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U.S. FISH AND WILDLIFE ~ Southeast

U.S. FISH AND WILDLIFE · of gopher tortoise habitats. Auffenberg and Iverson (1979) report a positive correlation between the amount of herbaceous ground cover and tortoise density,

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Page 1: U.S. FISH AND WILDLIFE · of gopher tortoise habitats. Auffenberg and Iverson (1979) report a positive correlation between the amount of herbaceous ground cover and tortoise density,

U.S.FISH AND WILDLIFE ~Southeast

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GOPHERTORTOISE

Gopherus polvphemus

RECOVERYPLAN

Prepared byWendell A. Neal

U.S. Fish and Wildlife Service

for

Southeast RegionU.S. Fish and Wildlife Service

Atlanta, Georgia

A.

Approved:R ion U.S. Fish and Wildlife Service

December 26, 1990Date:

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Recovery plans delineate reasonable actions which are believed tobe required to recover and/or protect the listed species. Plansare prepared by the U.S. Fish and Wildlife Service, sometimeswith the assistance of recovery teams, contractors, Stateagencies, and others. Objectives will only be attained and fundsexpended contingent upon appropriations, priorities, and otherbudgetary constraints. Recovery plans do not necessarilyrepresent the views nor the official positions or approvals ofany individuals or agencies, other than the U.S. Fish andWildlife Service, involved in the plan formulation. Theyrepresent the official position of the U.S. Fish and WildlifeService only after they have been signed by the Regional Directoror Director as approved. Approved recovery plans are subject tomodification as dictated by new findings, changes in species’status, and the completion of recovery tasks.

Literature Citations should read as follows:

U.S. Fish and Wildlife Service. 1990. Gopher Tortoise RecoveryPlan. U.S. Fish and Wildlife Service, Jackson, Mississippi.28 pp.

ADDITIONAL COPIES ~AY BE PURCHASEDFROM:

Fish and Wildlife Reference Service:5430 Grosvenor Lane, Suite 110Bethesda, Maryland 20814

A.

301/492—6403 or1/800/582—3421

The fee for the plan varies depending on the number of pages.

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ACKNOWLEDGEMENTS

Ed Wester of Auburn University circulated this plan amongknowledgeable persons on the Gopher Tortoise Council for review.Robert H. Mount, Professor Emeritus, Auburn University; JoanDiemer of the Florida Game and Fresh Water Fish Commission andRen Lohoefener of the U.S. Fish and Wildlife Service aregratefully acknowledged for making significant contributions tothe development of this plan.

The U.S. Fish and Wildlife Service thanks Ellen Nicol, an artist,writer, and reptile breeder from Anthony, Florida, for the coversketch.

A.

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EXECUTIVE SUMMARY

Current Status: The western population of the gopher tortoise islisted as threatened. This population lies west of the Tombigbeeand Mobile Rivers in Alabama, across south Mississippi andincluding extreme southeastern Louisiana. Threats includehabitat alterations and illegal taking.

Habitat Recmirements and Limiting Factors: The species is foundon droughty, deep sand ridges which originally supported longleafpine and patches of scrub oak. The most significant threats tothe species are adverse habitat alteration, taking, anddevelopment of occupied habitats.

Recovery Objective: The two objectives of this plan consistof an immediate objective which is prevention of the listedpopulation from becoming endangered and a long—term objectivewhich is delisting.

Recovery Criteria: The necessary criteria for the aboveobjectives are:

(1) Successful prevention of endangeredstatus would beconsidered by evidence of an average of 5 gophertortoise burrows per hectare (ha) on deep sandy soils(1.52 meters(+)) for a period of 30 years on the DeSotoNational Forest. This would equate to an estimatedpopulation of 22,400 gopher tortoises on 7,343 ha ofsuitable habitat.

(2) For delisting, evidence is required of an averageof 3 gopher tortoise burrows per ha on deep sandy soils(1.52 meters(+)) on private lands. This would equateto an estimated population of 34,000 gopher tortoiseson 18,594 ha on privately-cyned lands.

7 -

Actions Needed

:

(1) Survey, monitor and assess status of populations asbaseline for recovery actions.

(2) Protect and manage habitat on Federal lands.(3) Encouragemanagementof populations on private lands.(4) Develop law enforcement strategy to curb illegal

taking.(5) Conduct population viability studies.(6) Conduct telemetry studies to determine extent of

reproductive isolation as a threat.(7) Conduct genetic studies.(8) Relocate threatened isolated individuals/colonies to

protected and managed lands.

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-~ Total Estimated Costs of Recovery: Implementation of therecovery tasks for which cost estimates have been made total$433,000.00.

Date of Recovery: Unable to determine at this time due to theunknown response of the gopher tortoise population to improvedmanagement activities.

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TP1BLE OF CONTENTS

I. INTRODUCTION

A. Background

B. Description and Taxonomy

C. Life History and EcologyDistributionHabitatLongevity and ReproductionFoodActivity/MovementAdult MovementsBehavior

II.

III.

IV.

D. Threats and Causes for DeclineHabitat AlterationPredationOther MortalityPopulation Viability

RECOVERY

A. Biological Perspective

B. Objectives

C. Narrative Outline

D. Literature Cited

IMPLEMENTATION SCHEDULE

APPENDIX

List of Reviewers

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PAGE

1

1

1

22256777

88

101112

13

13

14

15

20

24

27

27

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I. INTRODUCTION

A. Background

The gopher tortoise (Gopherus polyphemus) is the onlytortoise indigenous to the southeastern United States.It is found in varying numbers in xeric sandy habitatsfrom South Carolina- through Florida and west to extremesoutheastern Louisiana. Within xeric sandy habitats, therange of G. polyphemus nearly coincides with the originalrange of the longleaf pine (Pinus palustris)

.

On July 18, 1984, Drs. Ren Lohoefener and Lynn Lohmeierpetitioned the U.S. Fish and Wildlife Service to list thepopulation of G. polv~hemuswest of the Tombigbee and MobileRivers under provisions of the EndangeredSpecies Act. Thepetition and accompanying report (Lohoefener and Lohmeier1984) presented substantial information on numbers anddistribution of the western population. The Fish andWildlife Service reviewed the petitioned action and onJuly 7, 1987, listed the western population as threatenedunder the Endangered Species Act (52 FR 25376—25380).

The basic biology of the tortoise has been reasonablywell documented, although many specific details remainunknown. Many biological parameters for this species varyconsiderably, including: age (or size) at sexual maturity,clutch size, growth rates, phenological characteristics,burrow depths, specific food habits, and others (Diemer1986). Biological information on G. polyphemus mostlyoriginates from Georgia and Florida. This plan drawsprimarily from the research in Georgia by Landers andBuckner (1981) since their study sites are more similar tothe western population (by latitude) than to populations inFlorida. This recovery plan is aimed specifically at thewestern population, but of necessity relies greatly upondata sources and expertise developed elsewhere.

A.-p

-pB. Description and Taxonomy

Gopherus polyphemus (Testudines, Testudinidae), describedin 1802 by F.M. Daudin, is the only Gopherus in thesoutheastern United States. The gopher tortoise has alarge shell, 15—37 centimeters (cm) (5.9—14.6 inches) long.It is a dark-brown to grayish-black terrestrial turtle withelephantine hind feet, shovel-like forefeet, and a gularprojection beneath the head on the yellowish, hingelessplastron or undershell (Ernst and Barbour 1972). Gophertortoise hatchlings are yellowish—orange, have a soft shell,and are 4-5 cm (1.5—2.0 inches) long at hatching.

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Go~herus polyphemus is sexually dimorphic. In most cases,the sex of adults can be determined by shell dimensions.The male has a greater degree of plastral (lower shell)concavity, and a longer gular projection. However, the sexof tortoises around the size of maturity can be almostimpossible to assess.

C. Life History and Ecolocry

Distribution

Historically, the western population was found in thelongleaf pine hills of northern Mobile, Washington,and southeastern Choctaw Counties in Alabama; in thesoutheasternupland areas of the pinehills province inMississippi (a 14-county area); and in the upland pineridges in St. Tammany, Washington, and Tangipahoa Parishes,Louisiana (Lohoefener and Lohmeier 1984) (Figure 1). Theamount of gopher tortoise habitat, as defined by Lohoefenerand Lohmeier (1984), for the listed population by State isas follows: southwestern Alabama — 40,770 hectares (ha) or100,741 acres (A); Louisiana — 4,815 ha or 11,898 A; andMississippi — 102,084 ha or 252,246 A. The entire westernpopulation is found within the original range of thelongleaf pine.

Habitat

Gopher tortoises occupy a wide range of upland habitattypes; however, general physical and biotic featuresprovided by Landers (1980) with slight modifications,characterize most suitable habitat. These are:

1. the presence of well-drained, sandy soils, which alloweasy burrowing (because of lower ambient temperatures,the western population may require a meter or more ofsandy soil depths);

2. an abundance of herbaceous ground cover; and

3. a generally open canopy and sparse shrub cover, which

allow sunlight to reach the forest floor.

Juvenile habitat is generally considered to be similar to

that of adults.

The traditional habitats of the western population of gophertortoises are natural xeric communities, mostly of thelongleaf-pine-scrub oak type, located on sand ridges. Theoriginal ecology of these xeric, fire—dependent communities

2

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has been significantly altered. Gopher tortoises may alsobe found in ruderal habitats such as fence rows, pastures,and field edges and power lines.

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3

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C,—C—

0~

C—’.

Figure 1. Range of Western Population of the Gopher Tortoise.

4

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Soil conditions are responsible for the xerophytic natureof gopher tortoise habitats. Auffenberg and Iverson (1979)report a positive correlation between the amount ofherbaceous ground cover and tortoise density, with grasses,grass—like plants and legumes being the most importantfood plants (Garner and Landers 1981). The amount andkind of low growing (within reach of a gopher tortoise)herbaceous plants may be a function of many variables,including timber age, density and species composition,burning history, nature and timing of past soil disturbance,and inherent soil fertility.

A relatively open canopy is necessary not only forherbaceous food plants but also for egg incubation. Thefemale gopher tortoise selects a bare spot for nestexcavation, normally in the mound of excavated sand atthe burrow entrance. Landers and Buckner (1981) notedthat when overstory overshadowedthe burrow entrance, nestswere selected in openings such as firelanes or roadsides.

The burrow is the focal point of many above groundactivities and a major portion of the gopher tortoise’slife is spent in the burrow. Most burrows have a singleentrance, and adult burrows average about 4.5 meters (in)(15 feet) in length with a depth of 1.8 m (6 feet) (Hansen1963). Small juveniles use similarly small burrows, oftenas shallow as a few inches. Single tortoises often excavatemore than one burrow. Lohoefener and Lohmeier (1984)reported a correction factor of 0.625 in Mississippi forconverting burrows counted to burrows occupied. The burrowprovides protection from fire, predators, and climaticextremes, and habitat for a host of unique species. Jacksonand Milstrey (1989) reported more than 60 vertebrate and302 invertebrates species using gopher tortoise burrows.Some of the more commonly known burrow associates includethe eastern diamondback rattlesnake (Crotalus adamanteus)

,

the gopher frog (Rana areolata) , A.and the eastern indigosnake (Drvmarchon corais couper~i)

.

Loricrevity and Reproduction

Longevity is estimated at 40-60 years (Landers 1980) andmay extend to 80—100 years (Landers et al. 1982). Growthannuli on scutes become worn at 20—40 years, making agedetermination imprecise. Age at sexual maturity in theGeorgia study (Landers et al. 1982) ranged from 19-21 yearsfor females. These animals had a plastral length of25—26.5 cm (9.8—10.4 inches). Males normally reachreproductive maturity at a smaller size and younger agethan fomal~. Growth rates vary with environmental andgenetic factors among gopher tortoise populations.

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Breeding periods may begin as early as February and extendinto September, depending on location. The period ofmaximum reproductive activity reported by Landers et al

.

(1980) is May 18 through June 27. Iverson (1980) reportedthe nesting peak in Florida also to be May and June. Clutchsizes in Mississippi average 4.8 eggs (Lohoefener andLohmeier 1984); however, this report was based on a rathersmall sample (N=14). Landers et ~ (1980) reported arange in clutch size of 4-12 eggs with a mean and SD of7.0 + 1.7. He also found that clutch size increased withthe size of the female. The lower value reported byLohoefener and Lohineier (1984) may have been due to limitedsampling, the result of human depredation (leaving primarilysmaller nesting females), or a combination of both. Thenest is usually 15—25 cm (6—10 inches) beneath the surface(Landers et al. 1980). Incubation periods range from80-90 days in northern Florida (Iverson 1980) to 110 daysin South Carolina, the northern limit of the gophertortoise’s range (Wright 1982). Most gopher tortoise eggsnever hatch becauseof predation.

Food

The gopher tortoise is the primary grazer in its xerichabitats (Landers 1980) and aids in seed dispersal fornative grasses (Auffenberg 1966). Observations and studiesof food habits come mainly from Georgia and Florida wherewiregrass (Aristida stricta) is often considered animportant food plant and is a common member of the longleaf-scrub oak community. However, in western parts of thecoastal plain, bluestem grasses (Andropocron) are often themost common herbaceousspecies in mature longleaf pineforests (Wahlenberg 1946). Lohoefener and Lohmeier (1981)observed tortoises in Mississippi eating crabgrass(Dicritaria sancruinalis) and panic grasses (Panicum)

.

Garner and Landers (1981) found that broad-leaved grasseswere staple foods while wiregras~. was used mainly in earlyspring and summer. Their studyX~lso sh6wed that wildlegumes (Fabaceae), which are high in protein, were usedextensively by juveniles. Garner and Landers (1981) alsofound that fleshy fruits were readily consumed, includingblackberry (Rubus cunefolius), sloeplum (Prunus umbellata)

,

blueberry (Vaccinium), maypop (Passiflora lutea), andhawthorne (Crataecrus). Regardless of the specific plantsavailable for forage, the conclusion reached by Garner andLanders (1981) that “grasses, grass—like plants and legumesare the most important food plants and evidently determinecarrying capacity” is likely a statement equally applicableto the western population.

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‘Act ivitv/Movement

McRae et al. (1981) found activity to be very restrictedduring winter months. In fact, from late November throughFebruary, feeding activity was observed only five times. Onunusually warm winter days when maximum temperature exceeded260 Celsius (C) or 790 Fahrenheit (F), tortoises wereoccasionally observed at the burrow entrance (McRae ~1981). No crepuscular or nocturnal activity is reported.As temperatures rose during the spring (March and April),outside burrow activity was most often observed in theGeorgia study during the warmest part of the day, 1600-1800h(hours). During July and August, McRae et al. (1981) founda bimodal movementpattern, the feeding forays peaking atmid—morning (l000—1200h) and mid—afternoon (1600—1800h),with much reduced activity during the hottest part of theday, 1300-1500h. They concluded that “activity throughoutthe year was correlated with ambient temperature; movementfrom the burrow was rare at coolest temperatures (<220 C or72~ F), was greatest at 28 to 310 C (82 to 880 F), and wascurtailed at >320 C (900 F).”

Adult Movements

McRae et al. (1981) studied movement related to feedingseparately from movements related to other behavior anddetermined 95 percent of all feeding activity took placewithin 30 m (33 yards) of the burrow being used. Auffenbergand Iverson (1979) reported increasing foraging radii fromthe burrow in areas with reduced ground cover. Thissuggests that food availability can increase or decreaseforaging distances. McRae et al. (1981) trailed 13 adultsand determined their movements to be in a nearly circularor elliptical pattern around the burrow. Depletion ofpreferred foods near burrows by late summer is thought tocontribute to larger movements later in the year. In theGeorgia study, the home ranges of =ales were much largerthan females; males had a home x=nge of~ 0.06—1.44 ha(0.14—3.56 A) with a mean of 0.47 ha (1.16 A), while femaleshad a home range of 0.04-0.14 ha (0.10—0.35 A) with a. meanof 0.08 ha (0.20 A) (McRae et al. 1981). The sexualdifferences are attributed to breeding forays by the males.Landers and Speake (1980) found the average colony typicallyused an area less than 4 ha (9.88 A).

Behavior

Gopher tortoises have a well—developed social structure,courtship, and territorial combat (Auffenberg 1966, Douglass1976, McRae et al. 1981). Males bob their heads to attractfemales during the breeding season. The speed and amplitude

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of the head bobbing increases as the male draws closer to areproductively active female, and the first contact betweenindividuals consists of males biting females on theforelimbs and around the gular area, perhaps seekingolfactory cues (Auffenberg 1966). When males confront eachother, there is usually some manifestation of dominance orsubmissive behavior. According to !4cRae et al. (1981),there is a dominance hierarchy in males based on size. Indense populations, smaller males are found around thecolony’s periphery rather than in the middle, close to thebreeding females, as is the case with larger males.

D. Threats and Causes for Decline

Habitat Alteration

An understanding of the reasonsbehind the threatened statusof G. polv-phemus is perhaps the most essential step indeveloping this recovery plan. The gopher tortoise,historically and currently, is a component of xeric plantcommunities originally identified mostly by the occurrenceof longleaf pine. The changes altering the originallongleaf pine communities also changed the ecosystem of thegopher tortoise. This species was an animal of theseforests, and to the extent maintenance of the listedpopulation is possible, that goal is inextricably tied toforestland conditions.

Before the arrival of European colonists in the New World,the longleaf pine was the principal tree species onsoutheasterncoastal plain upland soils. Croker (1987)cites 60 million acres in the original stands which heconcludes are now reduced to about 4 million acres. Afterthe red and white pine forests of New England and the GreatLake States were cut, lumbermen turned to the virginlongleaf stands, the mining of which peaked in 1909(Croker 1987). Power skidders andrailroad logging

A.supported these final assaults.,-’- -

Second growth longleaf pine stands came from the ruins oftimber mining operations, but these second forestsconstituted a small fraction of the area of virgin stands.Becauseof planting difficulties with the longleaf pine,these droughty sites were often planted in slash(P. elliottii) and loblolly (~. taeda) pines. Thispractice, along with excessive burning intervals andintensive site preparation methods, continues on soilswhich originally supported longleaf pine.

Artificial planting of longleaf is now successful and manyforesters are rediscovering the valuable traits of longleafpine, including the fact that it can be successfully

8

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regenerated-naturally through a shelterwood system of -

cutting combined with burning just in advance of an adequateseed fall. The U.S. Forest Service recently has adopted apractice of regenerating only longleaf pines on longleafsites in the DeSoto National Forest. However, the agency’spreferred method is by planting. Most private landownerscontinue to regenerate longleaf pine sites to off—sitespecies.

The original longleaf pine community burned and reseedednaturally. It contained trees of many ages and a diverseground cover with much edge, which would be of particularimportance to the gopher tortoise. Landers and Speake(1980) found better gopher tortoise densities in longleafpine — scrub oak stands that were thinned and burned every2—4 years. Slash pine plantations, with a similar system ofthinning and burning, had sparser population densities.While it is apparent that gopher tortoises can be maintainedunder a modified (heavily thinned, frequently burned)plantation system of management,Landers and Buckner (1981)showedthat gopher tortoise densities are significantlygreater (32 percent) in more naturally managedstands oflongleaf.

The natural longleaf pine community and its associatedbiological diversity represent optimal forest habitat forthe gopher tortoise. This community occurred in purestands, constantly trending toward small even—agedgroups ofa few hundred square feet (Chapman 1909). Larger even-agedpatches and strips were found following blowdowns fromsevere weather. These were often interspersed with patchesor single survivors, creating open glades and a patchinesswhich favored the gopher tortoise. Management practiceswhich alter this system include: clearcuts of large blocks(including the crowded planting of off—site species),diversity—diminishing soil churning activities that oftenaccompanyeven—agedtimber manag9m~nt, and prolonged burningintervals. Timber practices that most nearly mirror thenatural system, such as a shelterwood regeneration systemwith frequent burning and natural regeneration, improve thesoil and herbaceous cover condition to optimally support thegopher tortoise.

Longleaf pine trees, as well as fire—dependent annuals andperennials, originally existed in a summerburning cyclewhich has long since been interrupted. The change in firefrequency and timing may be the single most important factorinfluencing other alterations which have changedtheoriginal xeric communities. For example, it has been acommon practice to remove most of the longleaf pines fromthese dry ridges and then to exclude fire (or at least fail

9

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to burn). This allows eventual occupancy by poor siteoaks (Ouercus laevis, 2. incana, Q. marilandica, andQ. marcraretta) and woody shrubs such as yaupon ~vomitoria) and gallberry (~. crlabra). When the leaf litterfrom oaks becomesa thick mat, it retards fires that wouldotherwise be carried by longleaf pine needles and the commongrass associates under the open longleaf pine canopy. Fireexclusion allows the oaks to mature and shade out herbaceousground cover neededby gopher tortoises. This situation isnot uncommon throughout the range of the gopher tortoise.Landers and Speake (1980) provided substantial evidence thatthese altered sites originally were good gopher tortoisehabitat but now support the fewest gopher tortoises.

Hedrick and Zimmermann (1988) monitored gopher tortoisedensities in various forest types and classes for atwo—year period on the Conecuh National Forest in Alabama.Their unpublished data indicate gopher tortoise densitiesthrough three stand conditions (seedling/sapling stands,pole stands, and sawtimber stands). Gopher density wasgreatest (1 active burrow/1.51 ha or 3.73 A) in theseedling/sapling stands, greatly reduced (200 percent) inpole stands (1 active burrow/3.10 ha or 7.66 A) and followedby a large recovery (177 percent) in sawtimber (1 activeburrow/1.75 ha or 4.32 A).

The current threats to the western population of the gophertortoise in terms of habitat loss or degradation consist ofcertain forest management practices, conversion of dry sitesto agriculture, road placement and other developments onthese higher ridges, and urbanization (Lohoefener andLohmeier 1984).

Predation

The gopher tortoise was a signific~nt food source during theGreat Depression, as reflected iiy the name “Hoover Chicken”(Hutt 1967). Gopher pulling reiri’oves an average of 20percent of the larger tortoises, according to Taylor (1982).The taking of gopher tortoises by pulling (use of a longflexible rod with a hook) remains a cultural ethos in ruralareas where the western population is found. The gophertortoise’s low reproductive rate, high mortality of eggs andyoung, slow growth to sexual maturity, and long lifeindicate a K-selected strategy adapting to xeric communities(Landers 1980). Annual population growth may only be3-5 percent (Landers et al. 1980); accordingly, humanpredation on mature adults may produce long term adverseeffects which are difficult to overcome. Because manygopher tortoises exist in degraded or declining habitats

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and populations are often fragmented, the adverse effects ofeven limited taking may be exacerbated. Lohoefener andLohmeier (1984) report a significant number of Mississippigopher tortoises being taken for pets.

Gopher tortoise predators, other than -human beings, aremany. The most important egg and hatchling predatorappears to be the raccoon (Procyon lotor) (Landers andSpeake 1980); however, a variety of mammals are reportedpredators of G. polyphemus, including gray foxes (Urocvoncinereoarcrenteus), striped skunks (Mephitis mephitis)

,

opossums (Didelphis vir~iniana), armadillos (Dasvnusnoveincinctus) (Landers et al. 1980), and dogs (Canisdomesticus) (Causey and Cude 1978). Imported fire ants(Solenopsis saevissima and/or ~. victa) are reported ashatchling predators (Landers et al. 1980, Lohoefener andLohmeier 1984). Snakes and raptors have also been reportedas preying on G. polyphemus. Reported clutch and hatchlinglosses often approach 90 percent (Landers et al. 1980).

Other Mortality

Road mortality is reported by Landers and Buckner (1981) andLohoefener and Lohmeier (1984) as a significant mortalityfactor. Lohoefener and Lohmeier (1984) believe nests andjuveniles are often destroyed by intensive site preparation(heavy equipment). Tanner and Terry (1981) report a majorreduction in burrow density in Florida which was believedattributable to roller chopping or web plowing. Diemer andMoler (1982) demonstrated that tortoises are able to digout following chopping treatment on deep sandy soils, butconcluded that additional data were neededregardingtortoise response to various site preparation techniquesin different soil types.

Lohoefener and Lohmeier (1981) believed that a seriousproblem for the Mississippi goph9rtortoise was isolation ofsexually mature animals becaus&of habitat fragmentationaggravated by forest management practices. Only 14 percentof the tortoises encountered in density survey transectsby Lohoefener and Lohmeier (1981) in Mississippi wereconsidered so situated that interactions with other sizeable(sexually mature) tortoises might occur. As further supportfor this hypothesis, the discontinuous nature and small sizeof Mississippi sand ridges, which are often separated bystreams or wet boggy areas, may serve as impediments tocourtship travels of adult males (Lohoefener and Lohmeier1984)

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Population Viability

Local populations of the western gopher tortoise can intheory become extirpated through chance events and theseextirpations (and thus more rangewide extirpations) areinversely related to population size. Shaffer (1981) citesfour sources of uncertainty to which a population may besubject: (1) demographic stochasticity, which arisesfrom chance events in the survival and reproductive successof a finite number of individuals; (2) environmentalstochasticity due to temporal variation of habitatparameters and the populations of competitors, predators,parasites, and diseases; (3) natural catastrophes, such asfloods, fires, and droughts, which may occur at randomintervals through time; and (4) genetic stochasticityresulting from changes in genetic frequencies due to foundereffect, random fixation, or inbreeding. Based on theconcern expressed by Lohoefener and Lohmeier (1984)regarding reproductive isolation, genetic drift andinbreeding may already be occurring.

Recovery, therefore, must consider population viability inestablishing both the objectives and the procedures formeeting those objectives.

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II: RECOVERY

A. Biolocrical Perspective

The listed population of G. polyphemus could be consideredrelatively abundant. Lohoefener and Lohmeier (1984)estimated 10,923 tortoises of >23 cm (9.1 inches) carapacelength (CL) in 102,084 ha (252,246 A) of Mississippihabitat; and 12,900 tortoises >23cm (9.1 inches) CL wereestimated to occur in 40,370 ha (99,753 A) of Alabamahabitat west of the Tombigbee and Mobile Rivers. However,the species is nearing extinction in an estimated 4,815 ha(11,898 A) of Louisiana habitat. About 80 percent(121,000 ha) of the available habitat occurs on corporately-owned lands.

Despite the relatively large number of extant individualsestimated, the long—term prospects for survival of thewestern population are dimming. In view of past, current,and predicted forest managementpractices, continued illegaltaking, development on dry uplands, and private ownershipof much of the gopher tortoise’s habitat, this speciesis truly threatened in the western portion of its range.According to Donner and Hines (1987), timberland ownershipin south Mississippi is mostly private (85 percent belongingto individuals, the forest industry and corporations,11 percent belonging to the Federal government, withthe remainder in State or county ownership).

Section 7 of the EndangeredSpecies Act requires Federalagencies to insure that their actions do not jeopardize thecontinued existence of listed species. Beyond the jeopardyprohibition, Section 7 requires Federal agencies to usetheir authorities to further the purpose of the Act. Theessential purpose of the Act is conservation of listedspecies. Section 7 is limited in scope to Federal actions.Thus, the role of Section 7 in re~overy of this species willbe limited becausethe majorit< of habitat is in non-Federalownership. However, any advice given by Federal forestersor soil scientists to manage forests on state, local, andprivate lands is also subject to Section 7. Outside ofSection 7, the Act may serve in protection, and therefore,possibly contribute to recovery, through exposure of certainactivities under Section 9 (prohibition of take).

Through consultations with Federal landowners, it isexpected that forest managementpractices will be designedto contribute significantly to recovery on these lands.However, becauseFederal ownership is comparatively small,rangewide recovery for this population requires significantsuccess on privately—owned lands as well. Examples of suchactivities can be found in Mount et al. (1988).

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Unfortunately, among private timberland owners, there areperceived problems with longleaf pine, its growth, value,and availability of seed stock. Individual small landownersoften high grade their longleaf stands with littleforethought to long—term timber production; they thenexclude fire, thus creating a situation where the longleafpine sites convert to scrub oak stands. If these landownersdecide to regenerate, they will most often, on the advice offoresters, choose the off-site slash or loblolly. Suchadvice from Federal foresters or foresters supported byFederal monies should be subject to Section 7 consultation.The corporate or industrial landowner usually farms thesesandy sites by clearcutting, replanting to off—site species,and starting over with the same practices at a 25—35 yearrotation, devoting little attention between planting andharvest. These managementpolicies, along with intensivesite preparation, thick planting rates, and fire exclusioncontinue to threaten the existence of the westernpopulation.

B. Recovery Oblectives

The immediate recovery objective is to prevent the westernpopulation from becoming an endangeredspecies. To achievethis, the species’ overall status must be stabilized orenhanced. Lohoefener et al. (in review) considersthree to seven burrows per hectare as representing arecovered population density for a land unit the size ofDeSoto National Forest. The upland forested habitatexpected to support this density is likely underlain byLakeland, Troup, or one of the more rarely encountered deep,sandy soils in excess of 1.52 meters (5 feet). On the DesotoNational Forest, these soils are estimated to comprise7,343 ha (18,144 A) (Arnold 1989). The best hope forrecovery of the gopher tortoise is on these 7,343 ha ofdeep sands that represent original sandhill communities [andpotentially provide the best ch~n~e for a largeblock of contiguous habitat being made available to gophertortoises]. A range of three to seven burrows perha = 22,029—51,401 x 0.61 (correction factor of tortoisesper burrow) = 13,437—31,354 gopher tortoises. If amid-range density of five gopher tortoise burrows per ha(approximately equating to a total of 22,400 gophertortoises]) is accomplished on the Desoto National Forest,and maintained for a period of 30 years, the immediate goalof preventing the listed population from becoming endangeredwould be reached. Although little is known about the ratesof gopher tortoise recruitment and present age-class

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distribution, this recovery objective assumesthat oncethe stated density is maintained for 30 years that therecruitment rate is adequate for short-term stability.

A long-term objective, that of recovery to the point of nolonger requiring protection of the Act, requires significantsuccesseson the privately—owned lands having these deepsand ridges. Within the range of the western population, onprivate land, there are approximately 18,594 ha (45,945 A)of what originally constituted sandhill communities.Attaining the lower range of the recovery density for deepsands based on Lohoefener et al. (in review) would meanthree burrows per ha (18,594 x 3 x 0.61) = (approximately34,000 gopher tortoises on privately owned forested deepsands. To measure these goals, some form of survey isnecessary and must be comparable to the originalstatistically derived estimate (Lohoefener and Lohmeier1984)

C. Narrative Outline

1. Survey, monitor. and assessthe status of copulations

.

The original survey work by Lohoefener and Lohmeier(1984) needs to be updated to monitor status. Thereremains controversy about the abundanceof the gophertortoise. A survey will clarify the tortoise’s status;moreover, it will provide an essential baseline formeasuring the effectiveness of recovery activities.Surveys should also attempt to determine recruitmentrates and age-class distribution, if possible.

1.1 Survey cro~her tortoise populations on Federal andother public lands not previously surveyed

.

Baseline surveys will be necessary to track theeffectiveness of habitat management.

1.1.1 Conduct status sui~reys on Camp Shelby

.

This requirement is incorporated intoSection 7 compliance.

1.1.2 Conduct status surveys on DeSoto NationalForest. This requirement is incorporatedinto Section 7 compliance.

1.1.3 Conduct surveys on State—owned Parklands

,

Wildlife Manacrement Areas and 16th SectionSchool lands. Colonies on public landsoffer possibilities for conservationunavailable on private lands.

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1.2 Conduct rancrewide surveys at 5—year intervals on -

public and private land. This is necessary todetermine the effectiveness of recoveryactivities. Surveys must be comparable bytechnique to existing data (Lohoefener andLohmeier 1984), repeatable, and carried out duringthe same month becausetortoise movementsandburrow use may vary monthly.

1.2.1 Assess the status of individual populationsand of the species ranaewide. The goal ofthe recovery plan is to eliminate factorsdetrimental to the survival and recovery ofthe gopher tortoise. As data are acquired,the status of populations throughout therange will be reviewed and assessed asappropriate.

2. Implement protection and managementof habitat onFederal lands. The principal threats on Federal lands,specifically the DeSoto National Forest, have been:(1) adverse timber managementpractices on the high,dry ridges where gopher tortoises occur, and (2) themilitary use of about 136,000 acres. These threats arebeing addressedthrough Section 7 consultationinvolving both Camp Shelby’s land-altering activitiesand a habitat managementplan by the Forest Service.The review of these actions will be an ongoingactivity.

2.1 Protect and manage all existing cro~her tortoisecolonies. The colony sites on Camp Shelby will beprotected either by staking burrows with steelposts or by fencing the colony site. Formanagementpurposes, a gopher colony is defined asthree or more active adult burrows (=9inches inwidth) within 300 feet of each other, or anycombination of active,--=dultand activehatchling/sub-adult burrows within 100 yards ofeach other; the colony site is defined as theactive burrows making up a colony plus a 200—footbuffer around them.

Timber stands on Federal lands, where a colony islocated, will be managed primarily for the gophertortoise. Such management considerations willaddress: canopy closure in the stand, mid—storymanagement, regeneration and site—preparation,planting rates, thinnings, burning and/or chemicaltreatment of hardwoods for colony sitereclamation, and scheduling of harvest to avoiddisturbance during nesting periods.

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2.2 Manacre habitat for Present and future expansion.-

In order to reverse declines in gopher tortoisepopulations, it will be necessaryto manageforoptimum habitat conditions on some part of Federalownerships. The Camp Shelby Section 7consultation has resulted in the establishment ofa 2,200—acregopher refuge where military use isrestricted and forest management is aimed atachieving and maintaining optimal habitatconditions.

2.3 Assess adeauacvof established and proposedmanacrementplans. This is a continuous taskaccomplished largely through Section 7 of theEndangeredSpeciesAct. All Federal agencies mustreview their established and proposed programs,and for those that may affect the species,initiate consultation with the Fish and WildlifeService. The Service will then review the actionand prepare a biological opinion which addressesthe likelihood of jeopardy to the continuedexistence of the species if the action is carriedout. If jeopardy is likely, alternatives toremove jeopardy are presented in the opinion. Allmanagementprograms for the species represent a“may affect” situation requiring consultation.

3. Encouracre protection and management on private lands

.

Private lands contain the vast majority of forestpossibly containing gopher tortoises. Accordingly,maintenance of the population is not possible withoutsome significant successes on privately—ownedtimberlands. Promotion of protection and management ofhabitat on private lands is difficult becauseof thefew legal responsibilities and the perceived economicinterests of landowners. Therefore, special effortsare neededon private lands. -~

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3.1 Provide information on manacrement and lecralreauirements to private landowners and manacrers

.

3.1.1 Develop informational articles andmanacrement cruidelines oriented to privatelands. Informational articles andmanagement guidelines oriented to privatelands should be developed. These articlesand guidelines should include informationand visual aids which identify the habitatof the species, and give detailed optionsby which the species’ welfare can be

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maintained or enhanced without altering thetotal land management objectives of theowner or manager. These educationalefforts could also emphasizethecompatibility of gopher tortoise managementwith deer and quail m~.nagement. Legalresponsibilities of private landowners,through Sections 7 and 9 of the EndangeredSpecies Act, should also be explained.

3.1.2 Distribute information to privatelandowners and managersthroughprofessional and industrial associations

.

The information developed in 3.1.1 shouldbe distributed through a variety ofprofessional and industrial associationsand agencies, such as the State and privateforestry branch of the U.S. Forest Service,county agricultural extension agents, andState forestry and wildlife associations.

3.2 Develop a cooperative aareement between the Fishand Wildlife Service and private landowners andimplement where feasible. This agreement shouldspecify management actions needed to protect thespecies and should identify the party responsible(landowner or Federal agency) for implementing thevarious actions. The agreementshould set forththe total commitments of the two parties includingland base, funds, equipment, manpower, and timeperiod, and provide a means and a time frame forterminating the agreement.

3.3 Protect gopher tortoise habitat through easements

,

acouisitions, and donations. Lands containinggopher tortoises should receive specialconsideration when thesfr lands would consolidateFederal ownership or cbntrol br would contributeto overall resource management objectives of theagencies. Private landowners should be encouragedto avail themselves of these options.

3.4 Recognize or reward protection and managementefforts. Management efforts on private landsshould be recognized and rewarded in view of thelimited legal responsibilities involved. Newsmedia should be contacted and encouraged toprovide favorable publicity to deservinglandowners. News articles should be prepared forthe news media where desirable or requested.

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4. - Develop law enforcement strategy to curb illegal takingof cro~her tortoises. Gopher tortoise depredation byhumans remains a practice in the rural areas where thelisted population occurs. Habitat protection may befor naught if “taking” pressures continue to impactpopulations. Law enforcement must be a cooperativeeffort among the Fish and Wildlife Service, U.S. ForestService, and the States. This effort may or may notinvolve the use of publicity.

5. Conduct research on population viability. This isneededto determine what densities and distributionsare necessary to achieve minimum viable populationsnecessary for recovery goals. These factors are stillunknown; yet they may eventually control the results ofany scheduled recovery activity. Three areas, criticalto understanding population viability, requiringbaseline data, are (1) recruitment rates, (2) presentage-class distribution, and (3) what constitutescontiguous habitat for the species.

6. Conduct telemetry studies. This is needed to determinewhether or not seemingly isolated tortoises(particularly males) are in fact interacting with othertortoises. Data from telemetry studies will also yieldinformation on what constitutes contiguous habitat forgopher tortoises.

7. Conduct genetic studies. This is needed to answerquestions on the effects of augmentation and relocationefforts.

8. Relocate reproductively isolated individuals toexisting Protected and managed colonies. Animals thatare determined to be in this category add nothing tomaintenanceor recovery. If introduced into anexisting small colony whichi~ protected and managed,they may contribute to the?recovery goal. Suchrelocation should be done in accordance with theprocedures outlined in Mount et al. (1988).

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D. Literature Cited

Arnold, Thomas. 1989. Soils most suitable for gophertortoise occurring on the Desoto National Forest.Unpublished memo. 1 p.

Auffenberg, W. 1966. On the courtship of Go~heruspolyphemus. Herpetologica 22:113-117.

Auffenberg, W. and J. Iverson. 1979. Demographyofterrestrial turtles. pp. 541—569. In M. Harless andN. Norlock (eds.). Turtles: Perspectives andresearch. Wiley—International, New York.

Causey, M.K., and C.A. Cude. 1978. Feral dog predation ofthe gopher tortoise, Go~herus polyphemus, in southeastAlabama. Herp. Rev. 9:94-95.

Chapman, H.H. 1909. Longleaf Pine in Texas. Soc. Am.Foresters Proc, Vol. 4 Pp. 207—220.

Croker, T. 1987. Longleaf Pine: A History of Man and aForest. USDA Forest Service Forestry Report R8-FR7.37 pp.

Daudin, F.M. 1802. Histoire naturelle generale etparticuliere des reptiles Vol. 2 F.Dufart, Paris. 432pp. Paris 2:1—326.

Diemer, J.E. 1986. The ecology and management of thegopher tortoise in the southeastern United States.Herpetologica 42:125—133.

_________ and P.E. Moler. 1982. flopher tortoise responseto site npreparation in nor~h~rn Florida. Proc. Ann.Conf. Southeast. Assoc. Fish and Wildl. Agencies36:634—637.

Donner, B.L. and F.D. Hines. 1987. Forest statistics forMississippi Counties - 1987. Res. Bull. 50—129. NewOrleans, LA. U.S. Dept. of Agriculture, ForestService, Southern Forest Exp. Stat. 79 pp.

Douglass, J. 1976. Mating system of the gopher tortoise(Gopherus polyphemus) in southern Florida. M.S.Thesis, Univ. South FL, Tampa. 76 pp.

Ernst, C.H., and R.W. Barbour. 1972. Turtles of the UnitedStates. The University Press of Kentucky, Lexington.347 pp.

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Garner, J.H. and J.L. Landers. 1981. Foods and habitat ofthe gopher tortoise in SouthwesternGeorgia. Proc.Ann. Conf. S. E. Assoc. Fish and Wildl. Agencies35:120—133.

Hansen, K.L. 1963. The burrow of the gopher tortoise.J. Florida Acad. Sci. 26:353—360.

Hedrick, L. and J. Zimmermann. 1988. Unpublished data fromthe ConecuhNational Forest, Alabama.

Hutt, A. 1967. The gopher tortoise, a versatilevegetarian. FL Wildl. 21(7):20-24.

Iverson, J.B. 1980. The reproductive biology of Go~heruspolyphemus. Am. Midl. Nat. 103:353—359.

Jackson, D.R. and E.G. Milstrey. 1989. The fauna ofgopher tortoise burrows. Pp. 86—98. In J.E. Diemer,D.R. Jackson, J.L. Landers, J.N. Layne and D.A. Wood(eds.). Gopher Tortoise Relocation Symp. Proc. FloridaGame and Fresh Water Fish Comm. Nongame Wildl. Prog.Tech. Rep. # 5.

Landers, J.L. 1980. Recent research on the gopher tortoiseand its implications. Pp. 8-14. In R. Franz andR.J. Bryant (eds.). The dilemma of the gophertortoise—is there a solution? Proc. 1st Ann. Mtg.,Gopher Tortoise Council. Florida State Museum,Gainesville, Fl.

Landers, L. and J.L. Buckner. 1981. The gopher tortoise:effects of forest managementand critical aspects ofits ecology. Southlands Exp. For. Tech. Note 56. 7 p.

Landers, J., and D. Speake. 1980. Managementneeds ofsandhill reptiles in southern Georgia. Proc. Ann.Conf. S.E. Assoc. Game and2F~ish Wildl. Agencies34:515—529.

_________ J.A. Garner and W. A. McRae. 1980.Reproduction of the gopher tortoise (Gopheruspolyphemus) in southwestern Georgia. Bull. FloridaState Mus. Biol. Sci. 36:353—361.

_______ W.A. McRae and J.A. Garner. 1982. Growth andmaturity of the gopher tortoise in southwesternGeorgia. Bull. Florida State Mus. Biol. Sci. 27:81-110.

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Lohoefener, R. and L. Lohmeier. 1981. Comparison of gophertortoise (Gopherus polyphemus) habitats in young slashpine and old longleaf pine areas of southernMississippi. J. Herp. 15:239—242.

Lohoefener, R. and L. Lohmeier. 1984. The status ofGo~herus ~olv~hemus (Testudines, Testudinidae) west ofthe Tombigbee and Mobile Rivers. Unpublished Rept.Submitted to U.S. Fish and Wildl. Ser. 104 pp.

Lohoefener, R., M. Bailey, A. Ziinmermann and L. Hedrick.(In review]. Gopher tortoise burrows on the ConecuhNational Forest, Alabama. [Submitted to Journal ofWildlife Management. J

McRae, W.A., J.L. Landers and J.A. Garner. 1981. Movementpatterns and home range of the gopher tortoise. Am.Midl. Nat. 106:165—179.

Mount, R., E. Wester and C. Swing. 1988. Biologicalassessment of land altering activities of theMississippi Army National Guard on the federallythreatened gopher tortoise (Gopherus polyphemus) atCamp Shelby Military Reservation, Mississippi. SpecialReport for National Guard Bureau’s Compliance withSection 7 of the Endangered Species Act. 67 pp.

Shaffer, M. 1981. Minimum population sizes for speciesconservation. Bioscience 31:131—134.

Tanner, G. and W. Terry. 1981. Effects of roller choppingand web plowing on gopher tortoise burrows in southernFlorida. Pp. 66—73. In R. Lohoefener, L. Lohmeier andG. Johnson (eds.). The future of gopher tortoisehabitats. Proc. 2nd Ann. Mtg. Gopher Tortoise Council.Florida State Museum, Gainesvflle, FL.

A.‘--p

Taylor, R. 1982. Human predation on the gopher tortoise(Gopherus polyphemus) in north-central Florida. Bull.Florida State Mus., Biol. Sci. 28(4).

U.S. Fish and Wildlife Service. 1987. Endangered andThreatened Wildlife and Plants; Determination ofthreatened status for the gopher tortoise (Gopheruspolyphemus) . Federal Register 52(129) :25376—25380).

Wahlenberg, W.G. 1946. Longleaf Pine. Charles LathropPark Forestry Foundation. Washington, D.C. 37 pp.

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Wright, 5. 1982. The distribution and population biology ofthe gopher tortoise (Gopherus polyphemus) in SouthCarolina. M.S. Thesis, Clemson University, Clemson,South Carolina.

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PART III

IMPLEMENTATIONSCHEDULE

Priorities in column one of the following implementation schedule

are assigned as follows:

1. Priority 1 - An action that must be taken to preventextinction or to prevent the species from decliningirreversibly in the foreseeable future.

2. Priority 2 - An action that must be taken to prevent asignificant decline in species population/habitatquality or some other significant negative impact shortof extinction.

3. Priority 3 - All other actions necessary to meet the

recovery objective.

Key to Acronyms Used in This Implementation Schedule

MDWFP = Mississippi Department of Wildlife, Fisheries and ParksUSFS = U.S. Forest ServiceLDWF = Louisiana Department of Wildlife and FisheriesALDNR = Alabama Department of Natural Resources

A.,7

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)

‘.3

___________ IIlhItHtNIAI_loll_SCIIW4JIt

R~5YOad5lMI.E PAth COST IIIIIIAIIS

(55)

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ralo.- lASS tASKiv ii ii CLWIENIS/NOIIS

Ill E lASS 5 VI5CMIPIION OU*AhlOhI aegloii Provraia Oilier lPfl 1~Z 1~95

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~ 4 ewE tuuipiUSI s

25 ‘Other agaiwlee’ reaponalbhtltlestaut be a coopirailve lialW4•possibly on projects j&ee~e~i

levier a Service contract. in s~ceces contracts way be tat to

prIvate ilbJlVi4Jals. lb. ArMyMat lonet Gueral ageS USfS are

obl1~eta~4 to certain actionslbrOti~l*h Sect Ion I of lit. Act.

2 5.5.1 Leap Sliellay eurvey ci year 4 IUE MDUIS/usfs

5

2 1.1.2 PsEolo NeCIwtel forestaurecy

‘I year 4 hOUfV/us, s

25

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20

2 5.2 Pupulatiwi eurvey/nut I

2 years CUE hIIJUfP/13Sf S

35 lepeat every 5 years.

2 1.2.5 Aceece rauagaul4e status 2 yeara 4 CUE W)UIP/us’s

5 Repeat Ovary 1 years.

3 5 Proiccllout seal aenaue-se~uu ad private lejele

Coopuaretlve agreaeacutts

contIg~oue 4 tUE CIL)lJfP/USfS

10

5 5.2 ‘5 year 4 CUE I4UUIVI13515

5 5 5 Costa Ia be deteratned.

5 3.3 Eaua~nte/douaatIo~s 5 year 4 CUE CIDUfP/us, S

5 5 Coals t@ Sue determined.

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Page 33: U.S. FISH AND WILDLIFE · of gopher tortoise habitats. Auffenberg and Iverson (1979) report a positive correlation between the amount of herbaceous ground cover and tortoise density,

)))

IHPIftENIAIION SCHEDUlE

tRio.-its a tASK ~

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U0115111E PARIr cost ESTIMATEs(SE)

C(~OhiMISjM0hE5 C

US, US

other IV1991 ii1992 IV1993RdMIon Protraus

3 3.4 t~uer&la ~I year 4 IUE WJUtP/US’S

5 5 5 Coats to be determined.

2 4 I.tw eguIu~cccuueu±u~t&lrelc~y

~1 year 4 tUE I4OUtPIUStS

S 5 5 ContifluJOtJa

3 5 Se~asrch pupulatlonhy

3 yasra 4 IUE l0UfS’/tilt S

20 20 20

3 6 Ra~earch venedca 1 y.er 4 fUE lJnIv.r-a lay

20

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IS IS

3 Kuhuaete raiiiJ~uctiveh~\ia”lsi~J turlola.es

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15 10

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Coothgajoua 4 fliP IIDUIPIUSfS

30

3 3.2 Coop~rsihv. sureaau.uaAta 43 year 4 IUE IlOUfl’/U515

5 5 5 Coats to tie determined.

3 3.3 E~nls/tIogu.ethons I veer 4 fliP IQIJIP/tilt S

5 5 5 Costa to be determined.

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IV: APPENDIX

List of Reviewers

C. Kenneth DoddU.S. Fish and Wildlife Service412 NE 16th Ave., Room 250Gainesville, FL 32601

Ren Lohoefener1900 Martin Bluff RoadGautier, MS 39553

Mr. Jack StoutUniversity of Central FloridaDepartment of Biological

SciencesOrlando, FL 32816—0368

Bob JonesMS Museum of Natural Science111 N. Jefferson StreetJackson, MS 39212

Ed WesterCary HallAuburn University, AL 36849-5508

Nora MurdockU.S. Fish and Wildlife Service100 Otis Street, Room 224Asheville, NC 28801

James H. BakerENSR Consulting & Engineering3000 Richmond AvenueHouston, TX 77098

Samuel K. McLellanLieutenant ColonelCorps of Engineers, MSARNGPost Office Box 5027Jackson, MS 39296—5027

Ms. Virginia Van SickleDepartment of Wildlife and

FisheriesPost Office Box 98000Baton Rouge, LA 70898

Joan E. DiemerFL Game & Freshwater Fish Comm.Wildlife Research Laboratory4005 South Main StreetGainesville, FL 32601

Dr. Robert H. MountCary HallAuburn University, AL 36849-5508

Joe DuckworthDistrict RangerU.S. Forest ServicePost Office Box 248Wiggins, MS 39577

Larry D. Hedrick, Staff OfficerU.S. Forest ServiceFisheries, Wildlife & RangePost Office Box 1270Hot Springs, AR 71902

Gopher Tortoise CouncilFL Museum of Natural HistoryUniversity of FLGainesville, FL 32611

Lisa A. LofdahlTenneco Gas CompanyPostOffice Box 2511101ff Milam ~StreetHouston, TX 77252—2511

John E. AlcockRegional ForesterU.S. Forest Service1720 Peachtree Road, NWAtlanta, GA 30367

Dave FlemmingU.S. Fish and Wildlife Service75 spring Street, S.W., Room 1276Atlanta, GA 30303

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A1ice~ R. ThomasCitizen Enforcement Project

NRDC40 West 20th StreetNew York, NY 10011

Fred C. SchmidtHead, Documents Dept.The LibrariesColorado State UniversityFort Collins, CO 80523

LibraryBrookfield ZooBrookfield, IL 60513

A-

28