwood hammocks,sandhill woodlands, and saline prairies.

In addition to the list in table 1.9,Noss and others (1995) reported24 communities that have lost atleast 50 percent of their area. Theseinclude pocosins (Coastal Plain), sandpine (Florida), baldcypress-tupelo(Mississippi, Tennessee), flatwoods-swale habitats (Florida), herbaceousmarsh (Florida), calcareous forest(Louisiana), scrub-shrub swamp(Louisiana), cove hardwood forest(Tennessee), and others.

Boyce and Martin (1993) alsorecognized several sensitive commu-nities that are under pressure from avariety of factors. Such factors included


Table 1.10—The Nature Conservancy’s summary of distributions and threats for rare communities of the South

Geographic Number ofarea Habitat communities Threats

Southern Spruce-fir 2 Nonindigenous species, recreation,Appalachian Beech 2 air pollution, past logging, hydrologicalMountains Bog, fen 7 alteration, succession.

Grassy bald 1Cliff, gorge 4Other 1

South Florida Tropical hardwood 2 Development, nonindigenous species,Slash pine 3 hydrological alteration, fire suppression,

burning, fragmentation, agriculture,recreation.

Coastal Plain Barrier island 9 Development, grazing, fragmentation,Longleaf pine 3 hydrological alteration, fire suppression,Other forests 3 nonindigenous species, agriculture, pastGlade, prairie 6 logging, mining, burning, recreation.

Continental Forest 7 Fire suppression, agriculture, recreation,Interior Glade, prairie 3 grazing, past logging, nonindigenous species,

Other 1 succession, mining, hydrological alteration.

Other Outcrop 1 Recreation, grazing, agriculture, hydrologicalForest 1 alteration, fire suppression.Canebrake 1

Source: Grossman and others 1994.

urban growth, land use conversion,water diversion, exotic species,and pesticide runoff. Everglades,mangroves, bottomland hardwoodforests, pocosins, mountain bogs,and Carolina bays were classifiedas threatened. They classified longleafpine, spruce-fir and other high-elevation forests, heath balds, maritimecommunities, rock outcrops, glades,grasslands, and sand-pine scrubas vulnerable.

Grossman and others (1994) listed57 rare communities in the South(table 1.10). Community types wereranked on a global scale based on thenumber of occurrences, areal extent,condition, threats, and fragility. These57 communities had global ranks of G1(found in 1 to 5 occurrences globally)or G2 (found in 6 to 10 occurrencesglobally). Twenty-one types occurin the Coastal Plain, 5 in south Florida,17 in the Southern Appalachians, and11 in the Continental Interior.

Communities can decline in arealextent or have their structuresimpoverished or compromised.

Communities covering smaller areastend to maintain smaller populationsthat are more vulnerable to extinctionthan larger populations (Soulé 1987).Communities also can lose vigorbecause of change in their structure,function, or composition. For example,intense livestock grazing entailsreplacement of native perennialgrasses with exotic annuals. Thefactors contributing to communityimperilment that are listed in table1.10 are further discussed in thefollowing section.

Profiles of selected rarecommunities—This section reviewssome selected communities ofconcern. Each general communitytype can include multiple associations.Each account includes distribution,composition, threats, and potentialmanagement. Where available, stepstoward restoration are presented.The accounts were developed fromWhite and others (1998), Boyceand Martin (1993), Noss and others(1995), and Walker (2001). Thediscussion of communities followsWhite and others (1998).

Profiles of selected rarecommunities: old-growth forests—Although forests predominate inthe South, less than 585,790 acresof old-growth forest exist (Whiteand others 1998). The remainingold-growth forests tend to be onsteeper, rockier, or mesic sites difficultto farm or harvest. Old-growth forestcomposition varies with forest type,but characteristics generally associatedwith old-growth forests include large,old trees; accumulations of woodydebris; and multilayered canopies.

Many vertebrate species occurin patches of old-growth forest.These include the Jefferson salamander,the Peaks of Otter salamander, theoak toad, and the scarlet king snake(Wilson 1995). Public lands such asthe Great Smoky Mountains NationalPark and several national forests protectsome of the largest tracts in the South.With the exception of these areas,old-growth remnants are often smallerthan 250 acres.

Threats to old-growth remnantsinclude invasions by nonindigenousspecies, interruption of natural


TERRESTRIAL

disturbance regimes, outbreaks offorest pests, and timber harvest(Walker 2001).

Management options vary by foresttype, but controlling nonindigenousspecies and herbivores, and choosingbenign methods to accomplish theseobjectives are factors to consider.Management actions that mimic naturaldisturbances are particularly importantbecause natural disturbance regimesare unlikely to be intact. Managementemphasis may also include theprovision of forested buffers aroundexisting old-growth remnants.

Profiles of selected rarecommunities: spruce-fir forests—The spruce-fir community is confinedto the highest peaks of Virginia,Tennessee, and North Carolina.Red spruce communities occur atan approximate elevation of 4,500feet. In the northern limit of its range,Fraser fir is replaced with balsam fir.This community is characterized byrelatively high moisture levels, shortgrowing seasons, acidic soils, andextreme weather conditions. Theflora is distinctive. The communityreproduces in small-scale patchesresulting from wind disturbance.

The presettlement extent of theSouthern Appalachian spruce-fircommunity has been estimated as30,000 to 35,000 acres (White andothers 1998). These remote forestsremained relatively undisturbed untilthe widespread harvests of the late1800s (White and others 1998). In1934, the majority of the remainingspruce-fir forest went into public pro-tection with the establishment of theGreat Smoky Mountains National Park.

Spruce-fir communities are threatenedby infestations of balsam woollyadelgids. The stresses induced by insectattack are exacerbated by additionalstresses of acid precipitation, whichinfluence soil and stream chemistry.Air pollution and the deposition ofheavy metals, such as lead, copper,zinc, nickel, and manganese, alsocontribute to the decline of thiscommunity (refer to chapter 18).They inhibit regeneration andcontaminate the understory. Airbornepollution is carried with prevailingwinds originating from industrialareas of southern Ohio and Indiana.

In addition, recreation activitiescompact soil and damage young trees.

As the southern population centersexpand, continued recreationalpressure may further adversely affectthe spruce-fir community.

Spruce-fir communities supportseveral terrestrial species that areuncommon elsewhere. Examplesinclude the endangered subspeciesof northern flying squirrel, Weller’ssalamander, the endangered spruce-fir moss spider, mountain ash, andthe threatened rock gnome lichen.The northern saw-whet owl, black-capped chickadee, and red crossbillalso inhabit the community.

Restoration centers on enhancingthe stocking of red spruce treesand increasing stand structuralcomplexity. Appropriate silviculturaltreatments include the release of sprucesaplings from the understory andthe removal of competing stems. Insome areas, restoration may involveconversion of open areas to forestsby planting seedlings.

Profiles of selected rarecommunities: wetlands, bogcomplexes, pocosins—In the lasttwo centuries, the Nation has lostapproximately 30 percent of itswetlands. Substantial losses haveoccurred along the southern CoastalPlain and along the lower reachesof the Mississippi River. In addition,Florida has lost 46 percent (9 millionacres) of its wetlands (Stein andothers 2000). Wetland loss is of specialconcern, because these habitats providecritical waterfowl and fish habitat.

Small wetlands occur in depressionsembedded in forested areas. Soils aresaturated for extended periods fromrainfall and ground water seepage.Among the most vulnerable areasare small (less than 2 acres), isolatedbogs that retain characteristic species.Bogs require distinct hydrologicalconditions to function ecologically.Intermittent fires and beaver activitiesmay contribute to the origin andmaintenance of this complex.

The exact number of remainingbogs is difficult to determine but ismost certainly fewer than 150 in theentire South. Over half of the existingbogs occur on private land, and arethreatened by development, grazing,off-road vehicle use, agriculturalpractices, and hydrological alteration.

Pocosins are freshwater wetlandsdominated by a dense cover of broad-

leaved evergreen shrubs or low-growingtrees. They have highly organic soilsthat developed in areas of poordrainage. This community occurs inupland interstream areas. Peat layersare thick, and vegetation is shrubby.

The bog complex provides habitatfor a diversity of herpetofauna. Wilson(1995) lists 37 species of reptiles andamphibians associated with Carolinabays, pocosins, and bogs in the South;41 are associated with swamp habitat.These species include the bullfrog,green frog, eastern tiger salamander,four-toed salamander, mountain chorusfrog, and snapping turtle. The bogturtle, threatened in the northernportion of its range, also inhabits theseareas. This turtle is collected illegally, asare rare orchids and carnivorous plants.Opportunities for species to recolonizeare minimal, and the community ispermanently diminished.

Avian species occurring in thesecommunities include cedar waxwing,Nashville warbler, northern water-thrush, purple finch, white-eyedvireo, and wood duck. Characteristicmammals include the long-tailed shrew,marsh rice rat, mink, muskrat, riverotter, southern bog lemming, southernshort-tailed shrew, and the star-nosedmole. Butterflies include the Atlantisfritillary and silver-bordered fritillary.

No vertebrates are endemic topocosins, but the community provideshabitat and refuge from adjacentlandscape development. In NorthCarolina, 41 species of mammalsinhabit pocosin and Carolina baysites (White and others 1998).

Conservation activities includeprotection from heavy equipment,off-road vehicles, and foot traffic;controlling changes in site hydrologyby providing buffers between adjacentsites, filling ditches and blockingdrains; and restricting livestock grazing.The retention of woody debris providesvaluable microhabitat for many species.Adjacent land management activitiesthat alter the surrounding watersheddegrade these sensitive communities.Restoration includes maintenance ofsite hydrology and woody plant control.Periodic prescribed burns adjustedto maintain vegetative conditions helpto maintain the community. Speciesreintroduction into selected sites alsomay be required.


Presettlement range of longleaf pineand transitional communities

Current extent of natural longleafpine stands older than 40 years

Profiles of selected rarecommunities: bottomland andfloodplain forests—The forestedwetlands of the Coastal Plain,Piedmont, and Continental InteriorProvinces include bottomlandhardwood forests and deepwateralluvial swamps. Bottomland hard-woods are located along waterwaysand in low-lying areas such as theMississippi Delta region. Common treespecies include ash, sycamore, watertupelo, cypress, willow, cottonwood,elm, oaks, river birch, silver maple,sweetgum, black walnut, and pine.Vegetative composition and structurevary with flooding duration. Treesare vulnerable to prolonged changesin hydrology and are characterized byrapid growth. Bottomland hardwoodsare found almost exclusively on alluvialsoils that are associated with oldriverbeds, existing streams, andimpoundments and their terraces.Soils are saturated year round or nearlyso; the understory is sparse with vinesand shrubby vegetation.

Beneficial characteristics of thiscommunity for wildlife includehard mast production, cavity treeprovision, and production of abundantinvertebrate biomass. In agriculturallandscapes, bottomland forests serveas refuges for many species. Speciesassociated with this community includewood stork, prothonotory warbler,marbled salamander, and the swamprabbit. The loss of bottomlandhardwood forests to agriculturalconversion contributed to the declineof the Carolina parakeet and the ivory-billed woodpecker (Dickson 2001).

Many bottomland sites are productiveand have been in agriculturalproduction for long periods. Severalcypress-oak reforestation projectsin the Mississippi Alluvial Valleyhave been successful in areas wherefrequent flooding precludes agriculturaldevelopment. Restoration of thiscommunity occurs primarily onpublic land.

Profiles of selected rarecommunities: glades, barrens,and prairies—Scattered throughoutthe South are naturally treeless areasreferred to as prairies, glades, andbarrens. Historical accounts suggestthat these open communities were oncewidespread (Delcourt and others 1993),but estimates of original extent areuncertain. These grass-dominated

communities occurred in the Piedmont,Interior Plateau, Ridge and Valley,and Coastal Plain Provinces.

Lightning fires, Native Americanburning, grazing by elk and bison,and soil conditions historicallymaintained these areas. Today, thesecommunities occupy only a fraction oftheir original extent due to agriculturalconversion, recreation use, exoticspecies invasions, fire exclusion,and the loss of large herbivores.

Forbs and grasses occurring onrocky or shallow soil dominate glades;composition varies with geology, soiltype, and soil depth (Walker 2001).The limestone glades of the Ozarks,dominated by perennial grasses, havea more open nature than glades of theInterior Low Plateau. Eastern redcedarwoodlands are commonly associatedwith glades of various types. Threatsto glade communities includeconstruction, quarrying, agriculture(pasture), fire suppression, andnonindigenous species invasion.

The barren and prairie communitiescontain the majority of the region’snative grasslands. In the South, theyinclude the Black, Jackson, and GrandPrairies. In these communities, grassesare dominant, and shrubs and treesare generally absent. The sites arehighly productive because they retainnutrients. As a result, they supporta vast array of animal and plant life.

Species composition varies with sitemoisture. Characteristic species includelittle bluestem, Indian grass, andbig bluestem. Composition variesdepending upon specific soil andgeologic types.

The size and isolation of these openareas preclude support of endemicvertebrates. Many rare species ofbirds, reptiles, and arthropods usethese communities. Vertebrate speciesthat have been extirpated from thesecommunities include the greaterprairie chicken, bison, and elk.

Restoration centers on the controlof woody species from adjacent foresthabitats and the use of prescribedburning to maintain the diversityof the grassland communities. Theretention of characteristic speciesrelies upon site-specific management.

Profiles of selected rarecommunities: longleaf pine andsouthern pinelands—Longleaf pinehistorically dominated Coastal Plainsites from southern Virginia to easternTexas. It also occurred on sites in thePiedmont, southern Ridge and Valley,and southern Blue Ridge Provinces(fig. 1.22). This community oncecovered over 40 percent of the entireregion, but it has declined by more than98 percent (Noss and others 1995).

The community came under pressureduring the mid-17th century. Demandbegan for naval stores and then turned

Figure 1.22—The historic and present distribution of longleaf pinein the South (White and others 1998).


TERRESTRIAL Table 1.11—Examples of soft and hard mast-producing species in the South

Scientific name Common name Scientific name Common name

Soft mastPomes

Amelanchier spp. ServiceberriesCrataegus spp. HawthornPyrus malus Common apple

DrupesBerchemia scandens Alabama supplejackCallicarpa Americana American beautyberryCeltis occidentalis HackberryCornus florida Flowering dogwoodGaylussacia spp. HuckleberriesGaylussacia dumosa Dwarf huckleberryIlex spp. HolliesIlex cassine DahoonIlex coriacea Large gallberryIlex deciduas PossumhawIlex glabra GallberryIlex myrtifolia Myrtle dahoonIlex opaca American hollyIlex vomitoria YauponMorus rubra Red mulberryMyrica cerifera Southern bayberryMyrcia pensylvanica Northern bayberryNyssa aquatica Water tupeloNyssa sylvatica Black tupelo and

Swamp tupeloPersea borbonia RedbayPrunus serotina Black cherryPrunus spp. Wild cherries and

plumsRhus copallina Shining sumacRhus glabra Smooth sumacRhus radicans Common poison ivyRhus typhina Staghorn sumacRubus spp. BlackberriesSabal spp. PalmettoSambucus canadensis American elderSassafras albidum SassafrasSerenoa repens Saw-palmettoViburnum spp. Viburnum

BerriesDiospyros virginiana Common persimmonJuniperus virginiana Eastern redcedarLonicera japonica Japanese honeysuckleSmilax spp. GreenbriersVaccinium spp. BlueberriesVitis aestivalis Muscadine grapeVitis rotundifolia Summer grape

Hard mastNuts

Aesculus octandra Yellow buckeyeCarpinus caroliniana American hornbeamCarya spp. HickoriesCarya aquatica Water hickoryCarya cordiformis BitternutCarya glabra PignutCarya ovata ShagbarkCarya tomentosa MockernutCastanea spp. ChinkapinFagus grandifolia American beechJuglans cinera Butternut

(white walnut)Juglans nigra Black walnutOstrya virginiana Eastern hophornbeamNyssa sylvatica Black gumQuercus spp. OaksQuercus alba White oakQuercus chapmanii Chapman oakQuercus michauxii Swamp chestnut oakQuercus prinus Chestnut oakQuercus stellata Post oakQuercus virginiana Live oakQuercus falcate Southern red oakQuercus ilicifolia Bear oakQuercus incana Bluejack oakQuercus laurifolia Laurel oakQuercus marilandica Blackjack oakQuercus nigra Water oakQuercus nuttalli Nuttall oakQuercus phellos Willow oakQuercus pumila Running oakQuercus rubra Northern red oak

Source: Halls 1977.

to timber needs. By the 1960s,extensive areas were harvested andconverted to commercial plantationsof loblolly and slash pine. Firesuppression and the introduction oflivestock further restricted the longleafcommunity to a few isolated locationscomprising about 5 million acres.At present, the majority occurs on

private land. Much of what remainsis largely degraded due to lack ofproper management.

Community composition varies withsoil moisture and geography. Wiregrassand bluestem dominate the herbaceouslayer. This herb layer is diverse andincludes grasses, wildflowers, andcarnivorous plants. In mature

communities, the trees are thinlydistributed and flat-topped, andhave limbless lower trunks.

The community harbors severalvertebrate species. The fox squirrelis a long-lived species with lowreproductive rates. It depends onlongleaf pine for late summer forage.The decline in longleaf communities


has limited its range and reducedpopulation levels. The red-cockadedwoodpecker occurs in the openpinewoods, using fairly mature treeswith minimal understory (Hamel1992). Trees also must have properheartwood conditions for nest cavities.This species has also declined, butactive management has stabilizedseveral populations. The sensitiveBachman’s sparrow breeds in dense,grassy places where scattered pinetrees and saplings are present.

Dodd (1995) reported that 74amphibians and 96 reptiles occurin the range of the longleaf pinecommunity. These include theflatwoods salamander, Red Hillssalamander, striped newt, Carolinagopher frog, eastern indigo snake,gopher tortoise, eastern diamondbackrattlesnake, Florida pine snake,and Florida scrub lizard.

Although the influence of longleafreduction on the herpetofaunalcommunity has not been assesseddirectly, several species may havebeen affected. The gopher tortoise,a keystone species in longleaf pinesavanna, has declined by 80 percentover the last century (White andothers 1998). Amphibians breedingin temporary ponds have beenparticularly affected by habitatalteration. The flatwoods salamanderhas disappeared from its eastern range;gopher frogs are nearly extirpatedin North Carolina, Alabama, andMississippi; and dusky salamandersappear to have declined in coastalSouth Carolina and peninsular Florida.

Conversion of longleaf pine foreststo agriculture, slash, or loblolly pineplantations and urban developmentthreaten the continued existence ofseveral herpetofauna species in Georgiaand Florida (Ware and others 1993).Hardwood encroachment stemmingfrom fire suppression also has contri-buted to the loss of longleaf pinecommunities. Historically, frequentlow-intensity fires reduced litteraccumulation, controlled competingwoody species, and improvedherbaceous vigor (Walker 2001).Recent awareness of the importanceof this sensitive community hasencouraged restoration efforts.

Profiles of selected rarecommunities: Atlantic white-cedarswamps—Atlantic white-cedar oncewas distributed from southern Virginia

to interior Georgia and from the FloridaPanhandle along the Gulf of Mexicoto Mississippi. Drainage, development,and harvest without regenerationhave reduced Atlantic white-cedarto 10 percent of its original extent.

Much of the original communitywas destroyed by European settlerswho cleared land for agriculture.Today, white-cedar swamps arerestricted to inaccessible freshwaterwetlands in small, isolated stands.Road construction and the dammingof waterways continue to diminishthis habitat, as does suburban en-croachment, industrial runoff,and pollution.

Atlantic white-cedar swamps areunique communities adapted tovariable hydrological regimes, fire,and peat soils. This community typeoften represents some of the only forestin regions of intense agricultural andurban development. Atlantic white-cedar areas provide habitat for manyspecies, including black bear, deer,rabbits, and other fauna. The diversityof bird species is relatively high inAtlantic white-cedar swamps, com-pared to adjacent areas. The Hessel’shairstreak is a butterfly that feedsexclusively on Atlantic white-cedar.

During restoration, these standsrequire frequent, light fires in thedry season. Fire removes competitivevegetation and clears the seedbedfor regeneration.

Hard and Soft MastSouthern species that produce

mast—Mast refers to specific kindsof fruits of woody species. Hardmast possesses a hard exterior, asin acorns, while soft mast has fleshyfruits, as in berries. Both forms of mastare important in the diets of southernwildlife. Many southern woody plantsproduce mast (table 1.11). Mast yieldsare unpredictable from one year to thenext and vary according to species,location, and weather.

Pomes are fruits that have severaltough, papery-walled cavities thathouse seed; the cavities are surroundedby thick flesh. These fruits may be largelike apples or small like serviceberries.Fresh pomes have a high moistureand carbohydrate content, but arelow in crude protein (Halls 1977).

A drupe is a pulpy fruit with an innerovary wall that encloses a seed. Drupes

are extensively eaten by wildlife. Thefruits tend to be low in crude proteinand high in carbohydrates; nutrientcontent varies considerably amongspecies. Drupe producers in the Southinclude wild cherries, plums, hackberry,and red mulberry (Halls 1977).

Berries are fruits with fleshy ovariesthat envelop one or more seeds. Mostspecies are eaten by wildlife. Fruits areusually high in carbohydrates and lowin crude protein. Species that produceberries include persimmon, blueberry,and grape.

Hard mast includes nuts and one-seeded fruits (or kernels). Most haveconcentrations of crude fat, and somealso are relatively high in crude protein(Halls 1977). Characteristic speciesinclude hornbeam, hickory, beech,walnut, black gum, and severalspecies of oaks.

Selected species that utilize mastin their diet—Mast is an essentialcomponent in the diets of manyvertebrates in the South (Combsand Frederickson 1996, Doherty andothers 1996, Jensen 1982, Wolff 1996).Table1.12 lists several mast-consumingmammals, including mice, voles,woodrats, rabbits, raccoons, and foxes.Several birds also consume mast(table 1.13) including game birds(doves, quail, pheasant, grouse, turkey),waterfowl (mallards, wood ducks),woodpeckers, and songbirds (finches,thrushes, jays, and towhees). Therelationship between mast and thefood habits of several game species,such as deer, bear, and squirrelshas been documented extensively(Fridell and Litvaitis 1991, Kirkpatrick1989, Kurzejeski 1989, Pelton 1989,Wentworth and others 1989).

Selected species that utilize mastin their diet: white-tailed deer—Hard mast is often an importantcomponent of the fall and winterdiet of white-tailed deer. Nutrition,reproduction, weight, and antlercharacteristics of individual animalsare influenced by acorn availability(Wentworth and others 1989). Inpoor mast years, reproduction ratesmay be low, and conception maybe delayed. Postnatal survival alsocan decline following years of minimalacorn production. Fawn weight alsocan be directly related to the sizeof the acorn crop.


TERRESTRIAL

Table 1.12—Selected mammals of the South that utilize hardand soft mast in their diets

Scientific name Common name

Castor canadensis BeaverClethrionomys gapperi Southern red-backed voleDidelphis virginiana Virginia opossumGlaucomys sabrinus Northern flying squirrelGlaucomys volans Southern flying squirrelMephitis mephitis Striped skunkNeotoma floridana Eastern woodratNeotoma mexicana Mexican woodratNeotoma micropus Southern plains woodratOchrotomys nuttalli Golden mouseOdocoileus virginianus White-tailed deerPeromyscus attwateri Texas mousePeromyscus boylii Brush mousePeromyscus floridanus Florida mousePeromyscus gossypinus Cotton mousePeromyscus leucopus White-footed mousePeromyscus maniculatus Deer mouseProcyon lotor RaccoonSciurus carolinensis Gray squirrelSciurus niger Fox squirrelSpermophilus variegatus Rock squirrelSus scrofa Wild boarSylvilagus palustris Marsh rabbitTamiasciurus hudsonicus Red squirrelTamias striatus Eastern chipmunkUrocyon cinereoargenteus Gray foxUrsus americanus Black bearVulpes vulpes Red fox

AlabamaArkansas

FloridaGeorgia

KentuckyLouisiana

MississippiNorth Carolina

E. OklahomaSouth Carolina

TennesseeE. TexasVirginia

South

State timberland area (percent)0 2 4 6 8 10 12 14

23

5

7

1312

34

355

44

56

6

444

42

53

56

7

7

9

Percent other public Percent National Forest

Selected species that utilizemast in their diet: black bear—The abundance and distributionof oak mast (particularly white oak)also can influence black bear natality,mortality, and dispersal. Shifts in homerange sometimes occur in response tofluctuations in hard mast availability.The birth and survival of young bearscan be directly associated with oakmast crops (Pelton 1989). Poor mastyears often result in increased bearmovement, which can result inincreased mortality due to vehicularaccidents and human-bear interactions.The loss of the American chestnutlikely had a significant influenceon the population dynamics of blackbears in the Southern Appalachians(Pelton 1989). In addition, the relianceon soft mast in the seasonal diet ofblack bear highlights the importanceof early successional habitats in theprovision of this food source (Traniand others 2001).

Selected species that utilize mast intheir diet: squirrels—The availabilityof hard mast also can influence squirrelpopulations. Poor mast crops can resultin population declines, while abundantmast crops may result in substantialpopulation increases (Kurzejeski 1989).Mast comprises the majority of the fall,winter, and spring diets of red, gray,and fox squirrels. Acorns, walnuts,and hickory nuts are major foodsources for these squirrels as wellas for the eastern chipmunk.

Selected species that utilize mastin their diet: game birds—Hard mastprovides a high-energy resource forruffed grouse, wild turkey, bobwhitequail, and several waterfowl. Thesespecies consume acorns in proportionto their availability throughout the year;foraging for mast requires little energyexpenditure (Kirkpatrick 1989). Redoak acorns have an elevated phenoliccontent and are less palatable thanwhite oak species.

Factors affecting mast supplyavailability—In recent years, therehave been concerns about the declineof mast-producing species (particularlyoaks) in the South. Chapter 16 presentstrend information from the FIA on oakand other overstory mast-producingtrees. In addition, an examination ofoak decline in the South is presentedin chapter 18. The factors that mayhave contributed to the decline, and

Figure 1.23—National forest and other public ownership of timberlandin the South (U.S. Department of Agriculture, Forest Service 2000a).


Table 1.13—Selected birds of the South that utilize hard andsoft mast in their diets

Scientific name Common name

Aix sponsa Wood duckAnas platyrhynchos MallardAnas strepera GadwellAphelocoma coerulescens Scrub jayBombycilla cedrorum Cedar waxwingCarpodacus purpureus Purple finchCatharus guttatus Hermit thrushCerthia americana Brown creeperColaptes auratus Northern flickerColinus virginianus Bobwhite quailColumba fasciata Band-tailed pigeonColumba flavirostris Red-billed pigeonCorvus brachyrhynchos American crowCyanocitta cristata Blue jayCyanocitta stelleri Stellar’s jayIxoreus naevius Varied thrushMelanerpes carolinus Red-bellied woodpeckerMelanerpes erythrocephalus Red-headed woodpeckerMelanerpes formicivorus Acorn woodpeckerMeleagris gallopavo Wild turkeyMimus polyglottos Northern mockingbirdParus bicolor Tufted titmouseParus inornatus Plain titmousePhasianus colchicus Ring-necked pheasantPheucticus ludovicianus Rose-breasted grosbeakPhilohela minor American woodcockPicoides pubescens Downy woodpeckerPicoides villosus Hairy woodpeckerPipilo erythrophthalmus Rufous-sided towheeQuiscalus quiscula Common grackleSitta carolinensis White-breasted nuthatchSphyrapicus varius Yellow-bellied sapsuckerSturnus vulgaris StarlingToxostoma rufum Brown thrasherTympanuchus cupido Greater prairie chickenZenaidia macrocroura Mourning dove

Contribution ofPublic Lands

Extent of public lands in theSouth—Public land comprisesapproximately 11 percent of timber-land in the South (chapter 16). Thedistribution of public land betweenStates varies considerably (fig. 1.23).For example, national forestsoccupy 3 percent of the timberlandin Alabama and Georgia but 13percent of the timberland in Arkansas(U.S. Department of Agriculture,Forest Service 2000a).

FIA data indicate that 4 million acresof timberland are managed by States,1 million acres by counties andmunicipalities, and 16 million acresby Federal agencies (U.S. Departmentof Agriculture, Forest Service 2000a).State land is contained in State parks,wildlife management areas, State for-ests, and State natural resource areas.Counties and municipalities hold landin local parks and recreation areas,many of which contribute importantlyto the conservation of habitat.

The primary Federal landmanagement agencies in the South arethe USDA Forest Service, the NationalPark Service, and the U.S. Fish andWildlife Service (fig. 1.24). Federal landis concentrated in the Appalachian andOzark Mountains, with less land in thePiedmont and Coastal Plain. The ForestService manages approximately 60percent of the southern Blue Ridge,the eastern edge of the AppalachianMountain chain. In contrast, less thanone-tenth of the mid-Atlantic CoastalPlain is under Federal management.

National parks and the NationalPark Service—The idea of preservingFederal land in national parks is rootedin the conservation movement of thelate 1800s. Created in 1916, themission of National Park Service wasto conserve scenic, natural, and historicresources (Loomis 1993). Congressprecluded timber harvesting, mining,and livestock grazing.

In the 1960s, the Leopold Reportshifted this preservation philosophytowards ecological management(Loomis 1993). Parks were managedto restore a more natural appearance,and visitor development was directedto areas outside the parks. Park policiesallowed fire as a management tool formaintaining the park environment.Recreational activities were limited

the subsequent reduction in hard mastproduction, are briefly mentioned here.

Many variables, including disease,insect infestation, advanced standage, drought, and disturbance influenceoak forests. Mature oaks are quitesusceptible to disease and droughtconditions. As these forests age,tree vigor is reduced. They becomesusceptible to windthrow and icestorms. Longevity varies by speciesand site characteristics. Lack of naturaldisturbance is another factor. Firesuppression has resulted in an increasein other species in former oak-dominated areas.

Chestnut blight had a dramaticinfluence on the American chestnut(chapter 18). Chestnut oaks, whichreplaced chestnuts in many places,are an important source of hard mastfor wildlife populations. Gypsy mothinfestations on the poor sites occupiedby chestnut oaks often inhibit oakregeneration. Infested trees havea reduced capability for stumpsprouting, and their acorns lackthe energy reserves to remain viable.Repeated defoliation kills many oaks.When this happens, yellow-poplaroften captures the site.


TERRESTRIAL

National Forest System

National Wildlife RefugesNational Parks

based upon soil and vegetationcharacteristics, concerns about waterquality, and sensitivity of wildlifeto human presence. Still, on NationalPark Service land there is ongoingconflict between preserving the naturalenvironment and providing forvisitor use.

The Agency’s current mandate is toperpetuate native plant communities;manipulation of vegetation is keptto a minimum. Species managementobjectives include the provision ofself-regulating populations. Impactson animal populations are avoidedwith restrictions on the removalof individual animals.

In 2000, the National ParkService managed 97 properties inthe South totaling over 5 million

acres (table 1.14). These propertiesare in seven different designations,each of which is managed with differ-ent objectives. National parks containoutstanding natural features andgenerally are of a sufficient sizeto ensure protection from outsideinfluences. National preserves alsoprotect selected natural features, butallow uses such as hunting or miningif they do not impair the resourcesof the preserve. National seashoresprotect water-related areas of naturalsignificance that occur on the Atlanticand Gulf coasts. National recreationareas emphasize recreational use.Recreational areas also may exist onnational forests. National parkwaysprotect scenic resources along travelcorridors such as the Blue RidgeParkway. National monuments

and national historic sites (includingnational battlefields) are establishedto commemorate historical events(Loomis 1993).

The following area accounts describeselected National Park Service proper-ties that provide valuable habitat fora variety of species in the South. Manyareas contain impressive vertebratediversity or provide examples ofapplied conservation biology. Propertyinformation is summarized fromU.S. Department of the Interior,Park Service (2000).

National parks and the NationalPark Service: Buffalo National River,AR—The Buffalo River is one of thefew remaining unpolluted, free-flowingrivers in the South. Stretching 135miles, the Buffalo River cuts its way

Figure 1.24—The distribution of national forests, national parks,and national willife refuges in the South (White and others 1998).


Table 1.14—National Park Service national parks and monuments in the South

National Park Service property Total acres

AlabamaNational Parks

Horseshoe Bend National Military Park 2,040Little River Canyon National Preserve 13,633Tuskegee Airman National Historic Site (Private) 87Tuskegee Institute National Historic Site 58

National MonumentsRussell Cave National Monument 310

Total 16,128

ArkansasNational Parks

Arkansas Post National Memorial 749Buffalo National River 94,328Fort Smith National Historic Site 75Hot Springs National Park 5,549Little Rock Central HS National Historic Site 18Pea Ridge National Military Park 4,300

Total 105,019

FloridaNational Parks

Big Cypress National Preserve 720,573Biscayne National Park 172,924Canaveral National Seashore 57,662De Soto National Memorial 27Dry Tortugas National Park 64,700Everglades National Park 1,508,607Gulf Islands National Seashore 135,607Timucuan Ecological and Historic Preserve 46,000

National MonumentsCastillo de San Marcos National Monument 21Fort Caroline National Memorial 138Fort Matanzas National Monument 228

Total 2,706,487

GeorgiaNational Parks

Andersonville National Historic Site 495Chattahoochee River National Recreation Area 9,206Chickamouga and Chattanooga National Military Park 8,119Cumberland Island National Seashore 36,415Jimmy Carter National Historic Site 71Kennesaw Mountain National Battlefield Park 2,884Martin Luther King, Jr. National Historic Site 34

National MonumentsFort Frederica National Monument 241Fort Pulaski National Monument 5,623Ocmulgee National Monument 702

Total 63,790

continued

through massive limestone bluffs inthe Ozark Mountains. The BuffaloNational River has three designatedwilderness areas within its boundaries.

Ninety-five thousand acres furnishhabitat for 250 species of birds anda variety of animals. It also contains70 mines that provide important habitatfor gray, Indiana, and Ozark big-earedbats. The Buffalo National River alsois along the migration route of thefederally listed Eskimo curlew.

National parks and the NationalPark Service: Mammoth CaveNational Park, KY—This park wasestablished in 1941 to preserve oneof the longest known cave systems(336 miles) in the Nation. The parkalso was designated as a World HeritageSite in 1981 and an InternationalBiosphere Reserve in 1990.

The park’s 52,830 acres support avariety of plants and animals includingseveral bat species of conservationconcern: southeastern bat, Rafinesque’sbig-eared bat, and eastern small-footedbat. There are several State-listedreptiles, including the northern coalskink, glass lizard, and the northernpine snake. Among the 872 floweringspecies that have been confirmed are21 listed plants.

National parks and the NationalPark Service: Congaree SwampNational Monument, SC—Thismonument was established to protectthe largest remaining tract of virginbottomland hardwood wetlands inthe South. The monument is an inter-national biosphere reserve, a nationalnatural landmark, a wilderness area,and a continentally important bird area.

Biodiversity is very high within theCongaree’s 22,000 acres. Amphibiansthat thrive in the deep floodplainsloughs include the marbled sala-mander, the eastern newt, the southerndusky salamander, and the greatersiren. Frogs include the southernleopard frog and the chorus frog.One hundred seventy-three speciesof birds occur in the monument,including several of conservationconcern. Among these are the barredowl, pileated woodpecker, andSwainson’s warbler. At different seasonsof the year, prothonotory warblers,Mississippi kites, and herons use therefuge. In addition, Congaree Swampsupports important sites for the silver-haired bat, hoary bat, Brazilian


TERRESTRIAL

Table 1.14—National Park Service national parks and monuments in theSouth (continued)


KentuckyNational Parks

Abraham Lincoln Birthplace National Historic Site 337Cumberland Gap National Historic Park 20,454Mammoth Cave National Park 52,830

Total 73,621

LouisianaNational Parks

Cane River Creole National Historic Park 207Jean Lafitte National Historic Park and Preserve 20,020New Orleans Jazz National Historic Park 4

National MonumentsPoverty Point National Monument 911

Total 21,142

MississippiNational Parks

Brices Cross Roads National Battlefield Site 1Gulf Islands National Seashore 135,458Natchez National Historic Park 108Natchez Trace National Scenic Trail 10,995Natchez Trace Parkway 51,747Tupelo National Battlefield 1Vicksburg National Military Park 1,736

Total 200,046

North CarolinaNational Parks

Blue Ridge Parkwaya 88,734Cape Hatteras National Seashore 30,319Cape Lookout National Seashore 28,243Carl Sandburg Home National Historic Site 264Fort Raleigh National Historic Site 513Guilford Courthouse National Military Park 220Moores Creek National Battlefield 88Wright Brothers National Memorial 428

Total 148,809

OklahomaNational Parks

Chickasaw National Recreation Area 9,889Oklahoma City National Memorial 6Washita Battlefield National Historic Site 315

Total 10,210

South CarolinaNational Parks

Charles Pinckney National Historic Site 28Cowpens National Battlefield 842Kings Mountain National Miliary Park 3,945Ninety Six National Historic Site 989

continueda Property is in two or more States.

free-tailed bat, Rafinesque’s big-eared bat, and southeastern bat.

Feral hogs in the park are placingthis unique resource at risk. Wetlandcommunities are subject to severedamage from hog rooting andother behavior.

National parks and the NationalPark Service: Great SmokyMountains National Park, NC, TN—The Great Smoky Mountains NationalPark is one of the largest protectedareas in the South (521,621 acres) andis World-renowned for the diversityof its plant and animal resources andthe integrity of the wilderness withinits boundaries. Established as a nationalpark in 1934, it was designated as anInternational Biosphere Reserve in 1976and a World Heritage Site in 1983.

The park protects some of the World’sfinest temperate deciduous forests.Due to the fertile soil and abundantrain, this area boasts 1,650 species offlowers and trees, 50 mammal species,and 27 salamander species. Migratingbirds abound in late spring.

Existing and impending threats in thepark include invasion by exotic species,air pollution, and forest diseases. Sincefire suppression was initiated in the1930s, oak regeneration has beenminimal at some sites with adverseconsequences for mast-utilizing species.

National parks and the NationalPark Service: Big Thicket NationalPreserve, TX—Big Thicket was thefirst preserve in the National ParkSystem to protect an area of richbiological diversity. Established in1974, it also was designated as anInternational Biosphere Reserve. Thepreserve consists of nine land unitsand six water corridors encompassingmore than 97,191 acres. The BigThicket is rich in biological resourcesand contains swamps, bayous, pinesavanna, sandhills, plains, and desert.

National parks and the NationalPark Service: Shenandoah NationalPark, VA—This park extends along theBlue Ridge Mountains, encompassingover 198,000 acres. The oak-hickoryforest is inhabited by deer, black bear,bobcat, and wild turkey. Species suchas the chipmunk, groundhog, raccoon,skunk, opossum, and gray squirrel arefrequently detected. Approximately 200species of birds have been recorded,including flycatchers, thrushes, vireos,35 species of warblers, and migrating


Table 1.14—National Park Service national Parks and monuments in theSouth (continued)


South Carolina (cont.)National Monuments

Congaree Swamp National Monument 21,867Fort Sumter National Monument 195

Total 27,866

TennesseeNational Parks

Andrew Johnson National Historic Site 17Big South Fork National River and Recreation Area 125,242Fort Donelson National Battlefield 552Great Smoky Mountains National Parka 521,621Obed Wild and Scenic River 5,173Shiloh National Military Park 3,997Stones River National Battlefield 708

Total 657,310

TexasNational Parks

Amistad National Recreation Area 58,500Big Bend National Park 801,163Big Thicket National Preserve 97,191Chamizal National Memorial 55Fort Davis National Historic Site 474Guadalupe Mountains National Park 86,416Lake Meredith National Recreation Area 44,978Lyndon B. Johnson National Historic Park 1,570Padre Island National Seashore 130,434Palo Alto Battlefield National Historic Site 3,357Rio Grande Wild and Scenic River 9,600San Antonio Missions National Historic Park 819

National MonumentsAlibates Flint Quarries National Monument 1,371

Total 1,235,928

VirginiaNational Parks

Appomattox Court House National Historic Park 1,775Arlington House, The Robert E. Lee Memorial 28Colonial National Historic Park 9,349Fredericksburg National Military Park 7,787George Washington Memorial Parkway 7,248Maggie L. Walker National Historic Site 1Petersburg National Battlefield 2,659Manassas National Battlefield Park 5,212Prince William Forest Park 18,661Richmond National Battlefield Park 1,078Shenandoah National Park 198,182Wolf Trap Farm Park for the Performing Arts 130

National MonumentBooker T. Washington National Monument 224George Washington Birthplace National Monument 550

Total 252,884Grand total 5,519,240

a Property is in two or more States.Source: U.S. Department of Interior 2000a.

hawks. Permanent residents includeruffed grouse, barred owl, raven,woodpeckers, and junco. The park alsosupports several salamander speciesand two poisonous snakes, the timberrattlesnake and the copperhead snake.

The hemlock woolly adelgid, anexotic insect, currently jeopardizesthe eastern hemlocks in the park. Firstdetected 10 years ago, the adelgid isan aphid-like insect that sucks sap frombranches of the hemlock. The tree losesstrength and sheds its needles, andoften does not survive (chapter 17).

National parks and the NationalPark Service: Blue Ridge Parkway,NC, VA—The Blue Ridge Parkwayconsists of 469 miles of road andprotects the natural features of theBlue Ridge while connecting theShenandoah National Park with theGreat Smoky Mountains. The parkwayencompasses 88,734 acres.

The parkway supports several speciesof rare plants and animals. Someof these, such as the Peaks of Ottersalamander and the Blue Ridgegoldenrod, do not occur in othersouthern areas. Ponds and wetlandsnear the parkway provide essentialhabitat for amphibians, reptiles,mammals, and birds.

Many Neotropical migrant speciesreturn to the parkway each spring.These include the scarlet tanager, veery,wood thrush, and Kentucky warbler.The autumn hawk migration alsooccurs along the Blue Ridge Parkway.Raptors recorded include the Americankestrel, red-tailed hawk, sharp-shinnedhawk, broad-winged hawk, goldeneagle, and peregrine falcon.

National wildlife refuges andthe Fish and Wildlife Service—A network of lands set aside for wildlifebegan in 1903 with the designationof Pelican Island, FL, as the firstNational Wildlife Refuge. The Fishand Wildlife Service has responsibilityfor the Refuge System. Refugeobjectives include the provision andenhancement of habitat, perpetuationof migratory bird resources, preserva-tion of natural diversity, and restorationof endangered and threatened species.

Land is acquired for game refuges,waterfowl production areas, and otherreasons. Many refuges were createdunder the authority of the EndangeredSpecies Act, providing anchors forbiodiversity and ecosystem-level


TERRESTRIAL

Table 1.15—U.S. Fish and Wildlife Service refuges within the South

TotalRefuge acres

AlabamaBlowing Wind Cave 264Bon Secur 6,678Choctaw 4,218Eufaula 7,953Fern Cave 199Grand Bay 2,496Key Cave 1,060Watercress Darter 9Wheeler 34,247FSA Interest ALa 743

Total 57,867

ArkansasBald Knob 14,760Big Lake 11,036Cache River 45,232Felsenthal 64,902Holla Bend 6,428Logan Cave 124Overflow 12,235Pond Creek 26,816Wapanocca 5,484White River 154,856FSA Interest ARa 3,459

Total 345,332

FloridaArchie Carr 127Arthur R. Marshall 145,787Caloosahatchee 40Cedar Keys 891Chassahowitzka 30,843Crocodile Lake 6,688Crystal River 80Egmont Key 328Florida Panther 26,529Great White Heron 192,584Hobe Sound 980Island Bay 20J.N. Ding Darling 6,315Key West 208,308Lake Wales Ridge 1,814Lake Woodruff 21,559Lower Suwannee 51,031Matlacha Pass 393Merritt Island 139,174National Key Deer 8,614Okefenokee 3,678Passage Key 64Pelican Island 4,824Pine Island 602Pinellas 394St. Johns 6,256St. Marks 67,122

TotalRefuge acres

Florida (cont.)St. Vincent 12,490Ten Thousand Islands 35,034FSA Interest FLa 3,124

Total 975,693

GeorgiaBanks Lake 3,559Blackbeard Island 5,618Bond Swamp 5,490Eufaula 3,231Harris Neck 2,762Okefenokee 391,402Piedmont 34,967Savannah 12,011Wassaw 10,070Wolf Island 5,126FSA Interest GAa 4,778

Total 479,014

KentuckyClarks River 5,017Ohio River Islands 410Reelfoot 2,040

Total 7,467

LouisianaAtchafalaya 15,255Bayou Cocodrie 13,169Bayou Sauvage 22,261Big Branch Marsh 12,642Black Bayou Lake 1,861Bogue Chitto 29,493Breton 9,047Cameron Prairie 9,621Catahoula 6,545D’Arbonne 17,420Delta 48,799Grande Cote 6,077Handy Brake 466Lacassine 34,379Lake Ophelia 17,306Mandalay 4,619Sabine 140,717Shell Keys 8Tensas River 65,746Upper Quachita 41,063FSA Interest LAa 14,026

Total 510,520

MississippiBogue Chitto 6,808Dahomey 9,167Grand Bay 5,120Hillside 18,678Mathews Brake 2,419

continued

conservation. These areas have beeninstrumental in the recovery of severalspecies including the whooping crane,Key deer, and American crocodile.

The Migratory Bird ConservationAct of 1929 directed the Agency topurchase areas as refuges for migratorybirds. In 1934, the Duck Stampprogram established permanent fundsfor the acquisition of waterfowlhabitats. The system has an outstandingrecord for the successful managementof these species. The emphasis onmigratory birds has now expandedto include colonial water birds,birds of prey, shorebirds, seabirds,and songbirds.

The earliest form of managementconsisted of law enforcement andperiodic counts of wildlife. As thesystem expanded, there was an evolu-tion from habitat management for afew species to ecosystem management.For example, planting vegetation forducks evolved to planting an arrayof native grasses and forbs to rebuildprairie diversity. Prescribed fire wasincorporated to reduce hazardousfuel loads and restore vegetationcommunities. Management has beenaltered to mimic natural disturbance formaintenance of a diversity of habitats.

One hundred seventy-two refugesspread across the South encompassapproximately 4 million acres (table1.15). The greatest concentrationof wildlife refuges is in Florida andalong the Mississippi and Atlanticflyways. Hundreds of species of birds,mammals, reptiles, and amphibiansare supported by the diversity ofhabitats in the Wildlife Refuge System.Several of these properties are discussedin greater detail in the followingsection. Information on species andcommunities are summarized fromU.S. Department of the Interior,Fish and Wildlife Service (2000).

National wildlife refuges and theFish and Wildlife Service: FloridaPanther National Wildlife Refuge—This refuge supports a variety ofhabitats, including cypress forests,swamps, pine forests, hardwoodhammocks, prairies, marshes, andsloughs. Permanent and seasonalwetlands cover a majority of therefuge area (26,529 acres). The refugeis closed to the public to minimizedisturbance to the Florida pantherpopulation that occurs there.


Table 1.15—U.S. Fish and Wildlife Service refuges within theSouth (continued)

TotalRefuge acres

Mississippi (cont.)Mississippi Sandhill Crane 19,713Morgan Brake 7,372Noxubee 46,914Panther Swamp 35,272St. Catherine Creek 24,931Tallahatchie 4,839Yazoo 12,940FSA Interest MSa 29,326

Total 223,499

North CarolinaAlligator River 156,125Cedar Island 14,482Currituck 4,317Great Dismal Swamp 24,812Mackay Island 7,150Mattamuskeet 50,180Pea Island 5,834Pee Dee 8,439Pocosin Lakes 108,692Roanoke River 17,977Swanquarter 16,411FSA Interest NCa 6,175

Total 420,594

OklahomaDeep Fork 8,387Little River 12,029Optima 4,333Ozark Plateau 2,858Salt Plains 32,057Sequoyah 20,800Tishomingo 16,464Washita 8,075Wichita Mountains 59,020

Total 164,023

Puerto RicoCabo Rojo 1,857Culebra 1,574Desecheo 360Laguna Cartagena 1,036

Total 4,827

South CarolinaACE Basin 11,772Cape Romain 65,225Carolina Sandhills 45,348Pinckney Island 4,053Santee 12,483Savannah 14,839Tybee 100Waccamaw 4,978FSA Interest SCa 1,430

Total 160,228

a Farm Service Agency.Source: U.S. Department of the Interior 2000b.

TotalRefuge acres

TennesseeChickasaw 22,376Cross Creeks 8,861Hathcie 11,556Lake Isom 1,846Lower Hatchie 9,353Reelfoot 8,409Tennessee 51,359FSA Interest TNa 685

Total 114,445

TexasAnahuac 34,296Aransas 114,397Attwater Prairie Chicken 9,199Balcones Canyonlands 16,481Big Boggy 4,526Brazoria 43,905Buffalo Lake 7,664Grulla 5Hagerman 11,320Laguna Atascosa 57,826Little Sandy 3,802Lower Rio Grande Valley 77,695McFaddin 56,181Moody 3,517Muleshoe 5,809San Bernard 30,267Santa Ana 2,088Texas Point 8,952Trinity Point 6,801FSA Interest TXa 1,718

Total 496,449

VirginiaBack Bay 8,315Chincoteague 13,598Eastern Shore 1,570Featherstone 326Fisherman Island 1,025Great Dismal Swamp 83,944James River 4,195Mackay Island 874Martin 146Mason Neck 2,276Nansemond 423Occoquan Bay 642Plum Tree Island 3,502Presquile 1,329Rappahannock River 2,975Wallops Island 3,373FSA Interest VAa 134

Total 128,647

Virgin IslandsBuck Island 45Green Cay 14Sandy Point 490

Total 549

Grand total 4,089,154

There are several listed species onthe refuge. Mammals include theFlorida panther and Florida black bear.Avian species include the wood stork,snail kite, bald eagle, and Floridagrasshopper sparrow. The Americanalligator, eastern indigo snake, stripedmud turtle, and loggerhead sea turtleare reptiles of conservation concern.

Habitat management objectives centeron the provision of optimum conditionsfor the panther. Other objectivesinclude restoration of natural diversityand implementation of environmentaleducation programs promoting Floridapanther and south Florida ecosystems.

National wildlife refuges and theFish and Wildlife Service: St. VincentNational Wildlife Refuge, FL—This12,490-acre island refuge is a red wolfpropagation site. Additional endangeredand threatened species that occur on St.Vincent Island include the bald eagle,piping plover, wood stork, easternindigo snake, and loggerhead sea turtle.

The primary refuge objective ismanagement and preservation of thenatural barrier island and associatednative plant and animal communities.Additional management objectivesinclude the provision of habitatfor migratory birds and protectionof listed species.

National wildlife refuges andthe Fish and Wildlife Service:Okefenokee National WildlifeRefuge, GA—Established in 1936,the Okefenokee Refuge covers 391,402acres. The swamp contains numerousislands and lakes, along with vast areasof nonforested terrain. Prairies coverapproximately 60,000 acres of theswamp. Once forested, these marshexpanses were created during periodsof severe drought when fires burnedvegetation and surface layers of peat.

A wide variety of bird species aresupported. The prairies harbor wadingbirds, including herons, egrets, whiteibis, sandhill cranes, wood storks, andbitterns. Scrub-shrub areas supportvarious warblers.

Refuge objectives encompassprotection of the unique environmentalqualities of the Okefenokee ecosystem,and the provision of optimum habitatfor a wide diversity of fish, birds,mammals, reptiles, and amphibians.


TERRESTRIAL

National wildlife refuges and theFish and Wildlife Service: TensasRiver National Wildlife Refuge, LA—This refuge lies in the upper basinof the Tensas River in northeasternLouisiana. It includes the site of thelast documented sighting of the ivory-billed woodpecker. The refuge supports65,746 acres of woodlands, croplands,reforested agricultural fields, and openwater. The area also is home to thethreatened Louisiana black bear.

Management objectives include watermanagement for waterfowl, wadingbirds, and shorebirds. Cooperativefarming provides habitat for migratorybirds and bear. Deer are managedvia public hunting.

National wildlife refuges and theFish and Wildlife Service: AlligatorRiver National Wildlife Refuge,NC—This 156,125-acre refuge wasestablished to preserve a uniquewetland habitat type, the pocosin,and its associated terrestrial species.Diversity of habitat types includesbogs, freshwater and brackish marshes,hardwood swamps, and Atlantic white-cedar swamps. Plant species includepitcher plants, sun dews, low-bushcranberries, bays, pond pine, redmaple, and a wide variety of herbaceousand shrub species common tothe South.

Refuge objectives center on thepreservation of the unique wetlandand the provision of habitat for thered wolf, red-cockaded woodpecker,American alligator, black bear,waterfowl, and migratory birds.

National wildlife refuges andthe Fish and Wildlife Service:Mississippi Sandhill Crane NationalWildlife Refuge, MS—This refugeoccupies 19,713 acres of pine-savannahabitat interspersed with cypress,rivers, and marsh on the Coastal Plainof Mississippi. Water bodies suchas Perigal Bayou, Old Fort Bayou,and Bluff Creek flow through variousunits of the refuge. Approximately100 endangered sandhill cranes inhabitthe refuge.

Refuge objectives center on theprovision of habitat for the sandhillcranes and protection of the diversesavanna communities used by cranes.Crane management includes populationmonitoring, captive bird release,predator control, and law enforcement.Habitat restoration is accomplished

via prescribed burning, vegetationmanipulation, and noxiousweed control.

National wildlife refuges and theFish and Wildlife Service: WhiteRiver National Wildlife Refuge, AR—Established in 1935, the White RiverRefuge contains the largest contiguousblock of bottomland hardwood forestunder a single ownership in the South.

White River supports one of thelargest concentrations of winteringmallard ducks in the Mississippiflyway on its 154,856 acres. Numerousspecies of wading birds, shorebirds,geese, neotropical migrants, andraptors (including the bald eagle)also inhabit the area.

Refuge objectives center on theprovision of optimum habitat formigratory bird and resident species,and support for a diversity of speciescommon to the White River bottoms.

National forests and the ForestService—The USDA Forest Service wasestablished in 1905 to provide qualitywater and timber for the Nation. Inthe subsequent years, the Forest Serviceembodied the concept of multipleuses. Multiple uses refer to resourcemanagement that benefits a varietyof purposes while ensuring theproductivity and quality of theenvironment. Benefits include theprovision of water, forage, wildlife,wood, and recreation.

The Weeks Act authorized purchaseof lands for the National Forest System,especially deforested land, which wouldbe reforested for watershed protection.The Clark-McNary Act (1924) furtherallowed the Agency to purchase privateland that was potentially valuable fortimberland production.

Acquisitions under the Weeks andClark-McNary Acts further addedarea to the National Forest System.

The mission of the Forest Servicecenters on four primary objectives: (1)protection and management of naturalresources on National Forest Systemland; (2) research on forests and forestresource utilization; (3) assistanceto State and local governments, forestindustry, and private landowners forland management; and (4) internationalassistance for the management of forestresources (Loomis 1993). The ForestService has recently issued policiesfor preservation of old growth andmaintenance of biological diversity.

National forests are found in 13Southern States, Puerto Rico, and theVirgin Islands (table 1.16). Over 15million acres in the South are managedby the Forest Service. National forestownership ranges from 27,831 acresin Puerto Rico to 2,586,074 acres inArkansas. In addition to Arkansas,the greatest concentrations of nationalforest are in Virginia (1,660,428 acres),Mississippi (1,158,967 acres), andFlorida (1,152,824 acres). Hundredsof animals and plants are supportedby the diversity of habitats in theNational Forest System.

National forests and the ForestService: roadless areas—Roadlessareas comprise nearly 1 million acresof the southern national forests (table1.17). Substantial acreages with thisdesignation are in Virginia (394,000acres) and North Carolina (172,000acres). Roadless areas have a rangeof habitat types and successionalseres. Habitat tends to be contiguous,providing refuge from humandisturbance that can disrupt speciesmovement and reproduction.

These areas possess ecologicalcharacteristics that are rare indeveloped landscapes, such aslarge, relatively undisturbed blocksof habitat (U.S. Department ofAgriculture, Forest Service 2000b).Invasion of exotic species, erosion,sedimentation, and disruption of waterflow are often less likely in roadlessthan in roaded areas. Species richnessmay be improved in roadless areas thatare large enough to offer a mosaic ofhabitat patches in various successionalstages following disturbance.

National forests and the ForestService: wilderness areas—Wilderness areas cover 698,513 acresin the South (table 1.18). Arkansas(116,937 acres), Georgia (114,789acres), and North Carolina (103,226acres) have the largest amountsof wilderness in the South (U.S.Department of Agriculture, ForestService 2000c). The Wilderness Actrequires that these areas retain theirprimeval character without permanentdevelopments or human habitation.Roads, timber harvesting, andmotorized access are prohibited, buthunting and fishing are permitted.

One objective of managing wildernessis to preserve naturally functioningecosystems. Relatively large blocksof undisturbed habitat are rare in


Table 1.16—National forest location and acreage in the South

Gross NFS OtherLocation acreage acreage acreage

AlabamaConecuh NF 171,177 83,858 87,319Talladega NF 740,334 389,328 351,006Tuskegee NF 15,628 11,252 4,376William B. Bankhead NF 348,917 180,548 168,369Talladega PU 11,706 0 11,706Pea River LUP 40 40 0

State total 1,287,802 665,026 662,776

ArkansasOuachita NFa 2,004,231 1,423,459 580,772Ozark NF 1,496,999 1,136,709 360,290St. Francis NF 29,729 21,201 8,528Ouachita PU 1,442 1,442 0Ozark PU 7,115 3,263 3,852

State total 3,539,516 2,586,074 953,442

FloridaApalachicola NF 632,890 565,543 67,347Chotawhatchee NF 1,152 1,152 0Ocala NF 430,441 383,573 46,868Oscala NF 190,932 158,255 32,677Nekoosa PU 674 223 451Pinhook PU 171,182 40,025 131,157Tates Hell-New River 6,863 4,053 2,810

State total 1,434,134 1,152,824 281,310

GeorgiaChattahoochee NF 1,515,885 749,352 766,533Oconee NF 260,883 115,231 145,652Chattahoochee PU 69,302 195 69,107Ocmulgee PU 10,000 250 9,750Yonah PU 46 46 0Forestry Sci. Lab. EA 4 4 0

State total 1,856,120 865,078 991,042

KentuckyDaniel Boone NF 1,360,692 547,686 813,006Jefferson NFa 54,614 961 53,653Land between the Lakes 170,310 170,310 0Redbird PU 686,399 145,099 541,300

State total 2,272,015 864,056 1,407,959

LouisianaKisatchie NF 1,022,373 603,230 419,143Bayou Beouf PU 2,264 980 1,284

State total 1,024,637 604,210 420,427

MississippiBienville NF 382,821 178,542 204,279De Soto NF 796,072 506,028 290,044Delta NF 118,150 60,015 58,135Holly Springs NF 519,943 155,661 364,282Lyndon B. Johnson NGL 115,438 20,309 95,129

continued


TERRESTRIAL

Table 1.16—National forest location and acreage in the South (continued)


Mississippi (cont.)Homochitto NF 373,497 191,505 181,992Holly Springs NF 119,155 66,874 52,281De Soto PU 240 240 0Homochitto PU 67 67 0Forest Hydro. Lab. EA 15 15 0Forestry Sci. Lab. EA

(state college) 7 7 0Forestry Sci. Lab. EA (Gulfport) 10 10 0Southern Hardwoods Lab EA 3 3 0

State total 2,309,980 1,158,967 1,151,013

North CarolinaCherokee NFa 327 327 0Croatan NF 308,234 159,886 148,348Nantahala NF 1,349,000 527,709 821,291Pisgah NF 1,076,511 505,420 571,091Uwharrie NF 219,757 50,189 169,568Nantahala PU 17,027 737 16,290Yadkin PU 194,496 0 194,496Forestry Sci. Lab. EA 27 27 0

State total 3,165,379 1,244,295 1,921,084

OklahomaOuachita NFa 723,552 350,845 372,707Black Kettle NGL 32,537 30,710 1,827Rita Blanca NGL 15,816 15,576 240

State total 771,905 397,131 374,774

Puerto RicoCaribbean NF 55,665 27,831 27,834

State total 55,665 27,831 27,834

South CarolinaFrancis Marion NF 414,699 252,288 162,411Sumter NF 960,805 360,868 599,937Silviculture Watershed Lab EA 15 15 0

State total 1,375,519 613,171 762,348

TennesseeCherokee NFa 1,204,520 634,198 570,322Cherokee PU 7,712 325 7,387Land between the Lakes 63,852 63,852 0

State total 1,276,084 698,375 577,709

TexasAngelina NF 402,231 153,180 249,051Davy Crockett NF 394,200 160,652 233,548Sabine NF 442,705 160,656 282,049Sam Houston NF 491,800 162,996 328,804Black Kettle NGL 576 576 0Caddo NGL 68,661 17,873 50,788Lyndon B. Johnson NGL 115,438 20,309 95,129

continued


Table 1.16—National forest location and acreage in the South (continued)


Texas (cont.)McClellan Creek NGL 1,449 1,449 0Rita Blanca NGL 77,413 77,413 0

State total 1,994,473 755,104 1,239,369

VirginiaGeorge Washington NFa 1,635,565 960,133 675,432Jefferson NFa 1,586,343 700,268 886,075Jefferson PU 1,145 0 1,145Kimberling Creek PU 271 27 244

State total 3,223,324 1,660,428 1,562,896

Grand total 28,882,907 15,644,482 13,287,425

PU = purchase unit; LUP = land utilization project; EA = experimental area; NGL = national grassland.a Property is in two or more States.Source: U.S. Department of Agriculture 2000a.

Table 1.17—Summary ofinventoried roadless areasin the South

TotalState acreage

Alabama 13,000Arkansas 95,000Florida 50,000Georgia 63,000Kentucky 3,000Louisiana 7,000Mississippi 3,000North Carolina 172,000Oklahoma 13,000South Carolina 8,000Tennessee 85,000Texas 4,000Virginia 394,000

Total 910,000

Source: U.S. Department of Agriculture 2000c.

Table 1.18—Wilderness areas in the South

NFS Other TotalState acreage acreage acreage

Alabama 32,167 80 32,247Arkansas 116,578 359 116,937Florida 74,495 4 74,499Georgia 114,537 252 114,789Kentucky 16,779 658 17,437Louisiana 8,679 0 8,679Mississippi 6,046 0 6,046North Carolina 102,634 592 103,226Oklahoma 14,543 1,425 15,968South Carolina 16,671 0 16,671Tennessee 66,349 40 66,389Texas 38,483 0 38,483Virginia 87,064 78 87,142

Total 695,025 3,488 698,513

Source: U.S. Department of Agriculture 2000a.

the South. These are of particularimportance to mammals that have largehome ranges. Importantly, wildernesscontributes to understandingwildlife in an unmanaged setting.

Implications of HabitatFragmentation onVertebrate Species

This section reviews the literature onhabitat fragmentation and the resultinginfluence on the species that inhabit

those landscapes. Two additionalchapters of the Assessment examinefragmentation in the South. Chapter 6presents an analysis of southernlocations using remotely sensedimagery. In addition, chapter 3examines the influence of roads andpower lines on habitat fragmentation.

The definition of fragmentation—The term “fragmentation” is oftenused to refer to the insularization ofhabitat on a landscape. The changein arrangement of existing habitats isoften accompanied by a loss of habitat

area. A landscape may cover hundredsof square miles or a much smaller area.The definition depends on the contextof its use and is shaped by the scaleat which ecological processes arediscussed (Trani 2002).

Fragmentation may occur when aforested landscape is subdivided intopatches. Fragmentation may also occurwhen numerous openings for suchthings as fields, roads, and power linesinterrupt a continuous forest canopy.It also can refer to discontinuities ofvegetation in the landscape. Wetland


TERRESTRIAL

habitat can become fragmented whenportions are drained for urbandevelopment, while prairie habitatcan become fragmented by agriculturaldevelopment. The resulting landscapepattern alters habitat connectivityand edge characteristics, influencinga variety of species.

Factors that contribute to landscapefragmentation—Landscape fragmen-tation may result from natural processessuch as hurricanes, wildfires, andfloods. Landscape fragmentation mayalso occur in association with landuse conversion for urban development,agricultural use, and timber harvesting.The ecological consequences of naturalor human-caused fragmentation differdepending on the pattern imposedby these factors.

Landscape modification has occurredfor thousands of years. Native inhab-itants modified landscapes by burningand clearing forested areas. The firstEuropean settlers divided vast forestsinto farmlands and settlements. Thistrend continues today. Much of thesouthern landscape is under intensivemanagement and is becoming anincreasingly complex mosaic offorest, urban, and agricultural areas.

Timber harvesting may fragmentthe landscape, depending on thenumber, size, and arrangement ofharvest units (Trani 1996). Higherlevels of fragmentation occur whensmall, numerous harvest units aredispersed over the landscape thanwhen units are clustered. A dispersedharvest scheme increases spatialheterogeneity, patchiness, and forestedge length. However, the changesin pattern resulting from timberharvest are often temporary becausethe harvested area regenerates andreverts to forest. The rate of successiondepends on the composition of theresidual stand, browsing by herbivores,subsequent management activities,weather, and other disturbances(Wigley and Roberts 1994).

It is important to note that a forestedlandscape supporting a mosaic ofdifferent seral stages is not ecologicallythe same as a landscape containingisolated forested patches surrounded byagricultural or urban areas. Each seralstage provides habitat that varies insuitability for a particular species as itmoves through the forested landscape.

Roads may contribute to forestfragmentation when their placementdivides large landscapes into smallerpatches and interior forest habitat isconverted into edge habitat. As roaddensity increases, the populationsof some species may become isolated(chapter 3). Roads located alongthe periphery of a landscape have theleast influence on the resulting pattern(Trani 1996). The influence of roadson habitat fragmentation varies withroad width and degree of permanence.A six-lane interstate highway has agreater effect on landscape patternthan does a 20-foot forest road. Someroads, such as unimproved dirt roads,may be temporary, while others arepaved and permanent.

Influence of landscape fragmen-tation upon terrestrial species—Harris (1988) cited fragmentation asthe most serious threat to biologicaldiversity in the Nation. Area-sensitivespecies requiring large tracts of habitatmay decline or be extirpated locally.The movement of species betweenpatches may be inhibited. Populationpersistence may be linked to thenumber, size, and degree of isolationof forest patches (Robbins andothers 1989).

Influence of landscape fragmen-tation upon terrestrial species—The influence of fragmentation onthe landscape can be associated withthree related factors: (1) patchiness,(2) edge, and (3) connectivity.

Influence of landscape fragmen-tation upon terrestrial species:patchiness—Changes in patch sizehave been recognized as a majorcomponent of fragmentation. Speciesrichness may decline as patch area isreduced (Ambuel and Temple 1983,Askins and others 1990, Lynch andWhigham 1984). Small remnantpatches of forest surrounded by openareas constitute unfavorable habitatfor many species; these remnantsalso have increased susceptibility towindthrow disturbance and otherprocesses. Robinson and Wilcove(1994) suggested that fragmentedlandscapes become population sinksthat are only sustained by immigrationfrom nearby forest tracts that arelarge enough to produce a surplusof individuals.

Matthiae and Stearns (1981) foundthat the density of red squirrel, graysquirrel, raccoon, and red fox increased

with habitat patch size. Fahrig andMerriam (1985) also reported thatcertain mammals were more commonin large forest tracts than in smaller,isolated patches. Populations of white-footed mice and chipmunks in smallforest patches declined to a point thatlocal extirpations occurred.

Rosenberg and Raphael (1986)reported that gray foxes, ringtail cats,and northern flying squirrels weresensitive to forest fragmentation. Picton(1979) found that the presence of largemammals was correlated with the sizeof the mountain ranges where eachspecies occurs. Mammal populationcan increase when minimum habitatsize requirements are met. Theinsularity of populations increaseswith continued landscape fragmen-tation while larger, undeveloped areasprotected these species from extinction.

Roads may or may not act as barriersto the movement of species betweenhabitat patches. Extensive networks ofroads have negative impacts on blackbears, white-tailed deer, and Floridapanthers (chapter 3). These negativeimpacts stem from loss of habitat,increased hunter accessibility, andvehicular mortality.

Long-term population declines havebeen observed for neotropical migrantsinhabiting small forest patches.Breeding bird censuses for isolatedforest patches indicate generalreductions in abundance and diversityof species over the past several years(Lynch and Whitcomb 1977). Criticalinformation for the conservation ofbird species includes understanding ofthe relationship between reproductivesuccess and habitat size and quality.The dependence of many breedingsongbirds on large blocks of forest iswell established (Robbins and others1989, Whitcomb and others 1981).

Species sensitive to patch size tendto be highly migratory, are forest-interior specialists, build open nests,and/or nest on the ground (Whitcomband others 1981). The worm-eatingwarbler, the hooded warbler, and theblack-and-white warbler are generallyabsent in patches less than 50 acres(Hamel 1992). Other species thatare sensitive to patch size include theswallow-tailed kite, broad-wingedhawk, barred owl, pileated woodpecker,and black-billed cuckoo (Hamel1992). While many species avoid


small patches, widespread permanentresidents and short-distance migrantstend to predominate in small patches(Askins and others 1990).

Habitat isolation has been associatedwith population declines in large snakesdue to increasing networks of roads(Gibbons and Buhlmann 2001). Thesenetworks divide forested habitat intosmaller and smaller parcels. Likewise,amphibian mortality is intensifiedwhen a heavily traveled road separatesindividuals from the forest they livein and the wetland they requirefor breeding.

Influence of landscape fragmen-tation upon terrestrial species:edge—An edge is the place wheretwo different plant communities,successional stages, or land uses cometogether. Fragmentation can increasethe amount of edge habitat in alandscape. Inherent edges are causedby changes in soil type or topography,whereas induced edges are thosecreated by disturbance. Induced edgescan be created by land uses, includingcultivation, fertilization, and harvest,and by environmental disturbancessuch as fires, blowdowns, and floods.

The creation of forest edge influencesseedling establishment and vegetativecomposition. For some species, theseeffects persist hundreds of yards intothe forest interior (Chen and others1992). For example, the edge habitatmay serve as an access point, attractingcowbirds into the interior of a forestedlandscape (Askins 1994).

Many species occur in edge habitat,particularly those that use one habitatfor food and another for cover. Gamebirds, such as the American woodcockand northern bobwhite quail, occur inedge habitats. Many species in urbanand agricultural landscapes are edge-adapted. Many woodland passerinesfavor edge habitat (Yahner andScott 1988), which may provideenhanced forage and/or improvedhabitat conditions.

In contrast, excessive edge maylead to reduced populations of speciesdependent on large blocks of forestinterior (Robbins and others 1989).Species that use continuous matureforest may be replaced by generalistspecies. Southern breeding birds thatnest only in the interior of forestsinclude the sharp-shinned hawk,Cooper’s hawk, hairy woodpecker,

winter wren, and veery (Hamel 1992).Edge can negatively affect these species,particularly in patches with largeperimeter-to-area ratios (Noss 1983).

An increase in density of forest-edgeand farmland species along edges mayexclude certain interior and long-distance migrant species. Competitionby the edge-adapted starling exertsa direct negative impact on manyforest species (Harris 1988). Thiscompetition may influence birdcommunity composition more thanarea-dependent changes in habitat(Ambuel and Temple 1983).

Species that occur in edge habitatsare subject to high rates of mortalityfrom predators attracted to thesehabitats. The raccoon, least weasel,and striped skunk often hunt forsmall mammals along edges. Groundnests receive predation pressure wheremammals and reptiles are the dominantpredators (Chasko and Gates 1982).Predation reduces the recruitment ofthe Kentucky warbler, scarlet tanager,wood thrush, yellow-throated vireo,and ovenbird (Temple and Cary 1988).Increases in edge density contributeto the escalation of nest predationand parasitism to levels that canbring reproductive success belowreplacement rates.

Nest parasitism by cowbird speciesmay be an important factor in thedecline of some breeding birds. Broodparasites lay their eggs in the nests ofother species, reducing the reproductivesuccess of their hosts. The brown-headed cowbird may have contributedto the population declines of theAcadian flycatcher, veery, Americanredstart, and Louisiana waterthrush(Brittingham and Temple 1983).

Influence of landscape fragmen-tation upon terrestrial species:connectivity—Connectivity, the degreeof continuity of a landscape, is alsoaffected by fragmentation. Connectivitymay facilitate dispersal and improvehabitat quality by connecting patchesof habitat. It has been suggested thatthe population dynamics of speciesare affected by the spatial pattern offragmentation (Haddad and others2000, Hanski 1991). There isdisagreement, however, on the valueof corridors for the conservation ofbiological diversity. One view is thatpopulations linked by corridors arevulnerable to the spread of disease

and several environmental stressors(Gilpin 1987, Quinn and Hastings1987). If corridors spread the riskof environmental stress amongisolated populations, persistencetime may actually be longer infragmented landscapes (Fahrigand Paloheimo 1988).

Another view suggests that speciespersistence is lower in fragmentedhabitats than in contiguous habitats(Tilman and others 1994). Thesestudies suggest that corridors arevaluable as a conservation tool.This point of view is discussed next.

Heany and Patterson (1986)presented an extensive review ofthe regional patterns of mammaldistribution as affected by habitatconnectivity. Pelton (1986) describedhow the loss of connectivity restrictsthe distribution of black bears. Whendisturbance causes local extirpation,populations may be reestablishedthrough the dispersal of individualsfrom source populations. Jackson(1987) reported corridors aided red-cockaded woodpeckers in colonizingexisting habitat. Forest birds can oftenuse small tracts of forest connectedto large tracts by wooded corridors(Robbins 1979). Forest-interiorbirds and small mammals (Merriam1990) persist in forest fragmentsconnected by woodland corridorsthat ease colonization.

Species that are able to movebetween connected habitat patchesoperate demographically as ametapopulation. Corridors maypermit the survival of extinction-pronepopulations through the immigrationof individuals. Corridors also mayfacilitate movement of an individualwithin its home range. Such movementmay be particularly important forspecies whose home range arearequirements exceed the averagepatch size. For example, Rosenburgand others (1997) reported thatmigratory amphibians, such as red-spotted newts, may require corridorsamong seasonally used habitats. Theloss of connectivity may cause localextirpation. Many amphibian andreptile species cannot move throughrelatively large, deforested areasto reach other suitable forest habitat.Where declines of herpetofaunalpopulations occur, population sizes


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will not be rebuilt quickly in afragmented landscape (Gibbonsand Buhlmann 2001).

Discussion andConclusions

Status and Trendsof Terrestrial VertebrateSpecies

Natural Heritage classifies 86percent of southern vertebrate speciesas secure or apparently secure. Thepopulations of these species appearto be resilient; some species suchas white-tailed deer and beaver haverebounded despite incredible odds.Population trends are positive forseveral big game, small game, andwaterfowl species. In addition, thelong-term population projectionsfor several furbearer species appearstable or increasing.

In contrast, declines in thepopulations of northern bobwhitequail, ruffed grouse, and woodcockwarrant further management focus.The decline in breeding populationsof grassland and shrubland nestingbirds also is a concern in the region.The numerous species with G1, G2,or G3 conservation ranks suggestthat these vertebrates are sensitiveto changes in their environment.Identifying the factors that contributeto the declines of these species may beuseful for predicting future conditions.Several of these factors, as well as theirassociated conservation measures,are examined in chapter 5.

Significant losses of communitybiodiversity have occurred throughoutthe region. Several communities havebeen classified as critically endangered,endangered, or threatened. An addi-tional 24 communities have beenidentified as having a 50-percent lossof presettlement area. It is criticalto halt further losses of thesecommunities and to raise publicawareness through education.

There appears to be a commonalityof threats to sensitive species andcommunities of the South. Manyspecies and communities experienceddeclines associated with humandisturbance and settlement patterns.The growth of human populationsin the South will continue to pressurespecies and the communities that

support them. Vertebrate speciesand their associated habitats areinfluenced by urban development,fire suppression, agricultural practices,forest pest and exotic species outbreaks,and recreation activity. Other speciesare rare due to restrictive or specializedhabitat conditions (chapter 2).

The future of a majority of thesesensitive species and communitiesin the South depends on active resto-ration and management. Restorationcomplements species conservation bymaintaining habitat composition,structure, and function. Activitiesthat mimic natural disturbance areparticularly important. Prescribedburning can enhance herbaceousdiversity and control structuralcharacteristics. Other treatments areuseful for suppressing woody growthand enhancing the vigor of otherspecies. These management techniquesare described further in chapter 4.

Hard and Soft MastFor many species, mast is an essential

food source. Thus, provision of hardand soft mast is important for themanagement of terrestrial speciesinhabiting southern forests.

Many silvicultural techniquesenhance mast production (chapter 4).Management of stocking densitycan encourage reproduction ofmast-producing species and limitinterspecific competition. Artificialregeneration has been successfulfor several species, including northernred oak, white oak, and black cherry.Genetic selection for acorn productionand seedling growth also has thepotential to be successful. Thesetreatments can play an importantrole in southern forest areas thatmay experience mast decline.

The Implications ofHabitat Fragmentation

Extensive literature suggests thatlandscape patterns affect the abundanceand persistence of terrestrial species.The fragmentation of the landscape,and the consequences of thatfragmentation on ecosystems andpopulation dynamics, are concernsshared across the region.

Natural processes and humanactivities may influence habitat lossand isolation. Changes in patchiness,edge, and connectivity may eliminate,

displace, or enhance species popu-lations and habitats. Isolated habitatpatches may reduce the numberof species present simply becausesmaller habitats support fewer species(MacArthur and Wilson 1967).Preservation of species compositionand integrity in these areas cannot beexpected. Corridors may increase themovement of habitat-restricted species,thereby improving overall habitatquality (Haddad and Baum 1999,Rosenburg and others 1998).

Understanding how spatial patternsalter species habitat may provideresource managers with a basis formaking land use decisions. Speciesrespond to patterns in various ways,using certain areas for feeding andreproduction, and avoiding other areasentirely. By altering the distribution andavailability of spatial resources, changesin landscape pattern influence manyof the components important for thepersistence of species (Merriam 1990).

The South’s growing humanpopulation raises the possibilityof a substantial impact on speciesand their habitats in the next severaldecades (chapter 6). In the midstof expanding populations, the provisionof biological diversity has becomea critical conservation issue.

The Influence of LandOwnership Patterns

The population increases projectedfor the South may continually increasedemands on natural ecosystems,species, and their habitats duringthe 21st century (Boyce and Martin1993). This prospect presents achallenge to forest resource manage-ment. Biodiversity often declinesas economic development proceeds.Natural habitats for native speciesare replaced by industrial and urbandevelopment, while other habitatsare modified or degraded. The futuremay also bring increased concern forconservation of endangered speciesand habitats and the reservation oflands for aesthetic and recreationvalues (Boyce and Martin 1993).

These changes highlight theimportant role that public landswill have in the conservation of speciesand their habitats. The Forest Service,Fish and Wildlife Service, and NationalPark Service manage millions of acresin the South. Other agencies, such asthe U.S. Department of Defense and


the Tennessee Valley Authority, alsomanage critical habitat areas. Thereare numerous Federal policies thatdictate the management andconservation of natural resources.

Without these public lands, manyspecies would be in trouble. Forexample, over 53 percent of the specieswith viability concerns in the Ozarkand Ouachita Highlands are knownto occur only on national forests (U.S.Department of Agriculture, ForestService 1999). The Peaks of Ottersalamander is an example of animperiled species that occurs solelyon Federal land—in this case, theGeorge Washington and JeffersonNational Forests and the Blue RidgeParkway. The Federal land in theFlorida Panhandle and the centralAppalachian Mountains supportsconcentrations of imperiled andlisted species (Stein and others 2000).National wildlife refuges play a keyrole in the protection of listed speciessuch as the red wolf and the Floridapanther, and in the provision ofkey areas of habitat for waterfowl,migratory birds, and many otherspecies. National parks are importantfor the preservation and managementof old growth, spruce-fir, and otherrare and sensitive communities of bothplants and animals. National forestsare key in the provision of wildernessareas, large blocks of forest interior,and a diversity of habitats.

Other public lands are also importantfor the conservation of species and theirhabitats. State agencies own significantareas designated as parks, wildlifemanagement areas, forests, or naturalresource areas. While the purposesof such areas vary, the conservationof biological diversity is often oneobjective for these properties. InFlorida, State agencies are carryingout aggressive land acquisitionprograms for conserving biodiversity,using shared Federal excise taxrevenues as a funding source. City andcounty governments also own a varietyof land in parks and recreation areasthat support species and their habitats.

Many imperiled and endangeredspecies are found on public land,and this land represents a relativelysmall percentage of forest land in theSouth. It seems clear, therefore thatpublic land is vital for maintainingimperiled and endangered species(Stein and others 2000).

The area of public land is beingsupplemented by acquisition effortsby private conservation organizations.The Nature Conservancy, the Trustfor Public Lands, and Ducks Unlimitedacquire land for conservation purposes.They either manage it or transferit to public agencies. The NatureConservancy has created its ownsystem of conservation propertiesin the South. In contrast, the Trustfor Public Lands acquires land forultimate ownership and managementby public resource agencies. Manyof the trust’s land transactions havebeen from forest industry lands thatwere important biologically.

The magnitude of private ownershipalso presents a significant challengefor southern forests. Individual land-owners are changing the characteristicsof future forest resources. For example,the absence of management on privateland may result in declines in earlysuccessional habitat in many areas(Trani and others 2001). The smalltracts typical of present land usepatterns often provide little opportunityfor forest management and naturaldisturbance sufficient to create earlysuccessional forest. A myriad of speciesmay be influenced by this condition.

The Forest Service and other partnershave initiated active reforestationprograms with the private sector as partof the Lower Mississippi Valley JointVenture. Land clearing and alterationof hydrology have resulted in environ-mental degradation throughout thevalley. This step towards changingprivate land use practices may leadto restoration of the bottomlandhardwood system, the provision ofquality habitat, economic opportunitiesfor landowners, and a reliable woodsupply to meet society’s needs.

The significant numbers of imperiledand endangered species inhabitingprivate land indicate the criticalimportance of this land for conservation(Stein and others 2000). For thisreason, a variety of strategies designedto encourage conservation on privateareas have been implemented bygovernment agencies. Incentiveprograms have been created toencourage reforestation of privateland. Recognizing the significanceof private land to the imperiled speciesof the region is essential. Often, wildlifeconservation may be more importantthan timber production on this land.

Industry land also offers opportunitiesto provide wildlife habitat. Given theincentive of green certificationprograms and the scale of theiroperations, many large corporationsare taking positive actions to protectsensitive biological resources on theirproperty (Stein and others 2000).

Industry land supports breeding birdspecies, game species, and other species(Wigley and others 2000). Individualcompanies work with governmentagencies to identify threatened andendangered species on their land.The Special Sites program withinthe Sustainable Forestry Initiativemanages ecological sites to maintainwetlands, longleaf pine, and otherunique communities (Weyerhaeuserand Price 2001).

Forest industry has also donatedthousands of acres to State agenciesand the Nature Conservancy (Owenand Helssenbuttel 1989). Donationsinclude the Beryl Anthony WildlifeManagement Area in Arkansas (7,000acres), Great Dismal Swamp NationalWildlife Refuge in Virginia (60,000acres), and several wildlife manage-ment areas.

The significance of many typesof landowners in the South inproviding wildlife habitat cannotbe overstated. Each major landownerhas an important role to play inthe conservation of species andtheir habitats.

Needs for AdditionalResearch

Data are needed on the distribution,population dynamics, and habitatrequirements of many southern species.Basic life history and managementinformation is lacking for severalthreatened and endangered species.For some nongame birds and gamespecies, standardized inventories lendthemselves to regional assessments.For most species, however, there is adearth of monitoring information fromwhich to evaluate regional conditions.

Centers of amphibian and reptilediversity should be identified insensitive communities. Long-termmonitoring of amphibian and reptilepopulations is needed to establishpopulation trends. Further studyalso is warranted to assess the impact


TERRESTRIAL

the expected climate changes mayhave on amphibians and othersensitive species.

Further research is desirable intomanagement techniques that mimicnatural disturbance for the creation oflandscape patterns that are consistentwith the evolutionary history of species.Applied research is needed to identifythe best approaches, including burning,for restoring degraded communitiesand maintaining sensitive communities.

Finally, methods should be developedto quantify and forecast influencesof human developments on southernbiodiversity. We must identify verte-brate species that may be influencedby future habitat fragmentation, andexamine how fragmentation attributeschange over time.

Acknowledgments

My appreciation is extended to JohnGreis and David Wear for theirencouragement and support duringthe Assessment process. John Pyegraciously handled the Internetconversion of the document.

Appreciation is extended to TomDarden and his staff in the SouthernRegional Office for their ongoingsupport and assistance. An earlierdraft benefited from insightful com-ments by Dennis Krusac and GeorgeBukenhofer, USDA Forest Service,and six anonymous scientific reviewers.Their comments strengthened andimproved the final manuscript.

Gratitude also is given to ChuckHunter, U.S. Fish and WildlifeService, for supplying the physio-graphic region map and drafts of hisupcoming publications. Paris Griep,USDA Forest Service, contacted Stateagency biologists for information on thecurrent status and habitat concerns forblack bear. Joe Johnston and KennethGraham, U.S. Fish and Wildlife Service,supplied recovery plans and otherinformation on listed species. TheSouthern Forest History Associationassisted with literature searches relatingto historical species and conditions.

Special thanks to James Dickson,Louisiana State University, for supply-ing chapters of his upcoming textbookon southern wildlife. Credit also isgiven to Curt Flather, Rocky MountainResearch Station, for the use of maps

and figures from the RPA. DavidMeriwether, USDA Forest Service,Sheila Colwell, National Park Service,and Roger Boykin, U.S. Fish andWildlife Service, provided informationon land ownership within theirrespective agencies. M. J. Mac,U.S. Geological Survey, and PeterWhite, University of North Carolina,supplied the maps showing Federalland ownership and distributionof longleaf pine in the South.

In addition, many other organizationsand individuals supplied informationand ideas for this chapter, includingBrant Miller, Ben Wigley, Lisa Stocker,National Council of the Paper Industryfor Air and Stream Improvement,Southern Environmental LawCenter, and Larry Walker.

I would also like to thank thequestion managers and participants(general public, industry, agencies,and organizations) for sharingtheir ideas and concerns duringAssessment meetings.

Finally, my appreciation is offeredto several USDA Forest Serviceindividuals who assisted with theInternet searches, and the productionof tables and figures. These includedPal Mattox, Mike Donahue, ChristineGabbard, Brenda Ayers, Kay Spangler,and Ron Underwood.

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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.

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