SPECIES: Gopherus polyphemus
Table of Contents


INTRODUCTORY


  A gopher tortoise in Naples, Florida. Photo courtesy of Charles Warren.
AUTHORSHIP AND CITATION:
Innes, Robin J. 2009. Gopherus polyphemus. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/ [ ].

FEIS ABBREVIATION:
GOPO

COMMON NAMES:
gopher tortoise
gopher turtle

TAXONOMY:
The scientific name for the gopher tortoise is Gopherus polyphemus (Daudin). The gopher tortoise is in the Testudinae family [40]. No subspecies are recognized [54,112], although there is considerable geographic variation in size at sexual maturity, duration of winter dormancy, color, adult size, and clutch size (review by [7]). There are 3 distinct genetic assemblages of gopher tortoises: western (i.e., populations in Louisiana, Alabama, and the Florida panhandle), eastern (i.e., populations in Georgia and most of the peninsula of Florida), and the Brooksville Ridge on the west-central coast of the Florida peninsula [122].

SYNONYMS:
Testudo polyphemus Daudin [133]

ORDER:
Testudines

CLASS:
Chelonia

FEDERAL LEGAL STATUS:
The segment of the population that occurs west of the Tombigbee and Mobile Rivers in Alabama, Mississippi, and Louisiana is federally listed as Threatened [145].

OTHER STATUS:
Information on state-level status of animals in the United States is available at NatureServe, although recent changes in status may not be included.

WILDLIFE DISTRIBUTION AND OCCURRENCE

SPECIES: Gopherus polyphemus
GENERAL DISTRIBUTION:
The gopher tortoise is a terrestrial turtle native to the southeastern United States. It occurs in the southeastern Atlantic Coastal Plain from southern South Carolina southwest to eastern Louisiana and south through peninsular Florida [54]. It was introduced on Cumberland Island, Georgia [36]. Gopher tortoise populations have been fragmented by urban and agricultural development (see Status and threats) [9]. In many regions, gopher tortoise populations are small and discontinuous, particularly south of Sarasota County, Florida (Auffenberg 1978 cited in [39]) and areas on the periphery of the gopher tortoise's range [9,45,95,137]. NatureServe provides a distributional map of the gopher tortoise.

PLANT COMMUNITIES:
Gopher tortoises occur in a variety of plant communities, including pine-oak (Pinus-Quercus spp.) sandhills, pine flatwoods, pine plantations, sand pine (P. clausa) scrubs, coastal strands, coastal prairie grasslands, hammocks, and a variety of ruderal communities ([80], reviews by [39,54]). Ecotones between communities are also used ([47,59], reviews by [7,9]). Gopher tortoises prefer open habitats that support a variety of herbaceous ground cover vegetation for forage (see Diet) and usually abandon densely canopied areas (see Preferred Habitat). According to a review, the gopher tortoise is associated with the following forested and nonforested plant communities throughout its range [7]:

Mean gopher tortoise densities, level of suitability, and potential for gopher tortoise forage in forested and nonforested habitats [7]
Habitat Level of suitability** Potential for forage** Mean gopher tortoises/acre
Forested habitats
Upland coniferous forests
     Pine flatwoods
          Longleaf pine (Pinus palustris) 9 10 5.5
          Longleaf pine/saw-palmetto (Serenoa repens) 8 10 4.2
          Longleaf pine/gallberry (Ilex spp.) 6 9 1.2
          Longleaf pine plantation 6 6 1.4
          Slash pine (P. elliottii)/saw-palmetto 6 8 0.6
          Slash pine/oak/gallberry 5 8 0.5
          Slash pine plantation—early stage (open canopy) 7 7 4.0
          Slash pine plantation—late stage (closed canopy) 1 2 0.1
  Pine-mesic oak hammock (longleaf, slash, and loblolly (P. taeda) pines, southern red (Quercus falcata), water (Q. nigra), chestnut (Q. prinus), and laurel (Q. laurifolia) oaks, hickory (Carya sp.), dogwood (Cornus sp.), and sweetgum (Liquidambar styraciflua))
3 4 0.2
     Pine-xeric oak sandhills 10 10 4.2
          Longleaf pine-blackjack oak (Q. marilandica) 10 10 4.2
          Longleaf pine-bluejack oak (Q. incana) 9 9 2.5
          Longleaf pine-turkey oak (Q. laevis) 10 10 4.3
          Longleaf pine-sand post oak (Q. margarettiae) 9 9 3.8
     Sand pine scrub/rosemary (Ceratiola ericoides) 6 6 1.2
     Sand pine plantation—early stage (open canopy) 5 6 1.2
Upland hardwood forests
     Xeric oak hammock 7 7 2.8
     Live oak hammock (mesic) 5 5 0.5
     Sand live oak (Q. geminata) hammock (xeric) 6 7 1.3
Nonforested habitats
     Natural grassland 10 10 5.2
     Pasture (e.g., bahia grass (Paspalum notatum) pasture) 8 9 3.5
     Abandoned pasture 8 10 4.2
     Cropland (row and field) 5 8 0.4
     Abandoned citrus (Citrus spp.) groves 9 9 5.0
     Fallow cropland 7 9 3.0
     Herbaceous rangeland 9 10 3.2
     Shrubby rangeland 7 9 2.5
     Coastal strand—herbaceous 8 8 1.8
     Coastal dune scrub—shrubby 8 8 4.5
     Ephemeral ponds* 7 7 ---***
     Freshwater marsh* 5 7 ---
     Wet prairie* 5 7 ---
     Disturbed areas (e.g., roads, railroad beds, or berms) 6 7 1.0
     Ruderal (e.g., fence rows, orchards, or golf courses) 8 8 3.2
*Used in drought or when seasonally dry.
**1=low, 10=high.
***No data.

This review subjectively ranked the "level of suitability" of forested and nonforested habitats for gopher tortoises based upon information available in literature, unpublished data, and expert opinion. Because herbaceous plant species composition is highly variable within and across habitats and dependent upon current and historic management practices, the "potential for forage" occurrence within each habitat was also ranked. Data from numerous published and unpublished studies were compiled to estimate gopher tortoise densities in each habitat presented in the above table [7].

According to the State of Florida Game and Fresh Water Fish Commission, gopher tortoises occupy the following habitats in Florida [15]:

Gopher tortoise habitats in Florida using 3 habitat classification systems [15]. Additional habitats listed by Cox and others [39] are highlighted in bold.
US Soil Conservation Service Florida Land-Use and Cover Florida Natural Areas Inventory
Florida coastal strand (north and south) Grassland Coastal strand
Sand other than beaches Overwash plain
 

Beach dune

  Maritime hammocks
  Coastal berm
  Shell mound
Florida flatwoods (north and south) Palmetto (e.g., saw-palmetto and scrub palmetto (Sabal etonia)) prairies Mesic flatwoods
Pine flatwoods Scrubby flatwoods
Coniferous planted forests Dry prairie
  Wet flatwoods
  Wet prairie
  Prairie hammock
Sand pine scrub Coastal scrub Scrub
Other scrub and brush Xeric hammock
Sand pine scrub  
Coniferous planted forests  
Longleaf pine-turkey oak hills Longleaf pine-xeric oak Sandhill
Coniferous planted forests  
Mixed hardwoods and pine Mixed forest Upland pine forest
Coniferous planted forests Upland mixed forest
Hardwood planted forests Xeric hammocks
Upland hardwood hammocks Other hardwood Upland hardwood forest
Hardwood planted forests Xeric hammocks
Oak hammocks Xeric oak forest Xeric hammocks
Ruderal Recreational  
Mixed  
Open land and other  
Cropland and pastureland  
Orchards, groves (except citrus)  
Citrus groves  
Grassland  
Clearcut areas  
Altered lands  
Other barren lands  

Forested habitats: Although gopher tortoises occur in many plant communities, longleaf pine ecosystems, which are divided into 2 general types—pine-oak sandhills and pine flatwoods—comprise about 80% of the habitats associated with the gopher tortoise (review by [7]). Longleaf pine and threeawn (Aristida spp.) are characteristic species of "virtually all" longleaf pine ecosystems associated with gopher tortoises [119]. Gopher tortoises occur at high densities in midsuccessional stages of longleaf pine ecosystems maintained by fire and are less abundant in more advanced successional stages such as hammocks [9]. Historically, large areas of longleaf pine ecosystems were converted to plantations with slash pine, sand pine, or loblolly pine. Gopher tortoises may occur at moderate to high densities in early-successional stages of pine plantations but tend to occur at lower densities in late stages with closed canopies [7].

Pine-oak sandhills: Gopher tortoises occur at high densities in pine-oak sandhills. These plant communities extend over a large portion of the Atlantic Coastal Plain in the southeastern United States. Widely spaced longleaf pine typically dominates the overstory of pine-oak sandhills [58]. Less frequently, slash pine dominates the overstory [117,140]. The understory is typically sparse and consists of a mixture of deciduous oaks, including turkey, blackjack, bluejack, and post oak, and a variety of shrubs (e.g., blueberry (Vaccinium spp.), lyonia (Lyonia spp.), and gallberry) [7,31,58,117,140,151]. The ground layer is typically dense and consists of diverse herbaceous plants [58]. The understory and herbaceous ground cover associated with pine-oak sandhills vary across the gopher tortoise's range [21]. In the eastern part of the gopher tortoise's range, the gopher tortoise is primarily associated with the longleaf pine/turkey oak/threeawn (e.g., pineland threeawn (Aristida stricta) and Beyrich threeawn (A. beyrichiana)) plant association. This plant association is the "most typical" plant association in pine-oak sandhills [31,58,117]. In the western part of the gopher tortoise's range, threeawn is replaced by bluestem grasses (e.g., Schizachyrium scoparium and Andropogon spp.) in the ground layer [21,117]. Pine-oak sandhills occur on ridgetops and slopes of gently rolling hills, and their soils are deep, well-drained, and sandy. Pine-oak sandhills are fire-maintained communities that succeed to xeric hardwood hammock communities in the absence of fire or other disturbance. Xeric hardwood hammock communities are characterized as either scrubby, dense, low-canopied forests with little understory other than saw-palmetto, or multistoried forests of tall trees (e.g., pignut hickory (Carya glabra), sweetgum, and southern red oak) with variable canopy closure and an understory dominated by deciduous scrub oaks [2,58,117]. The ground layer of xeric hardwood hammocks is typically sparse and thus supports smaller gopher tortoise populations, particularly when the canopy cover is >60% [7]. The scarcity of herbaceous plants and the relatively incombustible oak litter preclude most fires in xeric hammock communities. When fire does occur, it is nearly always stand replacing (see Fire Regimes) [21,58,140].

Pine flatwoods: Gopher tortoises occur at high densities in pine flatwoods. These plant communities are characterized by an open canopy of widely spaced longleaf and/or slash pine with a sparse understory and a dense ground cover. Pine flatwoods occur on flat topography, and their soils are moderately to poorly drained, consisting of 1 to 3 feet (0.3-0.9 m) of sand overlying an organic hardpan or "clayey" subsoil [2,58,140]. Because pine flatwoods are more mesic than pine-oak sandhills, they typically support a more well-developed shrub layer [119] and relatively fewer grasses and forbs [20,140] than pine-oak sandhills. Oaks (e.g., sand live, laurel, water, and running (Q. pumila) oak), saw-palmetto, gallberry (e.g., inkberry (Ilex glabra) and large gallberry (I. coriacea)), lyonia (coastal plain staggerbush (Lyonia fruticosa) and fetterbush lyonia (L. lucida)), and wax-myrtle (Myrica cerifera) are common in the understory [2,119]. Like pine-oak sandhills, the understory and herbaceous ground cover associated with pine flatwoods vary across the gopher tortoise's range [68,140], but gopher tortoises frequently occupy pine flatwoods with a threeawn understory. Pine flatwoods are fire-maintained communities that succeed to mesic hardwood hammocks and other communities (e.g., bayheads) in the absence of fire or other disturbance (see Fire Regimes) [2,140]. Mesic hammock communities are dominated by evergreen oaks, saw-palmetto, gallberry, and ericaceous (Ericaceae) shrubs [2].

Scrub: Gopher tortoises occur at moderate densities in scrub, often in ecotonal or disturbed, open areas [9,16,31,60]. Scrub is dominated by various evergreen oaks (e.g., sandhill oak (Q. inopina), Chapman oak (Q. chapmanii), sand live oak, and myrtle oak (Q. myrtifolia)). Saw-palmetto, scrub palmetto, lyonia, and Florida rosemary occur in the understory [117]. Sand pine may or may not be present in the overstory. The tree and shrub canopies vary from sparse to dense depending upon the frequency of fire. The ground cover is "always sparse", species poor, and distributed in scattered open areas [117,140]. Soils of scrub are sandy and well drained [31,140]. Without fire or other disturbance, scrub may succeed to xeric hammock communities [2,132,140].

Nonforested habitats: Gopher tortoises may reach high densities in ruderal or heavily disturbed habitats. High gopher tortoise densities (8.9-9.9/acre) were reported in "remnants of natural habitat and had in every case a 20% to 60% cover of (nonnative) bahia grasses as well as a high diversity of broadleaf forage of over 160 species" [7]. Auffenberg and Franz [9] reported that "ruderal habitats may contain higher gopher tortoise population densities than original ones if grasses and forbs are abundant". In Orange County, Florida, gopher tortoises were absent from orange (Citrus sinensis) groves that were annually disced, but were relatively abundant in old fields (range: 1.1-2.5 gopher tortoise burrows/ha) and abandoned orange groves with dense herbaceous ground cover (range: 0.3-2.3 gopher tortoise burrows/ha) [74]. Across 24 counties in southern Alabama, mean gopher tortoise densities were highest in old fields compared to 7 other habitats [137]:

Average gopher tortoise densities in 8 habitats in 24 counties in southern Alabama [137]
Habitat Mean gopher tortoises/ha
Old fields 1.53
Pastures 0.50
Agricultural lands 0
Edges 0.64
Clearcuts 0
Pine plantations 0.72
Burned pine/scrub oak 0.64
Unburned pine/scrub oak 0

Among 7 habitat types at the Kennedy Space Center in Florida, gopher tortoise densities were highest in disturbed habitats. In this study, disturbed habitats were historically scrub habitats that were clearcut ≥20 years before the study [16,18]. The authors suggested that gopher tortoises might benefit from disturbed areas in scrub and slash pine flatwoods [16]. For more information on this study, see Population density.

Gopher tortoises may benefit from having a variety of plant communities in close proximity to each other. Wetland habitats (e.g., ephemeral ponds, freshwater marshes, and wet prairies) adjacent to other habitats may be used for feeding when seasonally dry or during drought years [7]. At the Kennedy Space Center, gopher tortoises preferred coastal scrub oak habitat to 6 other habitats for burrowing. No or few burrows occurred in swales; however, swales were used for feeding and many burrows were located adjacent to swales. These results suggested that gopher tortoises may require a range of well-drained to poorly drained sites for feeding and burrowing [16]. Other researchers report that pastures or mowed sites may be used for feeding when adjacent to other habitats. At the Kennedy Space Center, gopher tortoises rarely burrowed in disturbed areas, such as periodically mowed grassy areas, pastures, citrus groves, and firebreaks, but used these sites for foraging and burrowed along their ecotones [16,136].

BIOLOGICAL DATA AND HABITAT REQUIREMENTS

SPECIES: Gopherus polyphemus
  A gopher tortoise at its burrow in Naples, Florida. Photo courtesy of Charles Warren.
Many of the citations used in this section are reviews [7,8,9,10,39,44,45,54,105,156]. In addition to information from surveys performed at the Ashton Biological Preserve in Florida, Ashton and Ashton [7] provide an extensive review of gopher tortoise life history and habitat requirements throughout the gopher tortoise's range. They are cited frequently in the biological and habitat sections of this review. Ernst and others [54] review gopher tortoise morphology, life history, and habitat requirements. Their review is part of a larger collection of information on Testudines in the United States. Several reviews compare gopher tortoise biology with that of other North American tortoises [8,9,10]. Diemer [44,45] summarized gopher tortoise population status, habitat and ecology, and threats to survival based on literature available in the late 1980s. Reviews of gopher tortoise biology and habitat with a management focus were provided in these sources: [39,105,156].

LIFE HISTORY:
Daily activity: The gopher tortoise is diurnal, typically spending the night in its burrow and emerging from its burrow during the day to feed and bask (see Burrows) [54]. As an ectotherm, the gopher tortoise must use environmental conditions to maintain proper body temperature; thus, its daily activities (e.g., foraging, basking, and resting) are influenced by its environment and require suitable temperatures and/or humidities before initiation. If the temperature of the end chamber in the burrow is <55 °F (10 °C), then a gopher tortoise will typically remain inactive in its burrow. If the temperature of the end chamber in the burrow rises above 55 °F, then it typically becomes active and may exit the burrow chamber to bask at the burrow mouth [7]. Basking elevates its body temperature and is necessary for maintaining optimal efficiency of physiological processes such as digestion (Hutchison 1979 cited in [6]). The body temperature of a foraging gopher tortoise is generally above 70 °F (21 °C) but below 89 °F (32 °C). When its body temperature is too high, the gopher tortoise tends to seek shade or water and may or may not return to its burrow to cool down. It is less active if the air temperature is >90 °F (32 °C) or the soil temperature is >110 °F (43 °C) [7]. Thus, the gopher tortoise may exhibit a unimodal or bimodal daily activity pattern dependent upon local and regional weather patterns [9,51,103,154]. The gopher tortoise is also less active during conditions of high or extremely low humidity [7]. Adult and juvenile gopher tortoises exhibit similar daily activity patterns, although body temperature may fluctuate at a greater rate in small gopher tortoises due to greater surface area to volume ratios (MacDonald 1976, Rose and Judd 1982, cited in [154]). For more detailed information on daily activity and thermoregulation of the gopher tortoise, see Douglas and Layne [51].

Annual activity: Like its daily activity, the annual activity of the gopher tortoise is highly reliant upon environmental conditions. Annual activity patterns of gopher tortoises vary geographically, depending upon the severity of winter weather. Researchers divided the gopher tortoise's range into 2 subranges based upon annual activity patterns caused by latitudinal differences in weather. The northern subrange included gopher tortoises that were typically inactive through most of the winter months (January-March). This subrange included gopher tortoises occurring from southern South Carolina southwest to eastern Louisiana and east through northern Florida. The southern subrange included gopher tortoises that were typically active year-round and included those occurring south of the Santa Fe River in Florida [7]. During winter, gopher tortoises bask at the mouths of their burrows on warm days throughout their range [156], and gopher tortoises in all regions may be inactive for long periods (weeks to months) during cold or very hot and dry weather [7]. Even in places where gopher tortoises are active year-round, gopher tortoise activity is not uniformly distributed temporally. A peak in activity typically occurs during April through October, but particularly from April to June and September to October [10,55,152]. For more information on this topic, see Home range and movement.

Time of breeding and nesting vary throughout the gopher tortoise's range, depending upon latitude. Variations in air and soil temperature, forage availability, rainfall, and average number of sunny days affect gopher tortoise phenology [7]. Courtship behavior occurs principally in spring (April-early June), but also in late winter (February-March) and fall (October-November), particularly in the South [7,42,54,85]. Nesting generally occurs in spring through summer (late April-mid-July), with peak nesting activity occurring from mid-June to mid-July [54]. Nesting occurs only on warm, sunny days [85]. Hatching and emergence of young from the nest typically occur from August through October [7].

A gopher tortoise may spend as much as 80% of its time (or about 19 hours/day) in its burrow [7]. In mesic longleaf pine/pineland threeawn habitat at Ichauway in Baker County, Georgia, female and male gopher tortoises monitored during the day (when gopher tortoises are most active) were observed aboveground only 0.8% of the time [55]. Fourteen adult female gopher tortoises in pine-oak sandhills and old fields at the Katharine Ordway Preserve in north-central Florida were monitored during the day for up to 505 days. Female gopher tortoises were above ground <2% of the time. When active above ground, they spent most of their time basking at or near the apron (i.e., the mound of soil outside the burrow mouth formed during burrow excavation) (76%), walking (17%), and feeding (4%). Mating and nesting comprised 2% of their activities [135]. In pine-oak/saw-palmetto/pineland threeawn sandhills in Hillsborough County, Florida, juvenile gopher tortoises monitored during the day were active above ground 10% of the time [152].

Social interactions: Gopher tortoises are typically solitary [7] but live in loose aggregations of spatially clustered individuals with overlapping home ranges [103]. These groups are often referred to as "colonies". A colony may include >50 individuals [9]. Researchers further subdivided gopher tortoise colonies into subgroups, called pods, of highly aggregated gopher tortoises and their burrows. Interactions among individuals within a pod are presumably greater than those among pods, and interactions among pods within the same colony are presumably greater than those between colonies [7].

A dominance hierarchy may occur in gopher tortoise populations [54], and outcome of social interactions and distribution (i.e., home range, movement patterns, and burrow locations) of gopher tortoises may be based in part upon body size and gender ([10,42,103], Berry 1986 cited in [27], Douglass 1976 cited in [51]). Gopher tortoises may defend their burrows and feeding areas from other tortoises, particularly during the breeding season [7]. Territorial defense involves using aggressive and defensive postures, including combative charging and ramming [42,47]. Male movements and burrow preferences are mostly determined by reproductive and dominance behaviors. Adult males are often more aggressive, active, and wide ranging than females or younger age classes (e.g., juveniles and subadults) ([55], Berry 1986 cited in [27]). In coastal sand pine scrub at Savannas Preserve State Park in St Lucie County, Florida, male gopher tortoises spent significantly more time walking (P<0.001) and in social interactions (P<0.001) than females [42]. In longleaf pine-oak habitat at the Southlands Experiment Forest in Decatur County, Georgia, large (i.e., greater than the average carapace (i.e., upper shell) length) males defended burrow sites near adult females throughout the spring breeding season, apparently excluding small, subordinate males from breeding by preventing access to adult females [103]. At the Lochloosa Wildlife Management Area in Alachua County, Florida, a burrow occupied by a juvenile gopher tortoise was usurped by an adult male [47]. However, some studies found a relative lack of any social hierarchy influencing social interactions and distribution [13,27]. In slash pine-oak sandhills of Jasper County, South Carolina, burrow location was more heavily influenced by food supply and other habitat conditions than social interactions [158]. McRae and others [103] suggested that the formation of social hierarchies in the gopher tortoise population at the Southlands Experiment Forest may have been induced by increased density, pronounced social activities (e.g., courtship and aggressive interactions), and minimal dispersal during the spring breeding season.

Reproduction and development: Body size and age at first reproduction appear to vary by geographic location and local conditions [156]. Sexual maturity may be reached at carapace lengths of 7.1 to 15.2 inches (18.0-38.7 cm) (about 10-21 years old) for female gopher tortoises and 7.0 to 11.8 inches (17.7-30.0 cm) (about 9-18 years old) for male gopher tortoises [7,54].

Gopher tortoise eggs take approximately 60 days to develop in the uterus. Female gopher tortoises produce 1 clutch annually, although a given female may not produce a clutch every year. The number of eggs/clutch varies geographically [7]. Heavy rainfall during late summer and early fall (when follicular development begins in the female gopher tortoise) partially explained a significant decline in reproduction and clutch size in the subsequent year in a gopher tortoise population at the Fort Stewart Army Reserve [110]. Gopher tortoise clutches typically contain 5 to 8 eggs (range: 1-25 eggs). Clutch size is correlated to female carapace length, with larger females having larger clutches (r=0.346, P<0.05, [135]; r=0.68, P<0.01, [85]). The typical hatching time is 80 to 100 days. Eggs may not hatch together, and neonates may not emerge together. The entire period from hatching to emergence may occur over 1 to 20 days. The number of neonates surviving upon hatching and emergence varies with location (see Survival). Gopher tortoises do not provide parental care [7].

Temperature-dependent sex determination occurs in the gopher tortoise. Neonates are 1.3 to 2.2 inches (3.2-5.5 cm) long at hatching. The yolk sac is absorbed in 1 to 3 days, and neonates typically do not forage for 48 hours after final absorption of the yolk sac. A neonate is referred to as a hatchling once the yolk sac is absorbed and it begins to forage, typically 2 days to 2 weeks after hatching. Neonates, hatchlings, and juveniles have soft shells, whereas subadults and adults have hard shells. Differentiation between age classes may also be determined by an individual gopher tortoise's behavior and its carapace size, shape, and coloration. Sexual differentiation, using external characteristics such as concave plastrons (i.e., lower shell) of males, begins to become apparent in subadults and is distinctive in adults; in addition, adults demonstrate sexual behaviors such as courtship, mating, and territorial defense. Nomenclature for age classes is frequently inconsistent among studies, making comparisons difficult [7]. This review uses the nomenclature presented in studies and makes no effort to reconcile any potential inconsistencies. See Ashton and Ashton [7] for more information on this topic.

Growth: The rate of development of an individual gopher tortoise may be influenced by forage and water availability, age, and weather [7]. There is a positive linear relationship between gopher tortoise body size and age [10]. Gopher tortoises grow more rapidly before sexual maturity than after sexual maturity (Landers and others 1982 cited in [115]). Growth increments are greatest for gopher tortoises with carapace lengths from 4.7 to 7.5 inches (12.0-19.0 cm) (about 4-11 years old); thereafter, growth rate gradually decreases in both sexes. Adult female gopher tortoises often become larger than males [54]. Over 90% of the gopher tortoise's annual growth occurs from April to October [54], when gopher tortoises are most active (see Annual activity). Further information on growth can be found in Ernst and others [54].

Longevity: Gopher tortoises are long lived but difficult to age [7]. Researchers have speculated that gopher tortoises may live as long as 40 to 60 years [81,105] or as long as 150 years [39]. The oldest known gopher tortoise as of this writing (2009) was an 86-year-old captive male at the Nova Scotia Museum of Natural History [7]; however, because of high mortality in wild populations due to predation and disease, life spans of captive individuals may not be comparable to that of wild individuals [61].

Survival: Gopher tortoise survival rates vary by age class. Survival from the egg through the juvenile stage is low, largely because these age classes are extremely vulnerable to predation [7]. Low mean survival (5.8%) between the time of egg laying and 1 year of age was attributed to high predation rates in ruderal and longleaf pine-turkey oak habitats in northern peninsular Florida [3]. Auffenberg and Iverson [10] reported that "several" colonies in Alachua County, Florida, produced no young for as long as 7 years due to high predation of eggs. In longleaf pine-oak habitat at the Southlands Experiment Forest, 89% of nests were depredated within a few weeks after laying [85]. Based on information from Alford [3], the authors estimated that a female gopher tortoise in that population would produce a successful clutch (about 6 individuals) once every 10 years [85]. In slash pine-oak sandhills in Jasper County, South Carolina, 74% of eggs were destroyed by predators; 92% of these were destroyed within 2 weeks of laying [158]. Predation rates appear high regardless of whether nests are close to or remote from burrows [10,30,81,83,100,135]. For more information on this topic, see Nests.

If nests are not destroyed by predators, hatching success may be high. In longleaf pine-oak habitat at the Southlands Experiment Forest, 86% of eggs hatched in nests protected from predation [85]. In Duval County, Florida, hatching success of protected eggs was 81% [30]. In pine-oak sandhills and old fields at the Katharine Ordway Preserve, 67% to 97% of eggs at protected nests hatched [135]. Thirteen eggs excavated from Alachua County, Florida, had a 92% hatch rate when incubated in the laboratory [5].

Neonate, hatchling, and juvenile gopher tortoises are vulnerable to predation because they are soft-shelled, small, and rely on protection of their burrows for safety [152]. Gopher tortoises are particularly vulnerable during their first few days and weeks of life after emergence from the nest [10]. At Camp Shelby Training Site in southern Mississippi, 65% of hatchlings were killed within 30 days of hatching [53]. In slash pine-oak sandhills in Jasper County, South Carolina, the predation rate on hatchlings during the first year of life was 70% [158]. Juvenile gopher tortoises may be particularly susceptible to predation when at their burrows to thermoregulate during cool months of the year. In pine-oak/saw-palmetto/pineland threeawn sandhills in Hillsborough County, Florida, predation of juvenile gopher tortoises was higher in October and November (survival rate: 69%) and April and May (survival rate: 74%) than any other 2-month interval of the year (range: 89-100%) [153]. In addition to predation, survival of neonate gopher tortoises may be affected by soil compaction and clay content at the nest. A study of 40 nests in Forrest County, Mississippi, found that while most of the eggs hatched, "only about half" of the hatchlings survived; the others were apparently unable to dig their way out of the nest due to the hard-packed overburden [22]. Impacts by human land use also affect survival of young gopher tortoises. Landers and Buckner [83] reported that 6 nests located in roads and along firelines were destroyed by heavy equipment (see Mechanical site preparation). Predation on young gopher tortoises may influence their activity patterns, movements, and burrow use. For example, juvenile gopher tortoises appear to have narrower foraging and activity ranges than adults (see Foraging activity and Home range and movement), perhaps in response to high predation [153]. See Wilson [152,153] and Wilson and others [154] for more information on this topic.

Subadult and adult gopher tortoises are typically protected from predators by their hard shells [152]. Survival of subadults and adults is typically high and is mostly affected by human actions (see Status and threats). Thus, high levels of predation may occur in areas with heavy human use [7]. Over 15 months, survivorship of subadult and adult (6.7-11.1 inches (16.9-28.2 cm) plastron length) gopher tortoises at Archbold Biological Station in Highlands County, Florida, was 93%, whereas that of younger age classes (1.8-6.2 inches (4.6-15.7 cm) plastron length) was 45% [90]. In Jasper County, South Carolina, all adult deaths in a slash pine-oak sandhill population were attributed to direct morality due to collecting or indirect morality due to habitat destruction by humans [158]. Populations might be dominated by large individuals due to low recruitment in young age classes and high survival and longevity of subadults and adults, although many researchers could not discount the possibility of underdetection of young individuals (e.g., [3,112,130,137,158]). A study of 51 gopher tortoise populations throughout Florida indicated that populations residing in habitats that experienced severe area reduction (>25% reduction over 20 years), had >50% tree canopy, or were <5 acres (2 ha) in size tended to have few small or few large burrows (an indicator that these populations had few small or few large individuals). The authors surmised that large individuals leave these sites, and recruitment of young individuals into these populations is reduced [102].

Population density: Gopher tortoise densities may range widely among habitats but are typically highest in ruderal habitats and fire-adapted plant associations such as longleaf pine-oak sandhills, pine flatwoods, and sand pine-scrub (see Plant Communities) [9,44]. In general, gopher tortoise densities are positively related to herbaceous ground cover [18,28,44,77,86]. Throughout Florida, gopher tortoise densities in longleaf pine-turkey oak sandhills, xeric live oak hammocks, and sand pine scrub habitats increased as the percent cover of herbs increased (r²=0.61, P=0.005; [10,16]). In a variety of habitats on the mainland and on islands in Florida, gopher tortoise densities increased with increasing herbaceous ground cover (r=0.461) [115]. At the Kennedy Space Center, gopher tortoise density was highest in disturbed habitat and similar among scrub and slash pine habitats; herbaceous ground cover (excluding threeawn) was greater and shrub cover was less in disturbed than in slash pine and scrub habitats [16]:

Mean (SD) gopher tortoise densities in spring and fall in 7 habitats at the Kennedy Space Center in Florida [16]
 
Habitat
Disturbed* Palmetto scrub Scrub Slash pine/
palmetto
Slash pine Slash pine/
oak
Oak scrub
Gopher tortoises/ha
Spring 2.2 (4.8) 1.7 (3.4) 1.6 (3.2) 1.4 (2.7) 1.3 (2.7) 1.2 (2.7) 1.1 (2.6)
Fall 2.7 (4.9) 1.1 (3.0) 1.1 (2.9) 1.4 (3.2) 1.3 (3.4) 1.2 (4.0) 1.1 (2.6)
Habitat characteristics
Herb cover (%) 75.7 (22.7) 60.8 (24.8) 51.7 (28.7) 51.6 (28.1) 43.0 (29.4) 16.4 (12.9) 28.3 (25.4)
Nonthreeawn herb cover (%) 65.0 (28.6) 29.0 (27.9) 22.2 (26.2) 19.9 (19.6) 16.2 (18.5) 5.0 (7.1) 4.6 (4.5)
Canopy cover (%) 15.7 (27.6) 0 (0) 0 (0) 35.9 (23.6) 30.7 (23.1) 14.6 (11.5) 0 (0)
Oak cover (%) 27.7 (29.5) 23.2 (15.0) 38.0 (28.1) 22.4 (13.2) 36.6 (28.9) 80.5 (17.0) 76.3 (14.0)
Palmetto cover (%) 7.0 (14.0) 59.0 (21.2) 52.7 (23.9) 51.2 (22.4) 45.4 (22.5) 27.7 (10.8) 36.3 (23.6)
Shrub cover (%) 63.0 (28.7) 98.9 (2.5) 99.1 (2.3) 95.2 (8.9) 95.9 (8.0) 98.2 (3.4) 99.6 (1.4)
Shrub height (cm) 129.8 (75.7) 104.7 (34.4) 116.3 (56.5) 102.6 (34.5) 103.9 (36.0) 107.9 (42.0) 146.1 (87.2)
Depth to water table (cm) 83.6 (39.1) 46.6 (24.6) 57.7 (34.4) 69.0 (21.9) 82.8 (32.2) 125.6 (17.7) 86.6 (40.0)
*Disturbed habitats were scrub and clearcut ≥20 years before the study [18].

Seasonal shifts in habitat may occur in gopher tortoise populations. At the Kennedy Space Center, shifts may have occurred between fall and spring in some habitats (e.g., disturbed, palmetto-scrub, and scrub) but not in others (e.g., oak scrub, slash pine-oak, and slash pine-palmetto) [16]. In longleaf pine-oak habitat at the Southlands Experiment Forest, gopher tortoises moved from burrows in xeric sites in spring to burrows in mesic sites beginning in late summer. Greater availability of preferred foods by late summer in the mesic sites or a preference for moist burrows may have explained shifts in habitat use in this study [103]. In coastal scrub and slash pine flatwoods at the Kennedy Space Center, Breininger and others [16] found that cover of all herbaceous species (r= -0.18, P=0.07) and herbaceous species excluding threeawn (r= -0.26, P<0.01) was greater on poorly drained soils than on well-drained soils. These authors suggested well-drained soils and sunlit areas may be particularly important for gopher tortoise reproduction in spring, and poorly drained areas may be particularly important in fall because poorly drained sites have comparatively more available forage [16].

Because herbaceous ground cover is often negatively correlated with canopy cover and shrub cover, gopher tortoise densities tend to increase as tree and shrub cover decrease [9,16,28]. Increased canopy closure and litter buildup associated with fire exclusion can reduce herbaceous forage production and gopher tortoise numbers. For more information on this topic, see Succession.

Some locations may exhibit elevated or reduced gopher tortoise densities due to factors not related to habitat. High population densities may signal a stressed population rather than a healthy population, particularly if high densities are the result of gopher tortoises being confined [115,156]. Some of the highest reported densities are associated with real or effective "habitat islands". An isolated gopher tortoise population on Cape Sable in Everglades National Park, Florida, had a density of 11.3 gopher tortoises/ha. This population was located in xeric coastal prairie grassland with button sage (Lantana involucrata), crimson bluestem (Schizachyrium semiberbe), broomsedge bluestem (Andropogon virginicus), alena (Boerhavia repens), coconut palm (Cocos nucifera), cabbage palm (Sabal palmetto), and agave (Agave sp.). Presence of small burrows suggested that population recruitment was occurring [80]. An island population on the Egmont Key National Wildlife Refuge had 12.4 gopher tortoises/ha [115]. Comparisons of gopher tortoise populations on true islands with populations on the mainland indicated that gopher tortoises on islands may be forced to live in suboptimal conditions. Both island and mainland gopher tortoise populations showed a positive relationship between the number of burrows and the area of the habitat. On the mainland, gopher tortoise burrow density decreased as area increased. The ratio of inactive to active burrows (a measure of the tendency of individuals to construct new burrows) increased with area of habitat, and burrow density increased with increasing herbaceous vegetation. However, none of these relations could be demonstrated for gopher tortoises on islands. These findings suggested that gopher tortoises had a greater selection of habitats on the mainland than on islands [115]. A study of 51 gopher tortoise populations throughout Florida found that burrow abundance was negatively correlated with the area of available habitat (r= -0.46, P<0.01), possibly because gopher tortoises were concentrated in the only remaining suitable habitat [102]. The loss of preferred habitats may have increased gopher tortoise densities in habitats where gopher tortoises were previously less common [7]. In 1982, Auffenberg and Franz [9] indicated that gopher tortoises were not common in maintained pastures, but more recent studies (e.g., [159]) found that gopher tortoises are more commonly found in pastures and ruderal habitats as other available habitats are being lost or degraded [7]. Conversely, low population densities might not be indicative of less suitable habitat. Because of low recruitment, gopher tortoise densities may remain low for long time periods even if the cause of the initial decline (e.g., high rates of human predation) was removed. Measures of reproductive success and survival are important indicators of habitat suitability [146] and are frequently lacking in research addressing gopher tortoise habitat and density (e.g., [18]).

Differences in methodology make comparisons among population density studies difficult. Some studies have inferred gopher tortoise population densities from gopher tortoise burrow densities using a correction factor (e.g., [9,16,17,27,28,80,83,115,121]). Other studies have used mathematical relationships between the widths of burrow mouths and the sizes of gopher tortoises occupying them to construct age class distributions of gopher tortoise populations (e.g., [3,18,50,101,102,121,135,137,155]). Because of burrow use patterns—for example, use of several burrows by 1 individual and cooccupancy of a burrow by >1 individual—and the difficulty of detecting burrows of young gopher tortoises, some researchers contend that burrow densities or their dimensions are not likely to produce accurate estimates of gopher tortoise population density and structure [7,28,156]. In addition, the relationship between burrow density and actual gopher tortoise population density often varies with geographic location, predation pressure, habitat, season, and year [16,28,29,39,72]. For information on different methodologies used to capture gopher tortoises and determine burrow density and use, see the following reviews: [7,28].

Home range and movement: The gopher tortoise's home range is considered "well-defined" [7,10], and gopher tortoise movements are described as "extremely habitual" [54]. In general, older and larger individuals have larger home ranges than younger and smaller individuals [54]. Although females are often larger than males (see Reproduction and development), males tend to have larger home ranges [47,103], perhaps due to reproductive activity and dominance behaviors (see Social interactions). Adult female gopher tortoises typically travel less frequently and move shorter distances than adult males, and young gopher tortoises travel less frequently and move shorter distances than adults [10,55]. Home range of individual gopher tortoises tends to overlap greatly within and between sexes [42,103]. In coastal sand pine scrub at Savannas Preserve State Park, gopher tortoises overlapped home ranges but did not overlap core areas (i.e., areas of concentrated use) [42], suggesting that core areas may be defended from conspecifics.

There is considerable variation among individuals with respect to home range size and movements. Home range sizes range from less than 0.01 acre (0.004 ha) to more than 8 acres (3.2 ha). In longleaf pine-oak habitat at the Southlands Experiment Forest, home ranges of males were larger than those of females (P<0.01). Males had home ranges from 1.5 to 3.6 acres (0.6-1.44 ha), and females had home ranges from 0.1 to 0.2 acres (0.04-0.14 ha) [103]. In several habitats (i.e., mature slash pine plantation, grassy roadside strip, clearcut, and pasture) at the Lochloosa Wildlife Management Area, mean home range sizes were 2.17 acres (0.88 ha) for adult males, 0.77 acre (0.31 ha) for adult females, 0.12 acre (0.05 ha) for subadults, and 0.02 acre (0.01 ha) for juveniles [47]. In coastal sand pine scrub at Savannas Preserve State Park, male gopher tortoises had significantly larger home ranges than females (P<0.01); home ranges of male gopher tortoises ranged from 1.5 to 1.6 acres (0.62-0.64 ha), and home ranges of female gopher tortoises ranged from 0.8 to 0.9 acre (0.34-0.35 ha) [42]. In longleaf pine/pineland threeawn habitat at Ichauway, the average male home range was 2.7 acres (1.1 ha), and the average female home range was 1.0 acre (0.4 ha). Females in this study had significantly smaller home ranges than males (P<0.0001) [55]. At the Kennedy Space Center, home ranges of adult gopher tortoises encompassed 7 habitat types (i.e., oak scrub; oak palmetto scrub; palmetto/lyonia/wax-myrtle; mesic woody; swales; forest; and open, disturbed habitats). Males tended to have larger home ranges (x = 4.7 acres (1.9 ha)) than females (x =1.5 acres (0.6 ha)), though not significantly larger (P>0.05). Habitat use within gopher tortoises' home ranges was generally in proportion to the amounts of habitat available, except that gopher tortoises used swales and disturbed areas (e.g., grassy, mowed road shoulders, pastures, citrus groves, and firebreaks) less than expected based on availability [136]. Although gopher tortoise densities were low in swales and disturbed habitats, they were "heavily used by gopher tortoises for feeding" (Smith and Breininger personal observation cited in [136]). In pine-oak/saw-palmetto/pineland threeawn sandhills in Hillsborough County, Florida, the mean annual home range of juvenile gopher tortoises was 0.18 acre (0.072 ha) [152].

In general, home range sizes and distance of movements of gopher tortoises increase as forage quality and/or quantity decrease [7,10,45,103,113]. At the Katharine Ordway Preserve, home ranges of adult females ranged from 0.287 to 3.55 acres (0.116-1.435 ha) in pine-oak sandhills and 0.005 to 1.18 acres (0.002-0.479 ha) in old fields; differences between these habitats were attributed to more abundant and evenly distributed forage in old fields than sandhills [135]. In Florida, mean recapture radius (an estimate of home range size) of gopher tortoises in 3 habitats increased as the percent ground cover of herbaceous plants decreased (r²=0.83, P=0.012; [10,16]), an indication that gopher tortoises had to move farther to find food in habitats with fewer herbaceous plants [10]:

Mean herbaceous groundcover (%) and recapture radii (m) of gopher tortoises at 6 sites in Florida  [10]
Habitat Site Ground cover* Mean recapture radius**
Longleaf pine-turkey oak sandhills A 92 17.8
B 43 23.6
Xeric live oak hammock A 68 18.2
B 31 36.3
Sand pine scrub A 13 53.5
B 3 65.4
*All plant species combined.
**Sexes combined; n=312.

Reproductive behavior and seasonal availability of preferred forage may influence gopher tortoise movements and home range sizes. Although differences in methodology among studies make it difficult to compare results, in general, movements of adult gopher tortoises peak in spring (April-June), when gopher tortoises are reproductively active [10,103]. A secondary peak in movements occurs in late summer and fall (September-October), when local food supplies are reduced [103]. In slash pine-oak sandhills in Jasper County, South Carolina, adult gopher tortoises foraged at the greatest distances from their burrows and spent proportionately more time foraging during spring (April and May) and fall (October) than other times of the year [158]. In Alachua County, Florida, movements of adult males and females peaked during spring (May and June). Movement distances of adult males during this time were greater than those of adult females, presumably because adult male gopher tortoises moved farther to access the activity ranges of several females [10]:

Average monthly movements (m) of gopher tortoises in Alachua County, Florida from 1963 to 1969 (n=383) [10]
Carapace length (mm) Gender Jan-Feb Mar-Apr May-June Jul-Aug Sept-Oct Nov-Dec
100 Male 3.5 5.0 7.7 5.8 5.0 5.4
Female 4.3 4.7 6.5 5.9 4.7 5.6
101-200 Male 6.3 7.9 13.3 21.6 26.6 15.3
Female 9.2 8.8 15.2 18.9 27.1 17.8
201-300 Male 11.2 31.4 78.6 48.8 43.7 21.6
Female 13.1 22.5 61.5 44.2 51.8 38.1
>301 Male 10.6 41.3 69.3 46.7 45.5 32.3
Female 12.3 38.2 56.1 53.3 56.2 21.8

Herbaceous ground cover, an important food source for gopher tortoises, is typically less abundant in late summer and fall; home range sizes and movements may increase during this time [10,103,154]. In longleaf pine-oak habitat at the Southlands Experiment Forest, mean feeding radius of adult gopher tortoises for summer and fall (after 15 June) was larger than that for spring (before 15 June; P<0.10). The authors suggested that some preferred forage, such as legumes (Fabaceae), may be depleted around gopher tortoise burrows by this time [103]. Juvenile gopher tortoises in sandhills in Florida spent significantly more time foraging, traveled farther, and ate more plants as the number of seasonally available plants increased [116] (P<0.02 for all variables), indicating that juveniles moved farther to access preferred seasonal forage. For more information on this topic, see Foraging activity.

Seasonal movements of juvenile gopher tortoises may or may not be as pronounced as that of adults [10,103,154]. In Alachua County, Florida, young gopher tortoises typically moved shorter distances than adults and exhibited no consistent seasonal pattern. Gopher tortoises <3.9 inches (10.0 cm) in carapace length moved relatively small distances throughout the year and exhibited a small peak during May and June, and movements of gopher tortoises 4.0 to 7.9 inches (10.1-20.0 cm) in carapace length peaked in September and October [10]. In pine-oak/saw-palmetto/pineland threeawn sandhills at the University of South Florida Ecological Research Area in Hillsborough County, Florida, home ranges of juvenile (1-4 years old) gopher tortoises were significantly larger in summer (June-September) than in other seasons (P<0.05), and 89% of long-distance movements (>200 feet (61 m)) by juveniles were made during this time [154].

Because environmental conditions influencing gopher tortoises may not remain constant over many seasons and years, the area required for an individual over its lifetime is likely considerably larger than the area required over a single year. In longleaf pine/turkey oak/pineland threeawn habitat at the Fort Stewart Army Reserve in southeastern Georgia, Mitchell [110] found a significant positive correlation between average monthly rainfall and average monthly movements of gopher tortoises (r²=0.350, P=0.01), perhaps because burrow flooding prompted increased movements. Douglas (1986 cited in [47]) reported home ranges of 10.4 acres (4.2 ha) for a male gopher tortoise observed occasionally over a 5-year period in south Florida slash pine-turkey oak habitat and 16.6 acres (6.3 ha) for a male gopher tortoise observed over a 6-year period in an old field. In longleaf pine/turkey oak/pineland threeawn habitat at the Fort Stewart Army Reserve, average home range of adult female gopher tortoises was 1.94 acres (0.79 ha) in 2002 and 3.02 acres (1.22 ha) in 2003. The multiyear home range (2002-2004) was larger (5.60 acres (1.86 ha)) [110].

Long-distance movement: Gopher tortoises typically travel short distances within their home ranges but may infrequently travel long distances [7]. In several habitats at the Lochloosa Wildlife Management Area, adult male gopher tortoises moved 10 to 1,555 feet (3-474 m) between recaptures, adult females moved 0 to 610 feet (0-186 m), subadults moved 20 to 2,441 feet (6-744 m), and juveniles moved 10 to 325 feet (3-99 m) [47]. In longleaf pine/pineland threeawn habitat at Ichauway, the longest distance between subsequent locations was 1,942 feet (592 m) and 2,093 feet (638 m) by a female and male gopher tortoise, respectively, although the average distance traveled by female gopher tortoises was 177.4 feet (54.0 m)/move and that of males was 279.5 feet (85.2 m)/move [55]. Exploratory (i.e., nondispersal) movements as far as 5 miles (8 km) were recorded at the Ashton Biological Preserve in Florida [7]. Because of their "considerable mobility" [112], gopher tortoises may respond quickly to changes in local environmental conditions [121]. Gopher tortoises "rapidly colonized" an abandoned orange grove adjacent to gopher tortoise habitat in Orange County, Florida [74]. On the Ocala National Forest, gopher tortoises immigrated to burned longleaf pine-turkey oak within postfire year 1 [121]. For additional information on this study, see Fire Management Considerations. Large movements may occur over suitable or "apparently unsuitable" habitats (Diemer personal communication cited in [39]). Suggested reasons for large movements were foraging for seasonally available food resources (e.g., fruits, flowers, and seeds), breeding activity of males, nesting forays of females, seasonal migrations to and from overwintering sites, and departure from unsuitable habitat due to extremely dry or wet conditions, physical disturbance of burrows, or plant succession [7,9,10,83].

Dispersal by gopher tortoises is largely unstudied, and information is mostly based upon anecdotal observations. Dispersal apparently occurs within all age classes, although adults appear to disperse at a "proportionately much higher rate" than hatchlings. Hatchlings may or may not disperse from their natal areas. If dispersal occurs, movements away from the natal area may not take place for several years after hatching, and movements may be slow. In northern Florida, "the entire process leading to dispersal of at least 2 neonates from their natal areas took several years" [10]. In longleaf pine-oak habitat at the Southlands Experiment Forest, juvenile gopher tortoises rarely dispersed more than 49 feet (15 m) from the nest during the first year of life [103]. When hatchlings disperse from their natal areas, movements away from the natal area appear random. Dispersal movements of adults are apparently "more directed" and farther than that of hatchlings. Dispersal of adults, particularly males, is most likely to occur during the beginning of the reproductive period [10]. Total distance traveled by 2 dispersing, adult male gopher tortoises in longleaf pine/pineland threeawn habitat at Ichauway was 3 and 4 miles (5 and 6 km) [55]. In southwestern Georgia, 5 adult males emigrated >0.6 mile (1 km) from their colonies [86]. In several habitats at the Lochloosa Wildlife Management Area, the longest movement (0.56 mile (0.74 km)) of 22 adult, subadult, and juvenile gopher tortoises was made by an emigrating subadult [47]. McRae and others [103] suggested that subadult male gopher tortoises were more likely to disperse or be forced to the periphery of breeding areas than other age and gender groups; however, Eubanks and others [55] found no evidence that dispersal movements were tied to competition among males for mating opportunities.

Barriers to movement: Physical features such as cliffs, fissures, caverns, and sink holes [7,10], human infrastructure (e.g., roads, urbanized areas, and railroad tracks) [7,10,42], water (e.g., major rivers, lakes, ponds, and deep marshes), wind-erosion features [7], coarse woody debris [25], and dense vegetation [94,105] may be barriers to gopher tortoise movements. In young slash pine plantations on the Desoto National Forest in southern Mississippi, unburned windrows and associated stands of dense deciduous woody plants were barriers to gopher tortoise movements (see Windrows) [94]. Means [105] suggested that especially dense groundcover vegetation may impede gopher tortoise movements. Gopher tortoises at the Southlands Experiment Forest modified foraging movements near roads. Foraging areas were typically elliptical areas centered around the burrow entrance, but when a gopher tortoise was located along a road edge its foraging area was typically linear [103], perhaps avoiding the road. Bryant and Boykin [25] speculated that on the Desoto National Forest following Hurricane Katrina in 2005, "the amount of timber blown down could have restricted gopher tortoise movements for foraging and reproduction".

PREFERRED HABITAT:
Because gopher tortoise populations were reduced 80% between the late 1800s and late 1900s [9], an assessment of historical habitats, plant associations, and environmental conditions preferred by gopher tortoises is difficult. Current and historical human land-use practices have altered habitat throughout the gopher tortoise's range (see Status and threats), and "the habitats occupied by the gopher tortoise since human expansion within its range are not necessarily the 'preferred' ones but in many cases the only ones remaining that meet the gopher tortoise's needs" for survival [7].

Gopher tortoises are associated with a wide range of habitats within their range (see Plant Communities). Reviews state that within those habitats, gopher tortoises are consistently associated with well-drained, sandy soils that are suitable for burrowing and have deep (>2 feet (0.5 m)) groundwater tables (see Burrows), an open canopy (20-30% canopy closure) (see Cover Requirements), and an abundance of diverse herbaceous forage (see Diet). An open canopy provides sunlight necessary for high production of herbaceous forage and open, sunny sites required for nesting and basking [7,39]. Based on these associations, a review characterized "marginal", "moderately suitable", and "very suitable" gopher tortoise habitat throughout its range as follows [7]:

Characteristics of marginal, moderately suitable, and very suitable gopher tortoise habitat [7]
  Marginal Moderately suitable Very suitable
Habitat examples Scrub, coastal strands, rocky habitats Ecotonal areas, drying marshlands, live oak savannahs, open mesic hardwoods Sandhills, nonflooded flatwoods, open treed savannahs, old pastures
Soils very dry, sandy or rocky with thin soils seasonally dry to moderately wet moderately dry to very briefly seasonally wet
Number of plant species in 1 season 30-100+ 130-170+ 75-200+
Number of plant species in 4 seasons 10-210+ 170-375+ 150-400+
Canopy cover (%) <60 <60 <60
Herbaceous ground cover (%) 20-70 50-80 50-100
Preferred forage species (%)* 0-65 45-90 60-95
Plant species diversity** low moderate high
Forage species frequency of occurrence*** low moderate to high high
Seed source security**** low moderate to high high
Carrying capacity
(gopher tortoises/acre)
0.8 2-4 4+
*Preferred forage species occurred at a moderate, high, or very high frequency in the diet.
**High species diversity indicates a large number of equally or nearly equally abundant species, whereas low species diversity indicates either a small number of species of high abundance or a large number of species of low abundance.
***High frequency of occurrence indicates commonness, whereas low frequency of occurrence indicates rarity.
****Threats to seed source security include diseases, insect pests, overgrazing, and environmental pollutants. High seed source security indicates good forage sustainability.

COVER REQUIREMENTS:
Burrows: Burrows are an important aspect of gopher tortoise ecology. The burrow is a defendable refuge that protects the gopher tortoise from daily and seasonal temperature extremes, fire, and predators [7]. A gopher tortoise may spend up to 80% of its time in its burrow (see Annual activity), where it rests and defecates [54]. When active above ground, its activities (e.g., basking, foraging, and nesting) are centered around its burrow [103,135]. The temperature and humidity within the burrow remain relatively constant throughout the year, providing a consistent, suitable microclimate where temperature is moderated and humidity is held relatively high; thus, the burrow protects the gopher tortoise from lethal temperatures and prevents desiccation [7,39,51,54].

Burrow description: All age classes use burrows, although some hatchlings and juveniles may bury themselves in sandy soil or push under litter to create a depression or excavation directly below the soil surface called a "pallet". Gopher tortoises excavate new burrows, use the abandoned burrows of other gopher tortoises, or reconstruct damaged burrows [7,54].

The burrow is typically a single-channeled tunnel [105] that gradually slopes downward from the soil surface and levels off under ground [36]. The deepest part of the burrow is typically enlarged into a chamber [54]. The chamber is commonly placed close to the groundwater table and may be in soils different from the overburden (see Burrow sites). Burrows may be as long as 35 feet (10 m) and as deep as 10 feet (3 m) below the soil surface [7]. The length and depth of the burrow vary with the depth of sand and depth of the water table (Hallinan 1923 cited in [39]). Burrows of juvenile gopher tortoises are typically narrower, shorter, and shallower than those of adults, possibly making juveniles more vulnerable to predators [39]. Burrows are wide enough so that the gopher tortoise can turn around at any point along its length [36]. During burrow construction, excavated soil is deposited in a "half-moon shape" outside the burrow mouth to form the apron, which may be up to 7 feet (2 m) in diameter [7]. Its size and shape may be influenced by microtopography, gopher tortoise movements, and vegetation (Kaczor 1988 cited in [77]). The apron is often used as a basking site [8] and as a nesting site for female gopher tortoises (see Nests) [51].

Burrow sites: Where available, gopher tortoise burrows are typically located on well drained (rapid to moderate percolation rate), sandy soils where the groundwater table or impermeable clay or rock layer is at least 2 feet (0.5 m) below the soil surface. However, burrows may be dug in a variety of soils such as "shallow shelly soils", "heavy periodically flooded soils", and "rich loamy soils" [7]. In Louisiana, availability of sandy soils is "limited". In this region, most burrows are excavated in loamy soils with a "rather high" clay content, and the aprons are "usually compacted into a hard cement" [9]. The burrow may extend into different soils throughout its profile. The burrow's end chamber is often in soils "much less pervious than the overburden" [7]. Burrows dug in sandy soils are typically longer than those in clay soils, and hypoxia and hypercarbia may be more pronounced in burrows dug in clay soils (Ultsch and Anderson 1986, 1988, cited in [54]). In general, soils with high clay content do not support large gopher tortoise populations [9].

According to Ashton and Ashton [7], the most important soil characteristic in burrow site selection by gopher tortoises may be the distance between the soil surface and the groundwater table, with gopher tortoises typically placing the burrow chamber close to the groundwater table and at least 2 feet (0.5 m) below the soil surface [7]. The close proximity of the burrow chamber to the groundwater table may provide a suitable microclimate by keeping the humidity in the burrow high (>60%) year-round [7], but may subject the burrow to flooding when the water table moves upward during wet periods, particularly in winter [16,104]. In Leon County, Florida, and Thomas County, Georgia, water was observed in >100 burrows for periods lasting as long as 48 consecutive days, and several occupied burrows contained ground water at least once every year of the 4-year study [104]. In coastal scrub and slash pine flatwoods at the Kennedy Space Center, most burrows were flooded in winter. However, gopher tortoises still used them and "it was obvious that in winter gopher tortoises did not select for well-drained areas and that use of flooded burrows was common" [16]. Conversely, on Cape Sable, gopher tortoises commonly located burrows on upland ridges, but few burrows were located on beaches or swales. Beaches and swales were frequently used for foraging, however, suggesting that gopher tortoises "avoided digging burrows in areas subject to flooding" [80]. Mitchell [110] suggested that burrow flooding prompted gopher tortoises to abandon burrows in longleaf pine/turkey oak/pineland threeawn habitat at the Fort Stewart Army Reserve. It is unclear whether gopher tortoises prefer burrows that flood, but gopher tortoises apparently tolerate water in their burrows. In flooded burrows, gopher tortoises were found partially or completely immersed in water even when water levels were low enough in the burrow to avoid immersion, suggesting that gopher tortoises may prefer immersion and gain some thermoregulatory advantages from it [16,47,104].

Burrow site selection by gopher tortoises may be more heavily influenced by physical characteristics (i.e., soil type and proximity of the groundwater table) of the habitat than plant associations [7]. However, vegetation characteristics, including species composition and structure, apparently influence burrow site selection at least in part. Gopher tortoises may place burrows at sites where herbaceous ground cover, especially preferred herbaceous forage, is abundant [130,144]. A review states that gopher tortoises consistently select burrow sites where herbaceous ground cover is moderate to very dense (35-80%) [7]. In longleaf pine/turkey oak/pineland threeawn sandhills at George L. Smith State Park and Fort Stewart Army Reserve in Georgia, gopher tortoises selected burrow sites where percent cover of herbaceous plants was significantly greater than that available (burrow sites: 36-40%, random sites: 12-29%; P≤0.05) [130]. However, because selective foraging close to burrows may result in decreased abundance of herbaceous ground cover close to burrows, studies of the relationship of burrow sites to ground cover are confounded. In longleaf pine/pineland threeawn habitat at Ichauway, the abundance of some preferred forage (e.g., fleshy-fruited plants and nonthreeawn graminoids) was significantly lower near active gopher tortoise burrows than random sites (P≤0.05), whereas the abundance of other preferred forage (e.g., legumes) was more abundant near active burrows compared with random sites (P≤0.05). This author attributed the reduced abundance of some forage items next to burrows to selective foraging. Greater abundance of other forage items was attributed to gopher tortoise activities (i.e., creation of openings during burrow excavation and seed dispersal) that encouraged greater reproduction and growth of these plants near gopher tortoise burrows [13].

Gopher tortoises consistently select burrow sites where tree and shrub cover is low to moderate (range: 0-60%, "optimum": 20-30%) [7]. In longleaf pine/pineland threeawn habitat at Ichauway, average canopy cover at burrows (30%) was lower than at random points (60%; P=0.0001) [13]. In slash pine flatwoods and scrub habitats at the Kennedy Space Center, gopher tortoise burrows were associated with less canopy cover and lower shrub height than average conditions across the study sites [16]. At George L. Smith State Park, longleaf pine/turkey oak/pineland threeawn habitat that had not burned for >20 years had significantly lower canopy cover at active gopher tortoise burrows (26%) than at random points (76%; P≤0.05). In the same habitat, maintained by prescribed fire on 3- to 5-year intervals at Fort Stewart Army Reserve, canopy cover was also lower (26%) at active gopher tortoise burrows than at random points (40%; P≤0.05) [130]. Also in longleaf pine/turkey oak/pineland threeawn habitat at Fort Stewart Army Reserve, Mitchell [110] found low canopy cover both at burrows (23%) and at random points (22%), a result attributed to the uniformity of the habitat at her sites. Some researchers suggested that an open canopy near the burrow may provide basking and nesting sites, and may also provide sites for growth and reproduction of herbaceous forage [6,39,44,86,98,130,156]. However, gopher tortoise burrows are sometimes found in dense vegetation. Active gopher tortoise burrows were found in dense cogon grass (Imperata cylindrica) on the edge of sandhill habitat at the Withlacoochee State Forest in Florida [92]. Burke and Cox [28] located burrows in "extremely dense vegetation" in northern Florida, and Ashton and Ashton [7] located over 30 young gopher tortoises in burrows and pallets in a dense thicket of greenbriar (Smilax sp.), blackberry (Rubus sp.), blueberry, and grape (Vitis sp.) at another location in northern Florida.

Obstacles may also influence burrow site selection. Burrows are frequently located at the base of natural or manmade obstacles such as logs, rocks, plant roots, fences, and buildings [7].

Burrow use: The gopher tortoise uses one or more burrows within its home range. The number of burrows used by an individual varies geographically, seasonally, with the age and sex of the individual, and with habitat type, quality, and size [29,47,103,115]. Juvenile gopher tortoises typically use fewer burrows than adults, and adult males often use more burrows than adult females. In longleaf pine-oak habitat at the Southlands Experiment Forest, 2-year-olds used the greatest number of burrows [103]:

Mean number of burrows used from April to November by different age classes of gopher tortoise [103]
Age class (years) Burrows used
Neonate-1 1.1
2 2.2
4-5 1.7
6-9 2.0

In pine-oak/saw-palmetto/pineland threeawn sandhills in Hillsborough County, Florida, juvenile gopher tortoises used 4.4 burrows annually [152]. In several habitats at the Lochloosa Wildlife Management Area, juveniles used an average of 2.1 burrows from April to December, while subadults used 2.9 burrows, adult females used 2.7 burrows, and adult males used 5.5 burrows [47]. In 7 habitat types at the Kennedy Space Center, adult female gopher tortoises used an average of 8.8 burrows and adult males used an average of 16.6 burrows for 8 to 16 months. In this study, the number of burrows used by adult males was larger but not significantly different from that of adult females [136]. In longleaf pine/pineland threeawn habitat at Ichauway, male gopher tortoises used significantly more burrows (x =10.0 burrows) than females (x =5.2 burrows) during a 13-month period (P<0.0001) [55].

Although gopher tortoises may use multiple burrows, many burrows are consistently occupied by the same individual for months to years; others are used only briefly, abandoned, and then reopened by the same or another individual [7]. In pine-oak/saw-palmetto/pineland threeawn sandhills in Hillsborough County, Florida, juvenile gopher tortoises used an average of 4 burrows annually but used a single burrow 75% of the time [152]. There are typically more burrows in a given area than are used by gopher tortoises. A review states that "typical" occupancy rates of gopher tortoise burrows in Florida were between 47% to 49% in pine-oak sandhills and 32% to 67% in pine flatwoods [7]. Mushinsky and McCoy [115] found 55% of burrows were active at 19 sites located throughout Florida. To differentiate between burrows experiencing different use, many researchers classify gopher tortoise burrows as "active", "inactive", or "abandoned" [9,16,39,115,156]. Patterns of burrow occupancy are not thoroughly understood, but periodic occupation and abandonment of burrows often occurs seasonally [100]. In 7 habitats at the Kennedy Space Center, the average number of burrows used by adult gopher tortoises was highest for both sexes in March, August, and September. Burrow use decreased greatly during winter (December-February) [136]. In longleaf pine/pineland threeawn habitat at Ichauway, females used an average of 2 burrows each month from May through October and an average of 1 burrow in all other months. Males used increasingly more burrows each month from May (about 2 burrows) though September (about 5 burrows), followed by a decline in number of burrows used in October (about 3 burrows) and thereafter. Males used an average of 1 burrow from November to April [55]. In longleaf pine-turkey oak sandhills in Mobile County, Alabama, burrows used in spring and early summer were not used in late summer and vice versa [100]. In longleaf pine-oak habitat at the Southlands Experiment Forest, adult male and female gopher tortoises moved less and used fewer burrows during the spring breeding season than at any other time except late fall [103].

Seasonal patterns in burrow occupancy and abandonment may be influenced by environmental factors. In longleaf pine-oak habitat at the Southlands Experiment Forest, food depletion in spring breeding areas may have triggered burrow abandonment by adult gopher tortoises in late fall [103]. At the Katharine Ordway Preserve, adult female gopher tortoises moved between burrows 2 to 3 times more frequently from June to August in pine-oak sandhills than in old fields. Infrequent interburrow movements in old fields were attributed to more abundant, evenly distributed forage [135]. Other suggested reasons for differences in burrow use among populations were differences in ground cover, soil composition, temperature extremes at different latitudes, and number of disturbances to burrows [156]. For more information, see Home range and movement.

The gopher tortoise is typically solitary (see Social interactions); however, 2 or more gopher tortoises may occupancy a single burrow. In several habitats at the Lochloosa Wildlife Management Area, cooccupancy by 1 or more adult male gopher tortoises and an adult female gopher tortoise occurred at least 14 times from mid-May to mid-November, and cooccupancy of 1 burrow by 2 adult males or by 2 immature gopher tortoises occurred several times [47]. Cooccupancy occurred on 10 occasions at the Kennedy Space Center from March through November; a majority of dyads sharing the same burrow were the same sex [136]. Gopher tortoises may cooccupy burrows at night [47], but cooccupancy during the day may be more common [47,103]. Where burrow sites are limited, burrows may be more closely distributed and cohabitation may occur at higher rates. Young gopher tortoises appear to tolerate cohabitation of burrows more readily than adults and are sometimes found clustered together in small areas outside burrows [7]. At the Kennedy Space Center, 42% of burrows were used by more than one gopher tortoise at the same or different times during the course of a 16-month study conducted in 7 habitats [136].

Burrow distribution: Gopher tortoise burrow distribution within available habitat varies across sites and is poorly understood [39]. Many authors describe clusters of gopher tortoise burrows [3,9,10,103], while others demonstrate a random or regular distribution of burrows throughout an area [3,80]. In ruderal and pine-oak sandhills habitats in northern peninsular Florida, the burrow distribution varied from widely scattered, approximately uniform dispersion to distinctly clumped aggregations [3]. A review speculated that varying degrees of aggregation may reflect historical, sociological, or environmental differences among the areas surveyed or may simply reflect random variation in burrow placement [39]. Gopher tortoise burrows in longleaf pine-oak habitat at the Southlands Experiment

  Forest were aggregated. At this site, gopher tortoise movements were concentrated on a small area (about 2.5 acres (1 ha)) during the breeding season, when a few dominant males occupied the burrows closest to female burrows and excluded juvenile males from the immediate colony area. After the breeding season, gopher tortoises were more widely dispersed, shifting activity from the interior to the periphery of the colony [103]. In longleaf pine/pineland threeawn habitat at Ichauway, all burrows were clumped (Z=2.602, P<0.05), possibly reflecting the use of multiple burrows by a single individual. Active burrows were randomly dispersed (Z=1.225, P>0.05), possibly "related to the distribution of optimal habitat patches", which also appeared randomly dispersed. This researcher expected to find reproductive advantages for females in aggregations, but found no difference in the frequency of interactions, courts, or mounts between aggregated and isolated individuals [13]. She suggested that burrow distribution may be more closely associated with open canopy cover than with any particular social advantage. In habitats where gopher tortoises are forced into marginal areas or where suitable habitat patches are small and isolated, burrows may be clumped due to increased gopher tortoise density, a pattern noted for an island population in south Florida [80] and other sites throughout Florida [102].
A gopher tortoise burrow in "fire-maintained" longleaf pine forest in Georgia. Photo courtesy of Pete Pattavina, US Fish and Wildlife Service.

Nests: Gopher tortoise nests are typically 6 to 12 inches (15-25 cm) from the soil surface to the bottom of the nest [7]. Nests are located in open, sunny sites [7,54,83]. Most nests are located at or near the female gopher tortoise's burrow, but occasionally nests are located a considerable distance away in borrow pits [85], dirt roads, driveways, trails [49,83,85,100], firelines [83], clearcuts [47,49], a sand live oak thicket [49], forage strips planted for wildlife [85], gardens, lawns [39], and grassy fields [30]. In longleaf pine-oak habitat at the Southlands Experiment Forest, most nests (85%) were located within 6 to 32 inches (15-80 cm) of the burrow mouth. Thick herbaceous cover and roots were thought to restrict nesting to burrow entrances and man-made clearings at this site [85]. However, in pine-oak sandhills and old fields at the Katharine Ordway Preserve, <2% of aprons contained nests, and only 11% of monitored females deposited eggs at burrow entrances. At this study site, mean distance of nesting forays was 253.9 feet (77.4 m) [135]. At 2 sites in longleaf pine-turkey oak sandhill habitat in Mobile County, Alabama, all nests occurred in aprons at one site, whereas only 29% of nests occurred in aprons at another site [100]. Anecdotal information suggests that nesting away from the burrow may be a response to poor nesting conditions, such as little sunlight, at the burrow [6,81,83,86]. Females may also nest in the aprons of inactive or abandoned burrows. In pine-oak sandhills and old fields at the Katharine Ordway Preserve, 78% of nests were located in active burrows, 19% were located at inactive burrows, and 3% were located at abandoned burrows [135]. In slash pine-oak sandhills in Jasper County, South Carolina, 74% of nests were found at a female's apron, 9% were at aprons of abandoned burrows, and 17% were at sandy sites remote from a burrow [158]. Smith [135] speculated that active burrows may be preferred over inactive or abandoned burrows because the actions of the resident female may keep the apron clear of vegetation and debris and keep the soils less compacted. Uniquely, McRae and others [103] reported that in longleaf pine-oak habitat at the Southlands Experiment Forest "it was not uncommon for females to temporarily relocate or even excavate a new burrow... for the purposes of egg-laying".

FOOD HABITS:
Diet: The gopher tortoise is an herbivore [54]. The bulk of its food is grasses (70-80% of total intake by volume). The young shoots, stems, and roots of herbaceous plants, and seasonally available forage from woody plants such as flowers, fruits, and leaves from vines, shrubs, and trees, comprise much of the remainder [7]. Gopher tortoises may consume dry grasses, pine needles, and broadleaf tree leaves (MacDonald 1986 cited in [39]). Lichens, mosses, fungi, insects, carrion, rocks, bones, charcoal, and feces of other animals may also be consumed; these foods typically do not make up a substantial portion of the gopher tortoise's diet [7] but may be an important resource, especially for conditioning females prior to and shortly after egg laying [59]. Gopher tortoises may obtain some water from food [59] but drink water when it is available [54].

Gopher tortoises eat many herbaceous plants. A review states that throughout the gopher tortoise's range, >1,100 plant species are known to serve as forage. Forage availability differs throughout the gopher tortoise's range and depends upon environmental conditions. An individual gopher tortoise may consume up to 160 to 400 plant species. Forty-five of the 83 families of plants consumed by gopher tortoise were identified in a review as providing "critical" forage for the gopher tortoise throughout its range [7]:

Important gopher tortoise forage [7]
Common name Family Some genera used as forage
Important most of the year
Carrot Apiaceae Apium, Daucus, Eryngium, Hydrocotyle, Centella, Ptilimnium
Palm Arecaceae Serenoa, Sabal
Aster Asteraceae Ambrosia, Artemisia, Krigia, Lactuca, Hieracium, Helenium, Solidago, Bidens, Conyza, Aster, Gnaphalium, Liatris, Carphephorus, Chrysopsis, Coreopsis, Circium, Eupatorium, Emilia, Erechtites, Elephantopus, Helianthus, Heterotheca, Hieracium, Hypochaeris
Midsorus fern Blechnaceae Belchnum, Woodwardia
Cactus Cactaceae Opuntia, Cereus, Hylocereus
Manosteen Clusiaceae Hypericum
Sedge Cyperaceae Bulbostylis, Cyperus, Fimbristylis, Kyllinga, Rhynchospora, Scleria
Fern Dennstaedtiaceae Pteridium
Pea Fabaceae Acacia, Aeschynomene, Amphicarpaea, Apios, Arachis, Astragalus, Baptisia, Canavalia, Cassia, Centrosema, Chamaecrista, Chapmannia, Clitoria, Crotalaria, Dalea, Desmodium, Galactia, Indigofera, Lespedeza, Lupinus, Macroptilium, Medicago, Melilotus, Mimosa, Phaseolus, Pisum, Pueraria, Rhynchosia, Senna, Stylosanthes, Tephrosia, Trifolim, Vicia, Vigna, Zornia
Grass Poaceae Amphicarpum, Andropogon, Axonopus, Bouteloua, Bromus, Cenchrus, Chloris, Cynodon, Dactylis, Digitaria, Dichanthelium, Echinochloa, Eragostis, Eremochloa, Eustachys, Muhlenbergia, Panicum, Paspalum, Poa, Schizachyrium, Setaria, Stenotaphrum, Sporobolus, Tridens
Purselane Portulacaceae Portulaca
Rose Rosaceae Agrimony, Amelanchier, Crataegus, Eriobotrya, Photinia, Potentilla, Prunus, Rubus
Madder Rubiaceae Diodia, Ernodea, Galium, Hedyotis, Richardia, Spermacoce
Secondary or seasonal importance
Agave Agavaceae Polianthes, Nolina, Yucca, Sansevieria
Amaranth Amaranthaceae Amaranthus, Iresine, Froelichia, Alternanthera, Blutaparon, Gomphrena
Sumac Anacardiaceae Schinus, Rhus, Mangifera, Toxicodendron
Custard apple Annonaceae Asimina, Annona
Mustard Brassicaceae Lepidium, Brassica, Raphanus, Descurainia, Capsella, Draba, Polanisia, Warea
Butterfly bush Buddlejaceae Polypremum
Goosefoot Chenopodiaceae Atriplex, Chenopodium, Salicornia, Salsola
Coco plum Chrysobalanaceae Chrysobalanus, Licania
Spiderwort Commelinaceae Commelina, Cuthbertia, Murdannia, Tradescantia
Gourd Cucurbitaceae Coccinia, Cucurbita, Melothria, Momordica
Yam Dioscoreaceae Dioscorea
Heath Ericaceae Vaccinium, Gaylussacia
Spurge Euphorbiaceae Acalypha, Chamaesyce, Cnidoscolus, Croton, Euphorbia, Phyllanthus, Stillingia
Beech Fagaceae Quercus
Mallow Malavaceae Abutilon, Alcea, Hibiscus, Malva, Modiola, Sida
Meadow beauty Melastomataceae Rhexia
Mulberry Moraceae Morus
Primrose Onagraceae Gaura, Oenothera, Ludwigia
Woodsorrel Oxalidaceae Oxalis
Passion flower Passifloraceae Passiflora
Plantain Plantaginaceae Plantago
Buckwheat Polygonaceae Antigonon, Coccoloba, Eriogonum, Polygonella, Polygonum, Rumex
Figwort Scrophulariaceae Agalinis, Aureolaria, Buchnera, Gratiola, Linaria, Penstemon, Seymeria, Verbascum, Veronica
Smilaz Smilacaceae Smilax
Nightshade Solanaceae Solanum, Lycopersicon, Petunia, Physalis
Vervain Verbenaceae Stachytarpheta, Lantana, Lippia, Phyla, Stylodon, Verbena, Callicarpa
Grape Vitaceae Ampleopsis, Cissus, Parthenocissus, Vitis
Important but of low biomass
Acanthus Acanthaceae Ruellia, Dyschoriste, Elytraria, Stenandrium
Borage Boraginaceae Heliotropium, Lithospermum, Myosotis, Onosmodium
Morning glory Convolvulaceae Cuscuta, Dichondra, Stylisma, Convolvulus, Ipomoea, Evolvulus
Turnerna Turneraceae Piriqueta
Violet Violaceae Viola

Grasses and legumes appear particularly important in the gopher tortoise's diet ([59], MacDonald 1986 cited in [39]). In longleaf pine-slash pine/oak sandhills in Hillsborough County, Florida, 68 genera and 26 families of plants were identified in the scats of gopher tortoises of mixed ages and genders. The most frequent components in the diet were: Poaceae (particularly threeawn, 98% of scats), Pinaceae (particularly pine (Pinus sp.), 84% of scats), Fagaceae (79% of scats), Fabaceae (65% of scats), and Asteraceae (60% of scats). Gopher tortoises in this study did not exclude any common plant genus from their diet [98]. The most common genus ingested was Aristida, but when compared with availability, gopher tortoises randomly selected or avoided it. Dominant plants in the Poaceae and Asteraceae families and the Pityopsis genus were also randomly selected or avoided [98]. In young slash pine/bluestem and mature longleaf pine/bluestem habitats on the Desoto National Forest, gopher tortoises of unknown age and gender ate mostly grasses (Poaceae), particularly crabgrass (Digitaria sanguinalis) and panic grass (Panicum sp.) [94]. In slash pine-oak sandhills in Jasper County, South Carolina, gopher tortoises of mixed ages and genders commonly consumed threeawn, prickly-pear (Opuntia sp.), broom sedge (Andropogon sp.), and fennel (Eupatorium sp.) [158]. In longleaf pine-oak habitats at the Southlands Experiment Forest, plants in the Poaceae, Cyperaceae, and Asteraceae families were the principal foods of adult gopher tortoises throughout the active season (80% of fecal contents). Adult gopher tortoises ate pineland threeawn, particularly in the early spring and late fall when forbs such as Florida pusley (Richardia scabra) and poorjoe (Diodia teres) were least available. Legumes (e.g., sensitive brier (Schrankia microphylla), hoary pea (Tehrosia spp.), milk pea (Galactia spp.), and butterfly pea (Clioria mariana)) were used extensively, especially by juveniles [59]. Ashton and Ashton [7] speculated that a diverse diet appears critical for the overall health of gopher tortoises. These authors suggested that "moderately suitable" and "very suitable" gopher tortoise habitats were those that had at least 100 species of native plants, of which >50% were available to gopher tortoises as forage throughout the active season [7]. See Preferred Habitat for additional information.

Legumes appear particularly important in the diet of juveniles [10,59,81,103]. Compared to adults, young gopher tortoises tend to ingest more legumes and other forbs, fewer Poaceae, and fewer plants with thorns or other external defense mechanisms (e.g., Rubus and Cnidoscolus) [59,98]. In longleaf pine-oak habitats at the Southlands Experiment Forest, legumes were used extensively by juveniles [59]. In pine-oak/saw-palmetto/pineland threeawn sandhills in Hillsborough County, Florida, juvenile gopher tortoises avoided Aristida and other Poaceae and selected Liatris and Dyschoriste when compared with availability. Grasses were generally consumed during months when forbs were not readily available (October-December) [116]. Young gopher tortoises, particularly neonates, have higher metabolic rates, protein demands, and evaporative losses than adults [112], so they may ingest plants accordingly. However, nitrogen content of ingested genera was not correlated with the probability of a plant being ingested by juvenile gopher tortoises in pine-oak/saw-palmetto/pineland threeawn sandhills in Hillsborough County, Florida [116].

Threeawn may be seasonally or irregularly consumed [7]. Gopher tortoises frequently select threeawn right after fire, when it is one of the first grasses available. However, threeawn is apparently not selected as forage when legumes and other forbs are available. In longleaf pine-oak habitats at the Southlands Experiment Forest, consumption of threeawn was highest in the early spring and late fall when other species were unavailable [59]. In longleaf pine-slash pine/oak sandhills in Hillsborough County, Florida, young gopher tortoises did not ingest threeawn [98]. In slash pine-oak sandhills in Jasper County, South Carolina, threeawn comprised the greatest biomass available to gopher tortoises and also formed the largest portion of their diet [158], but no statistical comparison between use and availability was provided. Mushinsky and others [116] indicated that in a study of juvenile foraging, pineland threeawn was avoided when compared with its availability. Threeawn is an important component of gopher tortoise habitats (see Fire Regimes) [12,35], but threeawn presence alone may be a poor indicator of habitat suitability for gopher tortoises. In Florida, a population decline of 30% occurred when threeawn ground cover went from 40% to nearly 95% [7]. For an annotated list of 83 families, 376 genera, and 1,103 species of plants used as gopher tortoise forage, see Ashton and Ashton [7].

Foraging activity: Gopher tortoises select forage species seasonally and according to plant phenology [7,116]. Although gopher tortoises may consume plants in all growth stages, most forage is selected when it is actively growing. Gopher tortoises may consume dry grasses, pine needles, and broadleaf tree leaves from late fall through early spring, when most plants are senesced and other foods are unavailable [7]. Critical periods for gopher tortoises are early spring—a time of recovery from winter dormancy and when nutrients are needed for egg formation—and fall, when food plants are fibrous and less available and gopher tortoises are preparing for winter dormancy. Seasonal shifts in forage help maintain protein and mineral intake. As grasses became more fibrous in summer and fall than in spring, gopher tortoises tend to shift to higher-quality forage such as legumes and ripe fallen fruits [59].

Gopher tortoises are restricted to foraging near the ground. Adults can typically access forage items up to 12 inches (30 cm) above the ground; forage items <4 inches (10 cm) above the ground are typically accessible to hatchling and juvenile gopher tortoises [7,105]. Many species of groundcover plants grow out of reach of gopher tortoises as the growing season advances [105].

Gopher tortoises typically feed close to their burrows. In Alachua County, Florida, they usually fed <160 feet (50 m) from their burrows [10]. In longleaf pine-oak habitat at the Southlands Experiment Forest, mean feeding radius of adult gopher tortoises was 43 feet (13 m) from their burrows. Mean feeding radius of juvenile gopher tortoises was 25.6 feet (7.8 m) [103]. All feeding activity of adult female gopher tortoises in pine-oak sandhills and old fields at the Katharine Ordway Preserve occurred within 56 feet (17 m) of their burrows [135]. Mean feeding radius of juveniles in pine-oak/saw-palmetto/pineland threeawn sandhills in Hillsborough County, Florida, was 26.2 feet (8.0 m) [154]. However, gopher tortoises may travel long distances to forage. Some gopher tortoises at the Ashton Biological Preserve traveled 2 miles (3 km) to feed on legumes [7]. Ashton and Ashton [7] state that grasses, which make up a bulk of the gopher tortoise's diet, are normally near the gopher tortoise's burrow (i.e., within 328 feet (100 m) diameter of the burrow), whereas forbs, which make up much of the remainder of the gopher tortoise's diet and are often only seasonally available, are often much farther from the burrow (i.e., as far as 3,281 feet (1,000 m)). See Home range and movement for additional information on this topic.

Foraging sites: Mesic and xeric habitats and their ecotones, including all of the sites associated with burrows (see Burrow sites), serve as foraging sites [7]. Additional sites include wetlands [16], annually tilled fields, periodically mowed grassy areas [16,60], dense thickets [7], and firebreaks [16]. Because foraging sites may differ from burrow sites, "solely protecting gopher tortoise burrow habitat may not preserve adequate resources needed to support a population" [136]. For more information on this topic, see Plant Communities.

SOURCES OF MORTALITY:
Predators: According to reviews, predators of gopher tortoise eggs, neonates, hatchlings, and juveniles are northern raccoon (Procyon lotor), common gray fox (Urocyon cinereoargenteus), Virginia opossum (Didelphis virginiana), striped skunk (Mephitis mephitis), spotted skunk (Spilogale putorius), domestic dog (Canis familiaris), domestic cat (Felis catus), bobcat (Lynx rufus), coyote (Canis latrans), wild boar (Sus scrofa), nine-banded armadillo (Dasypus novemcinctus), snakes (e.g., eastern diamond-backed rattlesnake (Crotalus adamanteus), eastern indigo snake (Drymarchon couperi), coachwhip (Masticophis flagellum), racer (Coluber constrictor), common kingsnake (Lampropeltis getulus), and Florida cottonmouth (Agkistrodon piscivorus conanti)), native fire ants (Conomyrma spp.), and nonnative fire ants (Solenopsis spp.). Red-tailed hawks (Buteo jamaicensis), red-shouldered hawks (B. lineatus), bald eagles (Haliaeetus leucocephalus), American crows (Corvus brachyrhynchos), and blue jays (Cyanocitta cristata) also prey upon hatchling and juvenile gopher tortoises. Gopher tortoises have also been found in the scat of the Florida black bear (Ursus americanus floridanus) [99].

Larger gopher tortoises have few predators except humans, but subadults and adults may occasionally be killed by domestic dogs, domestic cats, coyotes, bobcats, wild boars, northern raccoons, hawks, and eagles. Nine-banded armadillos may cause indirect mortality of gopher tortoises by entombing them during excavation of burrows for their use as hibernation dens [10]. For additional information, see Survival.

Humans: Humans are the leading killer of adult gopher tortoises [7,54]. Gopher tortoises may be sold in the pet trade, killed on roads by vehicles, or slaughtered for food. Taylor [143] has made an extensive study of human predation on gopher tortoises in north-central Florida. For additional information, see Status and threats.

Parasites and diseases: Gopher tortoises of all ages may have ticks (e.g., Amblymma spp. and Ornithodorus turicata), botflies (Cistudinomya cistudinis), and other parasites, which may be debilitating or lead to the death of the individual. Stressors, such as extremes of temperature and lack of moisture, may make gopher tortoises susceptible to disease. A review of infectious diseases and pathogens of gopher tortoises, including upper respiratory tract disease, herpes virus, and iridovirus, is provided in Ashton and Ashton [7].

MANAGEMENT CONSIDERATIONS:
Status and threats: From the late 1800s to the late 1990s, the gopher tortoise's range contracted, and populations declined an estimated 80% [9]. Gopher tortoise population decline is attributed to habitat destruction, habitat degradation, and human predation. Habitat destruction affecting the gopher tortoise includes urbanization, phosphate strip-mining, sand extraction, road building, and dam construction. Conversion of longleaf pine forests to other habitats, fire exclusion from longleaf pine habitats, establishment and spread of invasive species, and increased predation have contributed to habitat degradation for the gopher tortoise. Collection of gopher tortoises for food and pets, vehicular mortality, removal as "pests" from fields with livestock or agricultural crops, and gassing during rattlesnake (Crotalus and Sistrurus) round-ups have also contributed to a reduction in gopher tortoise numbers [7,9,44,45,48,81,86,151,156].

Perhaps the single largest contributor to gopher tortoise population decline was the historical reduction in the extent and quality of longleaf pine ecosystems (reviewed by [13]). As of 1985, area of longleaf pine-dominated forests was reduced by approximately 86% [106]. Those forests that remained were often fragmented or of poor quality for gopher tortoises, largely as a result of forest management such as conversion of large areas to pine plantations and fire exclusion [9,21,58,151]. Habitat loss and degradation have fragmented many gopher tortoise habitats, resulting in small and discontinuous gopher tortoise populations in some areas (see General Distribution). Habitat fragmentation may limit the gopher tortoise's ability to immigrate into or emigrate away from isolated areas, thereby potentially restricting the expansion of colonies, limiting the establishment of new colonies, or reducing a colony's ability to replenish its numbers or diversify its gene pool [100]. Fragmentation may also increase gopher tortoise predation by humans, domestic dogs and cats, and urban-associated wildlife (e.g., northern raccoons), or force individual gopher tortoises into unsuitable habitats and onto roads [100,156]. Low reproductive rates, delayed maturation (see Reproduction and development), and high mortality (see Survival) may make it difficult for gopher tortoises to recover from population declines and habitat changes [39,44,59,81,85,93].

Ecological significance: Because gopher tortoises may increase local animal and plant diversity, the gopher tortoise has been characterized as a keystone species [52,67]. Gopher tortoises excavate burrows and build aprons; burrows may be large (see Burrow description) and last as long as 10 years following abandonment [68,71]. Burrow creation produces small gaps in the understory vegetation, which may increase habitat heterogeneity and promote plant species richness and diversity by exposing mineral soil favorable for germination of early-successional species [77,144]. Gopher tortoises may affect the flora around their burrows and the surrounding habitat by ingesting and subsequently dispersing seeds ([138], Boglioli and others. 2000, Birkhead 2001, cited in [12]). In pine-oak/saw-palmetto/pineland threeawn sandhills in Hillsborough County, Florida, gopher tortoise aprons had lower concentrations of some soil nutrients and organic matter, higher pH and light intensities, and greater daily temperature fluctuations than adjacent undisturbed areas. Temperature maxima during prescribed fires were significantly lower on aprons than in surrounding undisturbed vegetation at sites burned annually and at sites burned every 5 years (P≤0.05 for all variables), suggesting that gopher tortoise aprons "may function as cool-temperature refugia during fire for seeds, perennating organs, or established plants" [77].

Because gopher tortoise burrows provide habitat for other species, gopher tortoise presence may increase local animal diversity. Many other animals seek temporary shelter or live permanently in gopher tortoise burrows. Jackson [75] reported over 60 vertebrate and 300 invertebrate species found in gopher tortoise burrows. Some commensals, including beetles (Coleoptera), crickets (Orthoptera), and mites (Arachnida), have not been found in habitats other than active gopher tortoise burrows, where they feed on gopher tortoise feces (see [157] and [75] for annotated lists). Other commensals, such as the Florida mouse (Podomys floridana) and gopher frog (Rana capito), may utilize gopher tortoise burrows whether a gopher tortoise is present or not. Foxes, skunks, bobcats, coyotes, nine-banded armadillo, and Florida burrowing owls (Athene cunicularia floridana) may use abandoned gopher tortoise burrows to rear their young or as hibernation dens [7,10]. Several species associated with gopher tortoise burrows are listed as endangered, threatened, or species of special concern by the US Fish and Wildlife Service, including the gopher frog, eastern indigo snake, Florida pine snake, and Florida mouse [39]. Some researchers speculated that a decline in gopher tortoise populations could trigger declines in other animal species in the same ecosystem [120]. Several reviews covering this topic are available: [7,39,54].

Succession: Gopher tortoises tend to occur at high densities in midsuccessional stages of fire-maintained longleaf pine ecosystems, which have a "savanna-like" structure, early-successional stages of sand pine scrub, and ruderal habitats. Low gopher tortoise densities tend to occur in advanced successional stages such as hardwood hammocks with dense canopies (see Plant Communities) [9]. Habitats associated with the gopher tortoise are subject to frequent disturbance by fire, tropical storms, insects, and pathogens [21,87,126]. At the Kennedy Space Center, disturbed habitats had nearly twice as many gopher tortoises as 6 other habitats, partially because disturbed sites generally had less oak and shrub cover but higher nonthreeawn herbaceous cover [16]. Without disturbance, open habitats become more shaded as tree and shrub canopies close, and gopher tortoise forage becomes less available [9,31,58,144]. Lemon [91] recognized over 35 plant species found in longleaf pine forests that are abundant during the first few years following fire but decline after 8 years. Edmisten (1963 cited in [16]) found that herb frequency declined as canopy cover increased after fire in pine flatwoods in northern Florida. While threeawn declined at an even rate, several forbs declined earlier and more rapidly. In addition to reducing herbaceous forage cover, excessive shade may prevent gopher tortoises from reaching minimum thermal requirements for daily activities (see Daily activity) [115]. As shade increases and foraging, nesting, and basking sites decrease, gopher tortoises become less common [9]. Without frequent disturbance, plant succession may cause gopher tortoises to emigrate from an area within 3 to 8 years [6,9,45,86]. In Lake County, Florida, gopher tortoises declined 1,800% over 11 years as abandoned agricultural land succeeded to mature sand pine scrub. These authors provided the following account of this time period [9]:

"During the first year the flora of the habitat is predominantly composed of annuals; the specific composition varies considerably due to the importance of chance colonization. Though gopher tortoises occur in these pioneer communities, colonies are never formed, and population densities are generally very low. During the second and third years, dog fennels (Eupatorium capillifolium) gradually increase and tortoise population densities remain low; by the third year, broom sedges (a common food of gopher tortoise) become a conspicuous element. Population density increases and tortoise colonies become established from the 4th to the 10th years, and young trees of species commonly associated with disturbed situations (such as persimmon (Diospyros spp.), hawthorn (Crataegus), and chickasaw plum (Prunus angustifolia)) are abundant. Tortoise nests often become common for the first time, and populations usually prosper for several years. Populations will remain at high levels if the area is subjected to recurrent fire, pasturing, or any other factor that tends to reduce the growth and density of taller tree species in the succeeding years. Without such modification, the period after 11 years is one during which larger native trees begin to shade out and otherwise compete with ruderal species, and tortoise densities are generally reduced in accordance with the rate and patterns of shade production."

In Covington County, Alabama, Aresco and Guyer [6] found a relationship between level of permanent abandonment of burrows and increase in midstory and overstory canopy closure in slash pine plantations that were winter-burned every 3 to 4 years. In this study, active burrows were located in areas with the most open canopy, and abandoned burrows were associated with a dense midstory of young hardwoods (e.g., oaks, dogwoods, and gallberry). Burrows were abandoned at a rate of 22%/year over 5 years. Potential long-term effects of fire exclusion that may adversely affect the gopher tortoise include increased canopy closure due to the expansion of understory and midstory hardwoods, reduced cover and richness of herbaceous plant species used as forage, changes in composition of the dominant species from fire-adapted to less fire-tolerant species, and increased litter [20,62,79]. Changes in forest structure and buildup of litter may increase fuel loads and support a more homogeneous, "hotter" fire, which may have a detrimental effect on gopher tortoises [62,113]. For more information on this topic, see Fire Effects and Management.

Habitat Management: Habitat management for the gopher tortoise is largely focused on reducing overstory trees and understory shrubs to increase the amount of light reaching the forest floor and on improving the quality and quantity of herbaceous forage [6,7,44,86,98,116,130,141,156]. Management to reduce canopy closure and woody understory vegetation in habitats associated with gopher tortoises include prescribed fire, tree harvest, mechanically treating understory shrubs, grazing, mowing, and herbicides [21]. Depending upon ecological conditions, a combination of treatments is sometimes recommended (e.g., [6,43,73]). Because fire provides many ecosystem services that other management methods may not, fire is frequently cited as being "more beneficial" to gopher tortoises than other methods [39]. For example, fire may release nutrients, scarify seeds for germination, reduce litter, and increase sunlight on the ground by removing groundlayer, understory, and overstory vegetation (see Indirect Fire Effects) [19,39]. However, because of concerns for safety and air quality, fire may be impractical in some areas [19,131], and mechanical methods may be used to keep habitat open and sustain plant species diversity [7,21]. Intensive mechanical site preparation and chemical treatments may be detrimental to gopher tortoises, and some researchers recommend that these techniques be avoided [43] or that lower-impact alternatives be selected [21,83,86].

Many undesirable effects of management activities on gopher tortoises and their habitats may be overcome by timing treatments so gopher tortoises are in their burrows and removal of seasonally important foods is avoided [82]. Depending on local and regional weather patterns, gopher tortoises may exhibit a unimodal or bimodal daily activity pattern (see Daily activity) [51,103,154]. In spring, gopher tortoises typically forage in midday and afternoon. At this time, gopher tortoises eat young growth from a diversity of species including grasses. In summer, gopher tortoises tend to bask and forage in early morning and in late evening, while staying in their burrows during the hottest part of the day. During summer, plants may be stressed due to dry, hot conditions and gopher tortoises may travel to low, moist areas to forage (see Foraging sites and Home range and movement). In fall, gopher tortoises often forage in midday and afternoon and stay in their burrows on cool days. At this time, gopher tortoises often forage upon ripe fallen fruits and legumes. In winter, gopher tortoises forage only at midday if the weather is sunny and warm, but during periods of cold weather gopher tortoises may stay in their burrows for several days or weeks. Gopher tortoise feed on legumes and cool-season herbaceous plants in winter [7]. Because gopher tortoises are relatively inactive, winter is often the "best" time for management activities [7]:

Best and worst times for management activities for the gopher tortoise based upon gopher tortoise's daily and annual activity patterns in areas south of the Suwannee River, Georgia [7]
Activity Best Dates Best Time of Day Worst Dates Worst Time of Day
Prescribed fire June-July or December-January any time September-October*, November-December, or February-April none
Tree harvesting December-February any time March-November midday
Mowing December-August before 10 AM in summer, before 10 AM and after 4 PM in winter, or on cool cloudy days September-November* after 10 AM in summer and between 10 AM and 4 PM in winter
Mechanical site preparation (e.g., plowing and fireline cutting) December-February before 10 AM March-November 10 AM to 4 PM
Plant surveys April-October any time November-March after dark
Burrow surveys after fire**, September-November, or April-June 1 hr after sunrise to 1 hr before sunset December-March or July-August when visibility is low or during fog or rain
*Hatching occurs during these months.
**Burrows are often highly visible after fire.

Because gopher tortoise diet and activity patterns differ throughout its range, these authors caution that timing of management activities is likely different throughout the species' range, and that management activities should take into account local site conditions and characteristics of local gopher tortoise populations. The authors recommended monitoring habitat and populations before and after treatments to determine whether management objectives, such as improving gopher tortoise forage, are met [7,16].

Fire: For information on the effects of fire on the gopher tortoise, see Fire Management Considerations.

Silviculture: A wide variety of silvicultural techniques may be compatible with maintaining or increasing gopher tortoise populations when applied at appropriate times and at intensities, scales, and frequencies that gopher tortoises can tolerate. Silvicultural practices favorable to gopher tortoises in longleaf pine forest include uneven-aged forest management involving selective thinning of overstory and midstory trees [7,9,43,83,85,93,94,156,158]. Uneven-aged management is often promoted for gopher tortoise habitats because it allows much of the forest structure to be retained between harvests [149]. Because longleaf pine regeneration is largely confined to canopy gaps (Wahlenberg 1946 cited in [21]), selective group thinning that creates canopy gaps may facilitate its regeneration [43,57,149]. Wilson and others [156] recommended natural regeneration of longleaf pine over planting because planting and associated soil disturbance may adversely impact the groundlayer vegetation and thereby negatively impact gopher tortoises. In longleaf pine forest, group selection may be compatible with gopher tortoise management by reducing canopy cover, thinning dense oaks in the understory, stimulating productivity of herbaceous forage, and creating suitable nesting and burrowing sites [7,9,44,72,83,86]. However, selective timber harvesting may be detrimental to gopher tortoises if tree and shrub cover exceeds 60% during the rotation [7,156], if concurrent management activities such as mechanical site preparation adversely impact grasses or other herbaceous species [156], or if harvesting equipment occludes gopher tortoise burrows (see Mechanical site preparation). In Florida, selective cutting of longleaf pine-oak habitat had varying effects on gopher tortoises. On average, selective cut longleaf pine-oak habitat at 5 sites in Florida showed an increase in density of 0.5 gopher tortoises/ha [9]:

Effect of management techniques on gopher tortoise density in 3 habitats in Florida [9]
Habitat type and location Management technique Years between data collections* Change in gopher tortoises/ha
Longleaf pine-oak
Alachua County Clearcut 8 -3.63
Selective cut 5 0.79
Selective cut 5 2.57
Selective cut 5 -1.61
Columbia County Slash pine planting 3.5 -4.92
Selective cut 7 0.84
Levy County Selective cut 6 -0.27
Pasture 3.3 -1.75
Sand pine-scrub oak
Lake County Clearcut 5 -0.77
Selective cut 5 -1.06
Xeric hammock
Levy County Pasture 7.5 0.22
Pasture 6 -0.07
*Data from more than 8 years were excluded because of successional effects.

A combination of even-aged management and prescribed fire is frequently recommended to maintain or improve gopher tortoise habitats (e.g., [6,43,73,107]). Farrar [57] suggested that since soil disturbance is "minimal compared to other methods", a group selection system including cyclical prescribed fire "as an integral component" may be compatible with gopher tortoise conservation. A study of the effect of stand thinning and summer burning in slash pine plantations on the Conecuh National Forest, Alabama, demonstrated that plant species richness (P=0.0041) and percent cover of groundlayer plants (<4 inches (10 cm) tall) available to gopher tortoises at active burrows increased on treated sites compared with untreated sites (P=0.02) [73]. In general, management for uneven-aged stands of longleaf pine regenerated naturally and maintained with fire is recommended over other management methods for maintaining gopher tortoise populations [105]. See Fire Management Considerations for additional information.

Silvicultural practices that appear less favorable to gopher tortoises in longleaf pine forest include clearcutting, particularly removal of longleaf pine and conversion to slash, sand, or loblolly pine plantations [9,83,85,93,94,158]. Pulliam (1987 cited in [73]) commented that the greatest threat to the gopher tortoise is habitat management that favors a closed canopy of young pines, such as that produced in pine plantations. Clearcutting is likely most detrimental if it damages pine seedlings and herbaceous cover [149]. However, gopher tortoises may occupy pine plantations if the canopy is sufficiently open to allow growth of abundant herbs and provide nest sites in nearly full sunlight [59]. For a short time after clearcutting, a pine plantation may provide open and grassy areas suitable for gopher tortoises [9,72,156]. In a 2-year-old clearcut that was mechanically prepared and planted to slash pine at the Lochloosa Wildlife Management Area, gopher tortoises frequently excavated burrows in windrows created during clearcutting operations (see Windrows), and 1 female used the clearcut to nest [47]. Gopher tortoise populations may decline on pine plantations once the canopy closes and groundcover vegetation decreases. Unthinned pine plantations may develop a closed canopy 10 to 15 years after clearcutting [6,9]. Auffenberg (cited in Thomas 1978, cited in [94]) reported that gopher tortoise populations in longleaf pine areas in Florida declined 60% to 100% within 8 years of clearcutting and replanting to slash pine. Limited information from 2 sites in longleaf pine-oak and sand pine-scrub oak habitats in Florida suggested that clearcutting and planting slash pine reduced gopher tortoise densities by 0.8 to 4.9 gopher tortoises/ha [9]. Landers and Buckner [83] reported that gopher tortoises at the Southlands Experiment Forest were found at "moderately high" densities in seedling (3 years old) and pole stages (burned every 1-2 years) of slash pine plantations but at low densities in dense, unburned sapling-stage plantations. This pattern was attributed to the small amount of herbaceous ground cover in unburned sapling stage plantations. In slash pine-oak sandhills of Jasper County, South Carolina, "a few" gopher tortoises migrated from logged sites into adjacent unlogged sites. One year after logging, gopher tortoise populations were reduced, and 66% of juveniles and 32% of adults were not recaptured in the logged area. The author speculated that many of these individuals were probably killed within collapsed burrows or emigrated out of the study site [158]. In pine plantations, forest management recommendations to benefit gopher tortoises include using low-intensity site preparation [83,86] and selective tree harvesting so trees are widely spaced or there is a mosaic of treeless or sparsely treed openings [105]. Planting and retaining fire-tolerant species such as longleaf pine, maximizing edge where low-growing herbs may be more abundant [59], and conducting frequent (1- to 4-year intervals) prescribed fire are also recommended [45,83,86].

Although some pine plantations may support dense gopher tortoise populations [43,83,86], many researchers recommend that pine plantations be restored to longleaf pine forest to return historical fire, hydrologic, and nutrient dynamics to these communities [6,21,86]. Selective tree harvesting to retain longleaf pine and thin slash pine, loblolly pine, and other pines and hardwoods from the overstory has been suggested to help restore longleaf pine forest [6,43,156]. Clearcutting and planting longleaf pine has also been suggested as a means to convert loblolly or slash pine plantations to longleaf pine forest [43,149]. For more information on restoring longleaf pine forests, see Fire Management Considerations.

Mechanical site preparation: Mechanical site preparation techniques (e.g., roller chopping, anchor chaining, and web plowing) may reduce understory shrub cover and increase herbaceous ground cover in gopher tortoise habitats in the short term [2]. In slash pine/gallberry-wax-myrtle flatwoods in Florida, herbaceous diversity increased substantially after clearcutting, stump removal, burning, windrowing, discing, bedding, and slash pine planting. In posttreatment year 2, crown cover of previously dominant pines and shrubs was reduced from 151% to 12% of surface area; grasses (e.g., bluestem, threeawn, and panic grasses) decreased from 30% to 23% cover, rushes (e.g., Juncaceae) and sedges (Cyperaceae) increased from 3% to 6% cover, and forbs (e.g., aster (Aster spp.), buttonweed (Diodia spp.), dogfennel (Eupatorium spp.), deergrass (Rhexia spp.), and violet (Viola spp.)) increased from 3% to 21% cover [37]. At the Southlands Experiment Forest, herbaceous biomass and gopher tortoise densities on 16-year-old slash pine plantations that had been anchor chained and roller chopped were comparable to those on unaltered longleaf pine-oak habitat. In part, this was attributed to frequent prescribed fire and removal of pineland threeawn [86], which is sensitive to soil disturbance (review by [117]). For more information on this study, see Direct Fire Effects. Mechanical site preparation and removal of threeawn from gopher tortoise habitats may result in short-term increases in cover of some gopher tortoise forage species but also of "undesirable weedy colonizers" (review by [2]). In the long term, disturbance of the soil and removal of threeawn may adversely affect gopher tortoises by altering the fire regime. Fire-adapted groundcover vegetation such as threeawn in unaltered longleaf pine forests may help to carry surface fires, and removal of this native ground cover by mechanical means may impair fire spread [35,140]. Once threeawn and other fire-adapted groundcover plants are removed, restoring the ground cover and fire regime may be difficult, possibly requiring planting and seeding herbs in addition to prescribed fire [6,14,117,119].

Because mechanical site preparation involves soil disturbance, it may adversely affect gopher tortoise habitats by encouraging the establishment and spread of nonnative invasive species (see Invasive species) [2,35,149]. On Pebble Hill Plantation in southern Georgia and northern Florida, old fields that had succeeded to shortleaf pine (Pinus echinata) and loblolly pine had approximately 50% lower species richness and greater abundance of nonnative species than nearby, fire-maintained longleaf-shortleaf pine sandhills that had never been plowed [123]. Plant species composition was different between the habitats. Longleaf-shortleaf pine sandhills sites had high grass cover and relatively lower forb and woody species cover than old fields, which had minimal grass cover and high cover of large-statured forbs and woody plants. The longleaf-shortleaf pine sandhills habitat contained many herbaceous plants that were not typically present in old fields. Many of these plants are important in the gopher tortoise's diet [7], including Maryland golden aster (Chrysopsis mariana), Curtis's spurge (Euphorbia curtisii), summer spurge (E. discoidalis), hairawn muhley (Muhlenbergia capillaris), switchgrass (Panicum virgatum), and Virginia tephrosia (Tephrosia virginiana) (see Diet) [123].

Mechanical site preparation can be destructive to plants and soil and cause greater vegetation changes than prescribed fire. Prescribed fire in pine flatwoods reduces the cover of woody understory species and litter, and increases the frequency, biomass, and diversity of herbaceous species; however, the composition of woody species generally changes "only slightly", and woody species diversity is generally unaffected by prescribed fire. In contrast, mechanical site preparation tends to change woody understory species composition and relative abundance and may reduce woody species cover substantially. In addition, differential responses of herbs such as threeawn to mechanical site preparation change herb species composition and relative abundance (review by [2]). In sand pine scrub in Orange and Oceola counties, Florida, vegetation recovered more quickly on burned sites through sprouting and rapid release of nutrients into the soil (rather than slow release through decay) and stimulation of fire-adapted reproduction than vegetation on roller-chopped sites [128]. Conde and others [37] provided a review that compared the short-term (≤2 years after treatment) effects of prescribed fire and mechanical site preparation on plant community structure in slash pine flatwoods and found that mechanical site preparation had a "similar but more pronounced" effect:

A review of vegetation response to prescribed fire and mechanical site preparation in slash pine flatwoods [37]
Plant community information
Treatment
February prescribed fire** February prescribed fire, clearcutting, double-chopping, and slash pine planting** Clearcutting, chopping, bedding, and slash pine planting*** Clearcutting, May prescribed fire, and very intensive site preparation*
General description tree-dominated woody- to herb-dominated woody- to herb-dominated woody- to herb-dominated
Woody plant response marginal reduction substantial (10 times) reduction substantial (10 times) reduction severe (100 times) reduction
Herbaceous plant response marginal, uniform increase substantial, uniform increase substantial, variable increase highly variable increase
Number of species marginal increase increase increase unchanged
Species diversity marginal increase substantial increase substantial increase delayed increase
*Very intensive site preparation included stump removal, windrowing, discing, bedding, and slash pine planting [37]. Prior to treatment, the site had not been burned in 40 years [38].
**Prior to treatment, the site had not been burned in "many" years [111].
***Prior to treatment, the site had not been burned in 40 years [38].

Breininger and others [16] recommended that, if mechanical site preparation is included in a habitat management plan, it should retain enough groundcover vegetation to carry fire. Some researchers suggested that removal of some of the shrub component from a site by mechanical or other means is likely to enhance gopher tortoise habitats by reducing canopy cover and increasing light on the forest floor, but caution that because predation on juveniles may be greater in large open areas than in areas with shrub cover, retaining some shrub patches may be beneficial for juvenile gopher tortoises [16].

Gopher tortoises may be killed or maimed by heavy machinery associated with mechanical site preparation, and heavy machinery may occlude gopher tortoise burrows and kill gopher tortoises that are unable to dig themselves out [7,9,39,94,158]. Landers and Buckner [83] reported several occasions of direct mortality from heavy equipment. An adult gopher tortoise was decapitated by a disc pulled along a fireline, a hatchling gopher tortoise was killed by a disc in another area, 4 young (<4 years old) gopher tortoises were dug up and crushed by a road grader, and 6 nests laid in roads and along firelines were destroyed by heavy equipment. Heavy equipment frequently occludes gopher tortoise burrows, but its effect on resident gopher tortoises appears to vary by soil conditions and type of machinery used. The majority of studies indicate that gopher tortoises can dig out of collapsed burrows following site preparation on deep, sandy soils. The potential for gopher tortoises to be entrapped in clay soils is unknown but may potentially be greater [9,44,47,48,81,83]. In dry prairie habitat in Desota County, Florida, roller chopping reduced the canopy cover more than web plowing, but roller chopping occluded more burrows (72%) than web plowing (49%). None of these burrows were reopened following occlusion, and newly excavated burrows on nearby untreated sites indicated that at least some emigration occurred from treated sites to untreated sites [142]. After site preparation on the Southlands Experiment Forest, 7 of 11 gopher tortoises present before site preparation left the treated sites [83]. Occlusion of gopher tortoise burrows during mechanical site preparation may make fewer burrows available to gopher tortoises and cause individuals to relocate more frequently and create or use additional burrows ([48], Douglass 1990 cited in [46]). Although gopher tortoises may emigrate from mechanically treated sites, use of such sites soon after treatment has been documented. Diemer and Moler [48] found hatchling gopher tortoises on a site that had been recently roller chopped. Nests and burrows of young gopher tortoises, which are often shallow, may be particularly susceptible to damage by mechanical site preparation; however, such effects had not been well studied as of this writing (2009). Wilson and others [156] recommended that heavy machinery be excluded from a 25-foot (7.6 m) buffer around known burrows to avoid burrow occlusion and other disturbance to resident gopher tortoises.

Windrows: After harvesting pine stands, slash may be piled in windrows, which may be burned. Lohoefener and Lohmeier [94] suggested that unburned windrows may be barriers to gopher tortoise movements in young slash pine plantations on the Desoto National Forest. At this site, dense stands of deciduous woody plants (e.g., oaks, flowering dogwood (Cornus florida), persimmon, yaupon (Ilex vomitoria), grape, and greenbriar) growing out of unburned windrows inhibited gopher tortoise movements and divided areas occupied by gopher tortoises into "islands" [94]. However, unburned windrows were frequently burrow sites of juvenile gopher tortoises in a 2-year-old clearcut slash pine plantation at the Lochloosa Wildlife Management Area [47]. According to Landers and Buckner [83], windrow burning had no apparent adverse effect on gopher tortoises at the Southlands Experiment Station.

Grazing: As of this writing (2009), limited information was available on the effect of livestock grazing on gopher tortoises. Grazed pastures may support large gopher tortoise populations when adjacent to habitats with more herbaceous plant species diversity [9,47], suggesting that livestock grazing and gopher tortoises may be compatible. However, conversion of gopher tortoise habitat to pasture likely reduces diversity of gopher tortoise forage [7]. Anecdotal evidence suggests that cattle grazing in gopher tortoise habitats (e.g., xeric hammock, pine-oak sandhills, and sand pine-scrub oak) may maintain open canopies and encourage forage species for gopher tortoises, as long as these habitats are not overgrazed [7,9,84]. Historically, gopher tortoises were probably the primary grazers in upland xeric habitats [81]. Because it is a grazer, the gopher tortoise may potentially compete with other grazers for food. Livestock grazing may kill pine seedlings and prevent the natural regeneration of pine in longleaf pine-turkey oak sandhills habitat and indirectly affect gopher tortoises by altering the successional trajectory of the habitat [74,84]. Livestock trampling may occlude gopher tortoise burrows [47], consequently killing resident gopher tortoises if they become entombed.

Mowing: Limited information on the effect of mowing on gopher tortoises was available as of this writing (2009). Mowed areas are frequently used by gopher tortoises (see Plant Communities). Mowing in gopher tortoise habitats may reduce shrub cover and maintain herbaceous plants. It removes less digestible parts of plants and stimulates new, more digestible and accessible growth [7]. Means [105] suggested that mowing can be used as "a substitute for fire, or in combination with it, especially in the early stages of reclaiming overgrown gopher tortoise habitat". Ashton and Ashton [7] suggested that mowing at least once every 2 years, at a time when gopher tortoises are least active, may benefit the gopher tortoise. However, Means and Grow [106] suggested that mowing is not an adequate substitute for fire because unlike fire mowing "does not stimulate the flowering response of groundcover grasses and forbs" and "indiscriminately culls young trees and plants".

Herbicides: Herbicides are frequently used in gopher tortoise habitats to reduce hardwoods and thus increase growth of herbaceous forage for gopher tortoises [14,23], although the effects of herbicides on gopher tortoises themselves are not known.

Herbicides are often viewed as an attractive alternative to prescribed fire to reduce understory hardwoods, particularly due to concerns over smoke management, air quality, and litigation [131]. Some researchers suggest that herbicides may offer potential advantages over prescribed fire, including increased selectivity and decreased risk of off-site damage (Hunter 1990 cited in [131]). However, Brennan and others [19] argue that herbicides cannot duplicate the multiple ecosystem benefits provided by fire. Herbicides may reduce understory vegetation, but unlike herbicides, fire may release nutrients, scarify seeds for germination, reduce understory litter, increase sunlight on the ground by reducing understory, midstory, and occasionally overstory layers, and encourage fire-tolerant plants [19,21]. Several researchers recommend herbicides in conjunction with fire [19,23] or to facilitate the eventual use of prescribed fire [21,43] for managing gopher tortoise habitats. DeBerry and Pashley [43] suggest that herbicides may be particularly useful where hardwood and shrub encroachment is excessive or where a site has been degraded by invasive species, which may proliferate after fire. Where chemical treatments are used to selectively reduce undesirable plants, Wilson and others [156] recommended that soil disturbance, destruction of groundlayer vegetation, and nontarget effects of herbicides be minimized.

The short- and long-term effects of herbicide use on gopher tortoises were unknown as of this writing (2009), so herbicides should be used cautiously in gopher tortoise habitats [131]. In loblolly pine sandhills habitat with a diverse hardwood understory (e.g., oaks, black tupelo (Nyssa sylvatica), sweetgum, sassafras (Sassafras albidum), tree sparkleberry (Vaccinium arboreum), hackberry (Celtis occidentalis), wax-myrtle, and winged sumac (Rhus copallina)) in Marion County, Georgia, gopher tortoises continued to use an area after herbicide treatment [23]. However, herbicides were suspected of killing gopher tortoises in at least one instance (Dobie personal communication cited in [44]).

Other Management Considerations:
Invasive species: Nonnative invasive plants may threaten gopher tortoise habitats by displacing native species, disrupting nutrient and fire cycles, and altering plant succession [24,92]. For example, cogon grass is considered a threat to gopher tortoises because it crowds out native plants and increases fire severity by increasing fuel loads [43,92] (see the FEIS review of cogon grass) . Nonnative invasive plants, such as kudzu (Pueraria montana var. lobata), air yam (Dioscorea bulbifera), and morning glory (Ipomoea spp.), may reduce the availability of native forage species and threaten long-term gopher tortoise population viability by reducing diversity of grasses and other herbaceous species. Brazilian pepper (Schinus terebinthifolius) and melaleuca (Melaleuca quinquenervia) appear particularly problematic in gopher tortoise habitats because they "often out-compete local forage species" or "shade out forage habitat". However, a review states that "managers should carefully weigh the goals for native species restoration with the need for sufficient quality forage for gopher tortoises". Some nonnative pasture grasses, such as bahia grass, centipede grass (Eremochloa ophiuroides), and St Augustine grass (Stenotaphrum secundatum), are "excellent" forage for gopher tortoises and may support high gopher tortoise densities. Kudzu, air yam, and morning glory may provide "a large supply of biomass" to gopher tortoises and are "fairly nutritious". Thus, removal of some nonnative plants may reduce the diversity and quantity of forage species, which may be detrimental to some gopher tortoise populations [7]. For more information on this topic, see Ashton and Ashton [7].

Relocation: When implemented appropriately, relocating gopher tortoises may augment populations and mitigate negative impacts of development, fragmentation, or other disturbances [7,54,95,156]. Consideration of the social structure of the resident population and the movements of relocated individuals may increase the likelihood of success [7]. Relocation of gopher tortoises to areas where they have been eradicated (i.e., rehabilitated forest and mine lands or areas where they have been eradicated due to fire exclusion) has had mixed results, and in many cases relocated gopher tortoises abandon sites to which they have been relocated [27]. Other considerations when relocating gopher tortoises include disease transmission and the genetic relationships of the relocated and resident gopher tortoises [7]. For more information on gopher tortoise relocation, see these reviews: [7,29,156].

Predator management: Because predation pressure on gopher tortoise populations may be substantial (see Survival), several researchers have suggested that "any management plan to sustain gopher tortoise populations must include serious predator control" [7]. Recommended management practices include nest protection and predator-proof enclosures for hatchlings [158], control of nest predators through hunting and trapping [81,135], and prescribed fire [76]. Establishment of preserves, protection of gopher tortoises from overharvest, and public education are suggested to reduce human predation of gopher tortoises [44,81].

FIRE EFFECTS AND USE

SPECIES: Gopherus polyphemus
DIRECT FIRE EFFECTS:
No cases of direct mortality of gopher tortoises caused by fire had been reported as of this writing (2009). Since fire generally only heats the top 2 inches (10 cm) of the soil during fire [134] and gopher tortoise burrows may be as deep as 10 feet (3 m) below the soil surface (see Burrow description), burrows likely shelter gopher tortoises from fire [33,104]. However, active gopher tortoises located away from their burrows may be killed by fire, particularly if the fire is fast-moving [82] or occurs during cold weather when gopher tortoises move slowly (Means 1988 unpublished letter cited in [129]). Because their higher surface area to volume ratios make them more sensitive to heating, young gopher tortoises may be more susceptible to direct mortality caused by fire (see Daily activity). Also, young gopher tortoises may use shallow pallets or burrows (see Burrow description) that may provide inadequate protection from the heat of fire [16], particularly if these shelters are under or near heavy fuel loads and fire is of high intensity or severity. Although no information is available on gopher tortoises, desert tortoises (Gopherus agassizii) located in shallow burrows were killed by fire (see FEIS review of the desert tortoise).

Since active gopher tortoises are susceptible to fire, the timing of fire likely affects the extent of fire-caused mortality and the age and gender classes most affected. Gopher tortoises may be most affected by early spring, late summer, and fall fires, when they are most active and thus more likely to be away from their burrows, without protective cover. Spring is a time of recovery from winter dormancy and increased reproductive activity (i.e., nesting and mating), and late summer and fall (September-October) is a time when food plants have become fibrous and less available, so gopher tortoises may travel long distances in search of food (see Home range and movement and Foraging activity). In late summer and fall, gopher tortoises may be particularly susceptible to the indirect effects of fire, since fire at this time may reduce already limited available forage prior to winter dormancy, a potentially critical time for gopher tortoises. Also, during August through October, gopher tortoises are hatching from their nests (see Annual activity) [7]. Fire occurring during summer, but prior to emergence of hatchlings from their nests (i.e., late June and July), may be best suited to gopher tortoise conservation [7,39,103,156]. For additional information on timing of fire, see Habitat management and Indirect Fire Effects.

Several studies report fire having a positive effect on gopher tortoises. However, most studies are anecdotal, use small sample sizes, or include no controls and/or replicates, so results should be interpreted with caution. General observations at the Ashton Biological Preserve indicated that gopher tortoises in pine-oak sandhills used formerly abandoned burrows after fire, perhaps due to reduced vegetation cover over the burrows [7]. On the Ocala National Forest in Florida, a gopher tortoise excavated a new burrow approximately 30 days after a stand-replacing February prescribed fire in sand pine scrub [41]. On a 2.5-acre (1 ha) site at the St Marks National Wildlife Refuge in Florida, gopher tortoise burrow abundance increased from 2 to 13 burrows over 2 years after a prescribed fire in March 1983. The width of 4 of the 11 new burrows indicated that adult gopher tortoises had immigrated to the site following fire. The remaining 7 new burrows contained recently hatched gopher tortoises, indicating successful postfire reproduction (Means unpublished data cited in [105]). In Hillsborough County, Florida, richness and abundance of reptiles and amphibians were consistently higher on 2.5-acre (1-ha) plots of slash pine-oak/saw-palmetto/pineland threeawn sandhills burned in late spring (late May-early June) at 3 fire-return intervals (1-, 2-, and 7-year intervals) than on a "control" plot protected from fire for 20 years. After 6 years of treatment, the plot subjected to annual prescribed fires had few standing mature trees and sparse herb cover, with many areas of exposed sand. The plot subjected to 3 fires at 2-year intervals had dense herb cover with few standing trees and no exposed sandy areas. The plot subjected to fire at a 7-year interval had many fire-damaged understory trees. In the year after its second burn, this plot had much exposed sand and moderate grass and herb cover. The following year, "more complete" herb cover had reduced the sand cover in the plot. The control plot had a well-established tree canopy and dense understory of saw-palmettos, with herb density intermediate to that of plots burned at 1- and 2-year intervals. Gopher tortoise numbers were higher in annually burned plots than other plots, but no statistical tests were performed. The authors described gopher tortoises as "very abundant" and gopher tortoise numbers "relatively constant year to year" throughout the plots [114]:

Gopher tortoise densities on prescribed burned plots in slash pine-oak/saw-palmetto/pineland threeawn sandhills in southern Florida [114]
Fire-return interval (years burned)
Study year
1983 1984
Gopher tortoises/acre
1-year intervals (1979-1984) 15.4 13.8
2-year intervals (1979, 1981, & 1983) 8.5 10.5
7-year intervals (1976 & 1983) 11.7 10.5
Control* 8.1 5.7
*Burned in 1965.

Across 24 counties in southern Alabama, gopher tortoise densities were not found in unburned pine-scrub oak habitat, while their density averaged 0.64/ha in burned pine/scrub oak habitat [137]. In 7 counties in southern Georgia, gopher tortoise burrows were denser on "frequently burned", open-canopied longleaf pine stands with a "robust" herbaceous ground cover than on agricultural lands (i.e., cropland, field edges, and grazed or mowed pasture), pine plantations (mostly densely stocked with a sparse herbaceous understory), and unburned areas that had succeeded to dense hardwoods with sparse herbaceous ground cover [72]. In southern Georgia, gopher tortoise density and herbaceous biomass were greater in habitat burned at 1- to 2-year intervals than in habitat not burned in 8 years [86]:

Management history, habitat composition, and gopher tortoise density at 4 sites in southern Georgia [86]
 
Site 1
Site 2
Site 3
Site 4
Plant community 16-year-old slash pine plantation with a "sparse tree canopy" harvested longleaf pine-scrub oak 16-year-old slash pine plantation with a "sparse tree canopy" "natural" longleaf pine-scrub oak
Management history
Site preparation
anchor chained and debris windrowed none double-chopped and debris burned none
Harvest history
clearcut and planted to slash pine "much" of the longleaf pine had been harvested from the overstory clearcut and planted to slash pine none
Fire interval
dormant-season fire every 2 years for an unknown period prior to the study devoid of fire for 8 years prior to the study dormant-season fire annually for 8 years prior to the study dormant-season fire annually for approximately 20 years prior to the study, but carried fire every 2-4 years
Habitat composition
Herbaceous plants
(g/10 m²)
1,420 618 1,997 1,272
Pines/ha
478 6 722 317
Oaks/ha
489 917 211 372
Gopher tortoises/ha 10.1 3.3 9.3 15.8

High gopher tortoise densities at sites 1, 3, and 4 were attributed in part to frequent prescribed fire, which reduced oaks and litter and increased herbaceous biomass production. These data suggest that pine plantations managed similarly to longleaf pine-scrub oak habitat may benefit gopher tortoises [86].

Some gopher tortoise research indicates little or no effect of fire on gopher tortoise abundance. On the Ocala National Forest, gopher tortoises were surveyed in mature sand pine forest (≥55 years old) and in sand pine forest 5 to 7 years after the following treatments: 1) severe wildfire, salvage logging, and natural regeneration, 2) clearcutting, roller chopping, and seeding, and 3) clearcutting and seeding. Reptile species richness, diversity, and evenness did not differ among the 3 treatments or the mature forest control site. Although predisturbance data were not available, gopher tortoises were found only at treated sites, and the rate of detection of gopher tortoises was high enough to estimate gopher tortoise density only at the severe-fire site [66]. At the Kennedy Space Center, gopher tortoises were more abundant at sites burned within the previous 3 years than at sites burned at longer intervals. Data were combined across disturbed, oak-palmetto, and palmetto habitats. However, the relationship between time-since-fire and gopher tortoise densities was marginal and explained only a small amount of variation in gopher tortoise density, a result attributed to an abundance of gopher tortoise forage near plots that were not recently burned. In this study, herbaceous cover decreased and shrub height increased with time-since-fire (P<0.01 for all variables) [18]. In Orange and Osceola counties, Florida, gopher tortoise abundance in sand pine scrub was frequently higher at clearcut and roller chopped sites than burned and control sites, but this pattern was not consistent [60]:

Number of active gopher tortoise burrows on control and treated sites in sand pine scrub of Orange and Osceola counties, Florida [60]
 
Site 1
Site 2
Site 3
Treatment
Gopher tortoise age class
Adult Juvenile Adult Juvenile Adult Juvenile
Prescribed fire 4 0 6 0 3 1
Clearcut and roller chopped 7 1 10 1 3 1
Control 0 1 1 6 3 0

INDIRECT FIRE EFFECTS:
Gopher tortoises reach their highest densities in fire-adapted habitats, such as pine-oak sandhills, pine flatwoods, and sand pine-oak scrub, indicating that fire in those habitats is likely to benefit gopher tortoises [9,44]. Without fire or under altered fire regimes, gopher tortoise numbers are typically reduced (see Succession). Fire favors growth and reproduction of herbaceous gopher tortoise forage by maintaining an open-canopy structure, reducing woody plants and litter, and exposing mineral soil [2,9,13,26,43,96,105,151]. In gopher tortoise habitats, species diversity in the groundlayer vegetation increases after fire and is greater on frequently burned than infrequently burned sites [86,91]. Legume species (see Diet) were abundant in longleaf pine/pineland threeawn habitat occupied by gopher tortoises at Ichauway, which had been winter-burned at 1- to 3-year intervals for about 70 years. At this site, legumes were present in all months of the year, but abundance peaked in June immediately following a March prescribed fire and declined thereafter [69]. Nutrient content of preferred forage species may increase after fire [26]. Fire consumes parts of plants that are less digestible for gopher tortoises and stimulates new, more accessible plant growth [7,39,43,103]. Fire may also benefit gopher tortoises by exposing mineral soil and reducing root density, thus facilitating burrowing and nesting [9,32,65,105].

Winter prescribed fire may aid in restoring pineland threeawn and forbs important as gopher tortoise forage on sites where fire has been excluded for long periods [4]. Several fire-dependent herbs often consumed by gopher tortoises (e.g., pineland threeawn, narrowleaf silkgrass (Pityopsis graminifolia), and shortleaf blazing star (Liatris tenuifolia)) "responded positively" to the conditions following a prescribed fire at the Archbold Biological Station by "sprouting, rapid vegetative growth, and abundant flowering and fruiting". The prescribed fire was conducted in February on a pine-oak sandhill site that had succeeded to dense scrub species including evergreen oaks (Chapman, myrtle, and sand live oaks) and palmettos (e.g., scrub and saw-palmettos). Prior to the prescribed fire, the site had not been burned for over 60 years, and only small remnant patches of slash pine, sand pine, turkey oak, and pineland threeawn were present. Approximately 85% of the ground surface was burned using a series of strip head fires 7 to 33 feet (2-10 m) wide. For shortleaf blazing star and narrowleaf silkgrass, measures of reproductive effort were higher on burned plots than unburned plots (P<0.01 for all variables). Pineland threeawn flowering was not observed in unburned plots or prefire plots, but flowering was common in postfire plots [4]. Additional information on the fire and its effects on herbaceous plants is available in Anderson and Menges [4].

Fire could reduce the diversity and/or availability of some gopher tortoise forage in the short term [114]. Abrahamson [1] reported responses of 5 habitats to 2 fires. A lightning-caused May fire reduced total vegetation cover in swales and gallberry-lyonia flatwoods on Lake Wales Ridge, Florida, to 0% cover. However, total vegetation cover had returned to 94% in swales and 95% in gallberry-lyonia flatwoods 232 days after the fire, and complete recovery to prefire levels occurred within 2 years. After a January prescribed fire, total vegetation cover in pine-oak/palmetto flatwoods recovered to prefire levels in 1 year, slash pine/pineland threeawn flatwoods recovered to prefire levels in 3 years, and slash pine/oak-scrub hickory (Carya floridana)/pineland threeawn-palmetto sandhills did not recover to prefire levels during the 5 year study period. The author noted a lack of change in plant species composition following fire in these habitats [1].

Some nonnative invasive plants that occur in gopher tortoise habitats may proliferate after fire [139]. Because nonnative invasive plants may threaten gopher tortoise habitats by displacing native species, disrupting nutrient and fire cycles, and altering plant succession [24,92], fire could degrade habitat for gopher tortoises by creating conditions favorable to invasive species. Since many native herbaceous groundcover plants are adapted to fire and help carry fire, sites with remnant native groundcover may burn more easily and recover more quickly after prescribed fire than sites where the ground cover has been substantially altered by mechanical site treatments, invasive species, or other alterations [21]. For additional information on the effects of fire on gopher tortoise forage, see Fire Management Considerations.

Since gopher tortoise densities are directly related to herbaceous biomass (see Population density), the effect of fire in gopher tortoise habitats depends largely on its effect on herbaceous ground cover, which varies with fire frequency, severity, and season [109]. During late summer and fall, gopher tortoises may be stressed because of limited forage, and gopher tortoises may move over large areas in search of food. Fire occurring during late summer and fall may reduce this already limited forage, subjecting gopher tortoises, particularly hatchlings, to starvation [39,103,105]. Conversely, fires occurring during summer—when herbaceous plants are still growing—may stimulate late-season growth in many plants and provide additional forage for gopher tortoises prior to winter dormancy [105]. Fires during transitions from spring to summer—when intervals between successive rainfalls are increasing and fires may be more intense—may result in flowering of many groundcover species, greater mortality of hardwoods, and more open canopies than fires at other times [126]. Summer fire may create an open vegetation structure, which Cox and others [39] suggested may make it easier for neonates hatching in subsequent months (August-October) to move about, initiate burrows, and find food. In longleaf pine-turkey oak sandhills on the Ocala National Forest, greater numbers of gopher tortoise burrows were recorded on summer-burned plots than on winter-burned plots (P=0.0001). The authors suggested that gopher tortoises may have immigrated to the summer-burned plots in postfire year 1 [121]. However, inference is limited in this study because prefire data on gopher tortoise burrow densities were not available, and no information regarding habitat differences leading to such differences were provided.

FIRE REGIMES:
Gopher tortoises occur in a wide variety of habitats (see Plant Communities), and the periodicity of fires is thought to have varied considerably among these habitats [9,118]. Under presettlement conditions, low-severity, summer surface fires occurred in longleaf pine forests of the southeastern Coastal Plain at intervals of 1 to 10 years, resulting in savannas with widely spaced overstories, minimal midstories of scattered hardwoods, and diverse herbaceous ground cover dominated by perennial bunchgrasses (i.e., threeawn or bluestem) [11,21,35,39,44,87,106,117,127,140]. These conditions provided the diverse herbaceous ground cover gopher tortoises needed for foraging and the open-canopy conditions needed for thermoregulation and nesting [149]. Because fire-return intervals were short in longleaf pine ecosystems, fuels did not accumulate to levels that would allow stand-replacing fires [149]. Among longleaf pine ecosystems, pine flatwoods are more mesic than pine-oak sandhills, so fuels accumulated at greater rates, and fires were more frequent (1-3 years) [21,87]. Dry conditions in pine-oak sandhills typically resulted in sparse fuel accumulations that could carry fire about every 2 to 4 years [83,84]. Without relatively frequent fires, dense understory and midstory oaks develop in pine-oak sandhills and pine flatwoods, and these communities may succeed to hardwood hammocks [2,9,20,44,65,86,131]. Development of dense understory and midstory oaks degrades habitat for gopher tortoises. Oak shade suppresses herbaceous ground cover; leachates from oak litter retard ground cover growth and reduce gopher tortoise forage; oak litter is often so dense that it may discourage burrow initiation; and oak leaf litter is relatively nonflammable and retards fire [105]. The scarcity of herbs, the relatively incombustible accumulation of oak litter and other fuels, and the expansion of understory shrubs and hardwoods preclude most fires from invading hardwood hammock communities. When fire does occur, it is nearly always stand replacing [21,58]. Depending upon site conditions, threeawn may persist in the understory for 20 to 40 years after fire exclusion; however, other herbaceous plants often only bloom the year following fire and decline during postfire years 2 and 3, becoming rare thereafter [35]. Conversely, fires that are too frequent or severe may eliminate longleaf pine recruitment [21,35,43,57,105,126] and may eventually transform longleaf pine forests into other plant communities where gopher tortoises are less abundant [58].

Sand pine scrub communities are adapted to and maintained by severe, infrequent fires and other disturbance, such as windthrow during storms [44,117]. Stand-replacing crown fires historically occurred on average every 20 to 40 years (range: 10-100 years) in sand pine scrub [31,39,41,44,132]. At less productive sites, where fuel accumulation was slow, fire-return intervals may have been longer [117]. Scrub communities lack fine-textured, flashy fuels and require high wind, low humidity, and low fuel moisture for fire to start and spread. When fire occurs in sand pine scrub, mature sand pines are typically killed and the aboveground vegetation consumed. However, because scrub vegetation is dominated by species that sprout (e.g., oaks) or seed (e.g., sand pine and rosemary) after fire, recovery is relatively rapid. Fire in sand pine scrub typically results in little change in species composition or richness, although dominance may shift to more rapidly growing species after fire [66,117,128,132,140]. Unlike pine-oak sandhills and pine flatwoods, a flush of annuals does not typically occur in sand pine scrub the year following fire, so herbaceous ground cover remains sparse [117]. However, fire creates a low, open habitat structure and large areas of exposed mineral soil beneficial to gopher tortoises for burrowing [66]. This is particularly beneficial if such areas are adjacent to other habitats with greater herbaceous forage [16,18]. Gap colonization and closure of open canopies in sand pine scrub is slow, thus suitable gopher tortoise habitat is maintained "for several years" after fire [66,117]. Over time, however, shrubs, lichens, and leaf litter increase as the canopy closes, and gopher tortoises become less abundant [66]. Sand pine is fire-adapted. If fire is excluded from sand pine scrub for long periods (>75 years), sand pines may eventually be eliminated, and sand pine scrub may succeed to xeric hardwood hammock. Frequent fire can also eliminate sand pine from the overstory [31,39,41,44,132,140].

Historically, fires in gopher tortoise habitats were ignited by lightning and indigenous peoples, primarily during the growing season but also during other times of year [39,87,117]. Lightning-caused fires occurred mostly from May through August; the number of fires and burned area peaked in May and June [129]. Many plant species in gopher tortoise habitats are adapted to growing-season fires but not necessarily to dormant-season fires [4,14,34,35,105]. Thus, changes in the seasonality of fires could change the structure and plant composition of gopher tortoise habitats [7,39,86]. Growing-season fires generally reduce hardwoods, maintain the threeawn-dominated herbaceous layer, promote flowering of annual herbs, facilitate seed production of grasses, and increase the abundance and species richness of understory vegetation more effectively than dormant-season fires [129]. Threeawn and associated species are generally denser and bloom more prolifically after a growing-season than a dormant-season fire [35]. On Lake Wales Ridge, Abrahamson [1] reported "vigorous flowering" by pineland threeawn, cutthroat grass (Panicum abscissum), and beardgrasses (Andropogon spp.) after May, June, and July fires; however, these species grew but showed little or no flowering after January fire. Because of weather and plant physiology, growing-season fires may lead to greater mortality of hardwoods [57,63,96,126], whereas dormant-season fires are more likely to encourage the growth of hardwood trees and a dense shrub midstory [39,86,96,106,113,129]. Because the moisture content of deciduous hardwoods is lowest in early spring, early growing-season fires are the most detrimental to their growth, whereas evergreens may be most affected by late growing-season fires [129].

The Fire Regime Table summarizes characteristics of fire regimes for vegetation communities in which gopher tortoise may occur. Follow the links in the table to documents that provide more detailed information on these fire regimes. Further information regarding fire regimes in gopher tortoise habitats is available in these sources: [2,21,58,117,118,126,149,150].

FIRE MANAGEMENT CONSIDERATIONS:
Prescribed fire is generally considered favorable and often recommended for management of gopher tortoise habitats [9,81,86,113]. Several researchers recommend maintaining presettlement annual and seasonal periodicity of fire within habitats occupied by gopher tortoises [105,107,129,156]. Recommended fire-return intervals for gopher tortoise management in pine flatwoods vary from 1 to 3 years [7,156]. Recommended fire-return intervals for gopher tortoise management in pine-oak sandhills are 1 to 3 years [6], 2 to 4 years [86], 2 to 6 years [7], and 2 to 5 years [156]. Because herbaceous ground cover and longleaf pine needles help carry fire, retaining longleaf pine in the overstory and herbs in the ground layer is important in maintaining this forest type [83,86]. Since longleaf pine seedlings <1 year old are susceptible to fire mortality, several researchers recommend that annual fires be avoided in longleaf pine forests [105]. A review states that the potential benefits for gopher tortoises in maintaining a fire regime of periodic, low-severity fire in longleaf pine ecosystems include maintaining the physiognomic character of longleaf pine-bunchgrass ecosystems by excluding plants that are poorly adapted to fire; preparing a seedbed favorable to longleaf pine regeneration; reducing the density of understory vegetation and thus providing microsites for herbaceous plants and gopher tortoise nesting and burrow sites; stimulating increased seed production by native grasses; and releasing nutrients to the soil for uptake by plants [21].

To maintain sand pine scrub habitat in conditions suitable for gopher tortoises, researchers recommend fire-return intervals of 15 to 30 years [7,156]. In sand pine scrub communities, fire stimulates flowering and seed production of many species, enhances regeneration by exposing bare mineral soil, reduces shading from woody understory species, reduces fuel loads, releases nutrients to the soil, accelerates growth in the herbaceous layer, and increases fruit and forage production [41]. Some researchers have speculated that clearcutting may "mimic the natural situation of infrequent crown fires to which the scrub fauna is adapted" [31]. However, other researchers demonstrated that timber harvesting in Florida sand pine scrub had a negative effect or mixed results on herpetofaunal species richness and diversity compared with prescribed fire [60,128], and that the 2 treatments resulted in different herpetofaunal assemblages [66,128]. On the Ocala National Forest, Custer and Thorsen [41] found that stand-replacing prescribed fire could be used in sand pine scrub habitat to stimulate regeneration of sand pines in gopher tortoise habitats. For more information on this topic, see these reviews: [65,117].

Growing-season fires are often recommended for management of gopher tortoise habitats because plant responses to growing-season fires benefit gopher tortoise survival and recruitment much more than plant responses to fire during other seasons (see Indirect Fire Effects) [86,105]. However, because higher fuel loads in fire-excluded longleaf pine forests may allow stand-replacing fires [149] and make it difficult to control prescribed fire [43], alternative management strategies are often suggested for gopher tortoise habitats to avoid killing overstory trees [21,43,105,131]. Several researchers recommend that if too much fuel has accumulated to make summer prescribed fire practical, one or more consecutive winter fires "may provide a way to begin a fire program" [43]. Once fuels are reduced, prescribed fire may be conducted during the growing season [21,43,86,105,148,156]. Landers and Speakes [86] found that longleaf pine-scrub oak habitats winter-burned at 2- to 4-year intervals sustained high gopher tortoise densities (15.8/ha). Nonetheless, the authors recommended summer fires every 5 to 10 years for management of gopher tortoises in longleaf pine-scrub oak habitats to control hardwoods and increase canopy openness. For more information on this study, see Direct Fire Effects. Brockway and Lewis [20] acknowledged the importance of growing-season fires in longleaf pine flatwoods but reported that frequent, winter prescribed fires maintained high grass cover in this community. Frequent (every 2 years) dormant-season prescribed fires conducted over 17 years in longleaf pine flatwoods in Berrien County, Georgia, resulted in increased cover of some understory shrubs (shiny blueberry (Vaccinium myrsinites)), decreased cover in other shrubs (e.g., inkberry), and increased cover of graminoids (e.g., bluestem grasses, pineland threeawn, and Curtis' dropseed (Sporobolus curtissii)) compared with an unburned control site. However, dormant-season fire resulted in no difference in forb cover among sites, a result attributed to high residual cover of gallberry and other shrubs on fire-treated plots [20]. At the Camp Shelby Training Site in Mississippi, prescribed fire was implemented in previously fire-excluded longleaf pine forests at 4 sites during January and February 2002 to reduce fuel loads and dense shrub understories. Prescribed fire was conducted at these sites again during April 2003. One cycle of "dormant-and-growing-season prescribed fire" did not restore desired habitat conditions for gopher tortoises. Although understory shrub biomass was reduced in the short term, herbaceous forage and canopy openness did not increase. Reintroduction of fire in this 4-year study did not result in large shifts of habitat use by gopher tortoises from adjacent habitats to burned areas, although 3 gopher tortoises used burrows in postfire habitat. These authors suggested that repeated application of dormant- and/or growing-season prescribed fires may increase gopher tortoise use of these forests in the long term [159]. At the Escambia Experimental Forest in Alabama, winter burning at 3-year intervals for about 40 years led to a "significant and increasing hardwood component". The authors indicated a "need to resort to a temporary series of spring burns to bring the hardwoods back under control". Because of safety and logistical concerns, the authors suggested that "winter burning should be the operational norm" [57]. Several researchers suggested that, at sites where hardwoods are too large to be controlled by winter prescribed fire, selective cutting or herbicide treatments prior to winter prescribed fire may be effective [21,57]. Other researchers report that it is necessary to reintroduce summer fire to restore longleaf pine ecosystems [129].

In general, results from seasonal fire experiments indicate that effects of prescribed fire season on vegetation may be minor relative to the effects of long-term fire exclusion [19,83,105]. Ultimately, using prescribed fire to restore longleaf pine habitat will vary with local site conditions and degree of ecological succession [21]. For more information on this topic, see these reviews: [21,129]. For details on employing prescribed fire in longleaf pine ecosystems, see reviews by Landers and others [84] and Robbins and Myers [129].

Fire historically provided a mosaic of habitat types in a variety of seral stages and fuel conditions over the landscape [21,84,147], thus several researchers suggested that longleaf pine ecosystems be managed for a variety of seral stages [7,84]. Ashton and Ashton [7] suggested that prescribed fire conducted in a mosaic pattern over large areas may be beneficial for gopher tortoises. However, the effects of patch mosaic burning and different burning patterns (shape and size) on gopher tortoises have not been tested and remain largely unknown [124].

APPENDIX: FIRE REGIME TABLE

SPECIES: Gopherus polyphemus
The following table provides fire regime information that may be relevant to gopher tortoise habitats. Follow the links in the table to documents that provide more detailed information on these fire regimes.

Fire regime information on vegetation communities in which gopher tortoise may occur. This information is taken from the LANDFIRE Rapid Assessment Vegetation Models [89], which were developed by local experts using available literature, local data, and/or expert opinion. This table summarizes fire regime characteristics for each plant community listed. The PDF file linked from each plant community name describes the model and synthesizes the knowledge available on vegetation composition, structure, and dynamics in that community. Cells are blank where information is not available in the Rapid Assessment Vegetation Model.
Southeast
Southeast
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Southeast Grassland
Floodplain marsh Replacement 100% 4 3 30
Everglades (marl prairie) Replacement 45% 16 10 20
Mixed 55% 13 10  
Palmetto prairie Replacement 87% 2 1 4
Mixed 4% 40    
Surface or low 9% 20    
Southern tidal brackish to freshwater marsh Replacement 100% 5    
Gulf Coast wet pine savanna Replacement 2% 165 10 500
Mixed 1% 500    
Surface or low 98% 3 1 10
Southeast Woodland
Longleaf pine/bluestem Replacement 3% 130    
Surface or low 97% 4 1 5
Longleaf pine (mesic uplands) Replacement 3% 110 40 200
Surface or low 97% 3 1 5
Longleaf pine-Sandhills prairie Replacement 3% 130 25 500
Surface or low 97% 4 1 10
Pine rocklands
Mixed 1% 330    
Surface or low 99% 3 1 5
Pond pine Replacement 64% 7 5 500
Mixed 25% 18 8 150
Surface or low 10% 43 2 50
South Florida slash pine flatwoods Replacement 6% 50 50 90
Surface or low 94% 3 1 6
Atlantic wet pine savanna Replacement 4% 100    
Mixed 2% 175    
Surface or low 94% 4     
Southeast Forested
Sand pine scrub Replacement 90% 45 10 100
Mixed 10% 400 60  
Maritime forest Replacement 18% 40   500
Mixed 2% 310 100 500
Surface or low 80% 9 3 50
Mesic-dry flatwoods Replacement 3% 65 5 150
Surface or low 97% 2 1 8
*Fire Severities
Replacement: Any fire that causes greater than 75% top removal of a vegetation-fuel type, resulting in general replacement of existing vegetation; may or may not cause a lethal effect on the plants.
Mixed: Any fire burning more than 5% of an area that does not qualify as a replacement, surface, or low-severity fire; includes mosaic and other fires that are intermediate in effects.
Surface or low: Any fire that causes less than 25% upper layer replacement and/or removal in a vegetation-fuel class but burns 5% or more of the area [70,88].

REFERENCES

SPECIES: Gopherus polyphemus
1. Abrahamson, Warren G. 1984. Post-fire recovery of Florida Lake Wales Ridge vegetation. American Journal of Botany. 71(1): 9-21. [9509]
2. Abrahamson, Warren G.; Hartnett, David C. 1990. Pine flatwoods and dry prairies. In: Myers, Ronald L.; Ewel, John J., eds. Ecosystems of Florida. Orlando, FL: University of Central Florida Press: 103-149. [17388]
3. Alford, Ross A. 1980. Population structure of Gopherus polyphemus in northern Florida. Journal of Herpetology. 14: 177-182. [74123]
4. Anderson, Roger C.; Menges, Eric S. 1997. Effects of fire on sandhill herbs: nutrients, mycorrhizae, and biomass allocation. American Journal of Botany. 84(7): 938-948. [28453]
5. Arata, Andrew A. 1958. Notes on the eggs and young of Gopherus polyphemus (Daudin). Quarterly Journal of the Florida Academy of Science. 21: 274-280. [74134]
6. Aresco, Matthew J.; Guyer, Craig. 1999. Burrow abandonment by gopher tortoises in slash pine plantations of the Conecuh National Forest. Journal of Wildlife Management. 63(1): 26-35. [30017]
7. Ashton, Ray E.; Ashton, Patricia Sawyer. 2008. The natural history and management of the gopher tortoise Gopherus polyphemus (Daudin). Malibar, FL: Krieger Publishing Company. 275 p. [73126]
8. Auffenberg, Walter. 1969. Tortoise behavior and survival. Patterns of Life Series: Biological Sciences Curriculum Study. Chicago, IL: Rand McNally and Co. 38 p. [74138]
9. Auffenberg, Walter; Franz, Richard. 1982. The status and distribution of the gopher tortoise (Gopherus polyphemus). In: Bury, R. B., ed. North American tortoises: conservation and ecology. Research Report No. 12. Washington, DC: U.S. Department of the Interior, Fish and Wildlife Service: 95-126. [34390]
10. Auffenberg, Walter; Iverson, John B. 1979. Demography of terrestrial turtles. In: Harless, Marion; Morlock, Henry, eds. Turtles: Perspectives and research. New York: John Wiley and Sons: 541-569. [74625]
11. Barbour, Michael G.; Billings, William Dwight, eds. 1988. North American terrestrial vegetation. Cambridge; New York: Cambridge University Press. 434 p. [13876]
12. Birkhead, Roger D.; Guyer, Craig; Hermann, Sharon M. 2005. Patterns of folivory and seed ingestion by gopher tortoises (Gopherus polyphemus) in a southeastern pine savanna. The American Midland Naturalist. 154(1): 143-151. [54515]
13. Boglioli, Melissa Dills. 1999. Burrow dispersion and occupancy patterns as they relate to habitat parameters and social behavior in the gopher tortoise, Gopherus polyphemus. Auburn, AL: Auburn University. 62 p. Thesis. [73133]
14. Boyles-Sprenkel, Carolee. 1993. Restoring a "grass-roots" forest. American Forests. 99(5&6): 43-45, 60-61. [21284]
15. Brantly, Robert M. 1989. Guidelines for gopher tortoise relocations. In: Diemer, Joan E.; Jackson, Dale R.; Landers, J. Larry; Layne, James N.; Wood, Don A., eds. Gopher tortoise relocation symposium: Proceedings; 1987 June 27; Gainesville, FL. Nongame Wildlife Program Technical Report #5. Tallahassee, FL: Florida Game and Fresh Water Fish Commission: 103-109. [75220]
16. Breininger, D. R.; Schmalzer, P. A.; Rydene, D. A.; Hinkle, C. R. 1988. Burrow and habitat relationships of the gopher tortoise in coastal scrub and slash pine flatwoods on Merritt Island, Florida. Final report: Project GFC-84-016. Tallahassee, FL: Florida Game and Fresh Water Fish Commission, Nongame Wildlife Program. 238 p. [74246]
17. Breininger, David R.; Schmalzer, Paul A.; Hinkle, C. Ross. 1991. Estimating occupancy of gopher tortoise (Gopherus polyphemus) burrows in coastal scrub and slash pine flatwoods. Journal of Herpetology. 25(3): 317-321. [74120]
18. Breininger, David R.; Schmalzer, Paul A.; Hinkle, C. Ross. 1994. Gopher tortoise (Gopherus polyphemus) densities in coastal scrub and slash pine flatwoods in Florida. Journal of Herpetology. 28(1): 60-65. [34385]
19. Brennan, L. A.; Engstrom, R. T.; Palmer, W. E.; Hermann, S. M.; Hurst, G. A.; Burger, L. W.; Hardy, C. L. 1998. Whither wildlife without fire? In: Wadsworth, K. G., ed. Transactions of the 63rd North American wildlife and natural resources conference: Proceedings; 1998 March 20-24; Orlando, FL. Washington, DC: Wildlife Management Institute: 402-414. [30010]
20. Brockway, Dale G.; Lewis, Clifford E. 1997. Long-term effects of dormant-season prescribed fire on plant community diversity, structure and productivity in a longleaf pine wiregrass ecosystem. Forest Ecology and Management. 96: 167-183. [29222]
21. Brockway, Dale G.; Outcalt, Kenneth W.; Tomczak, Donald J.; Johnson, Everett E. 2005. Restoration of longleaf pine ecosystems. Gen. Tech. Rep. SRS-83. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southern Research Station. 34 p. [64063]
22. Brode, William E. 1959. Notes on the behavior of Gopherus polyphemus. Herpetologica. 15: 101-102. [74133]
23. Brooks, Jeffrey J.; Johnson, A. Sydney; Miller, Karl V. 1993. Effects of chemical site preparation on wildlife habitat and plant species diversity in the Georgia sandhills. In: Brissette, John C., ed. Proceedings, 7th biennial southern silivicultural research conference; 1992 November 17-19; Mobile, AL. Gen. Tech. Rep. SO-93. New Orleans, LA: U.S. Department of Agriculture, Forest Service, Southern Forest Experiment Station: 605-612. [23325]
24. Brooks, Matthew L. 2008. Plant invasions and fire regimes. In: Zouhar, Kristin; Smith, Jane Kapler; Sutherland, Steve; Brooks, Matthew L., eds. Wildland fire in ecosystems: fire and nonnative invasive plants. Gen. Tech. Rep. RMRS-GTR-42-vol. 6. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 33-45. [70467]
25. Bryant, Danny; Boykin, Jay. 2007. Fuels management on the national forests in Mississippi after Hurricane Katrina. In: Butler, Bret W.; Cook, Wayne, comps. The fire environment--innovations, management, and policy: conference proceedings; 2007 March 26-30; Destin, FL. Proceedings RMRS-P-46CD. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 287-300. CD-ROM. [71107]
26. Buckner, James L.; Landers, J. Larry. 1979. Fire and disking effects on herbaceous food plants and seed supplies. Journal of Wildlife Management. 43(3): 807-811. [11966]
27. Burke, Russell L. 1989. Florida gopher tortoise relocation: overview and case study. Biological Conservation. 48: 295-309. [16420]
28. Burke, Russell L.; Cox, James. 1988. Evaluation and review of field techniques used to study and manage gopher tortoises. In: Szaro, Robert C.; Severson, Kieth E.; Patton, David R., technical coordinators. Management of amphibians, reptiles, and small mammals in North America: Proceedings of the symposium; 1988 July 19-21; Flagstaff, AZ. Gen. Tech. Rep. RM-166. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 205-215. [16261]
29. Burke, Russell. 1989. Burrow-to-tortoise conversion factors: comparison of three gopher tortoise survey techniques. Herpetological Review. 20: 92-94. [74599]
30. Butler, Joseph A.; Hull, Todd W. 1996. Reproduction of the tortoise, Gopherus polyphemus, in northeastern Florida. Journal of Herpetology. 30: 14-18. [74148]
31. Campbell, Howard W.; Christman, Stephen P. 1982. The herpetological components of Florida sandhill and sand pine scrub associations. In: Scott, Norman J., Jr., ed. Herpetological communities: a symposium of the Society for the Study of Amphibians and Reptiles and the Herpetologists' League: Proceedings; August 1977; [Meeting location unknown]. Wildlife Res. Pap. 13. Washington, DC: U.S. Department of the Interior, Fish and Wildlife Service: 163-171. [28592]
32. Carlile, Lawrence D. 1997. Fire effects on threatened and endangered species and habitats of Fort Stewart Military Reservation, Georgia. In: Greenlee, Jason M., ed. Proceedings: 1st conference on fire effects on rare and endangered species and habitats; 1995 November 13-16; Coeur d'Alene, ID. Fairfield, WA: International Association of Wildland Fire: 227-231. [52064]
33. Carr, Archie. 1952. Handbook of turtles: The turtles of the United States, Canada, and Baja California. Ithaca, NY: Cornell University Press. 542 p. [13691]
34. Christman, Steven P. 1983. Timber management is not wildlife management. In: Bryant, Rhoda J.; Franz, Richard, eds. The gopher tortoise: a keystone species: Proceedings of the 4th annual meeting of the Gopher Tortoise Council; 1983 October 29; Valdosta, GA. Gainesville, FL: Florida Museum of Natural History; Gopher Tortoise Council: 5-18. [74343]
35. Clewell, Andre F. 1989. Natural history of wiregrass (Aristida stricta Michx., Gramineae). Natural Areas Journal. 9(4): 223-233. [10092]
36. Conant, Roger; Collins, Joseph T. 1991. A field guide to reptiles and amphibians: Eastern and central North America. 3rd ed. Peterson Field Guide Series No. 12. Boston, MA: Houghton Mifflin Company. 450 p. [22902]
37. Conde, Louis F.; Swindel, Benee F.; Smith, Joel E. 1983. Plant species cover, frequency, and biomass: early responses to clearcutting, burning, chopping, windrowing, discing, and bedding in Pinus elliottii flatwoods. Forest Ecology and Management. 6: 319-331. [75794]
38. Conde, Louis F.; Swindel, Benee F.; Smith, Joel E. 1983. Plant species cover, frequency, and biomass: early responses to clearcutting, chopping, and bedding in Pinus elliottii flatwoods. Forest Ecology and Management. 6: 307-317. [9661]
39. Cox, James; Inkley, Douglas; Kautz, Randy. 1987. Ecology and habitat protection needs of gopher tortoise (Gopherus polyphemus) populations found on lands slated for large-scale development in Florida. Nongame Wildlife Program Technical Report No. 4. Tallahassee, FL: Florida Game and Fresh Water Fish Commission, Office of Environmental Services,. 75 p. [73129]
40. Crother, Brian I. 2000. Scientific and standard English names of amphibians and reptiles of North America north of Mexico, with comments regarding confidence in our understanding. Herpetological Circular No. 29. Lawrence, KS: Society for the Study of Amphibians and Reptiles. 82 p. [54172]
41. Custer, George; Thorsen, James. 1996. Stand-replacement burn in the Ocala National Forest--a success. Fire Management Notes. 56(2): 7-12. [50617]
42. Davis, Jennifer L. 2007. Home range and activity patterns in a South Florida population of the tortoise Gopherus polyphemus on Savannas Preserve State Park. Boca Raton, FL: Florida Atlantic University. 53 p. Thesis. [73505]
43. DeBerry, Drue; Pashley, David. 2004. Pine ecosystem conservation handbook for the gopher tortoise: A guide for family forest owners. Washington, DC: American Forest Foundation, Center for Conservation Solutions. 48 p. Available online: http://www.conservationforestry.org/Documents/Handbook_Gopher_Tortoise.pdf. [74899]
44. Diemer, Joan E. 1986. The ecology and management of the gopher tortoise in the southeastern United States. Herpetologica. 42(1): 125-133. [16418]
45. Diemer, Joan E. 1989. Gopherus polyphemus, gopher tortoise. In: Swingland, Ian R.; Klemens, Michael W., eds. The conservation biology of tortoises. Occasional Papers of the IUCN Species Survival Commission (SCS): No. 5. Gland, Switzerland: IUCN--The World Conservation Union: 14-16. [74130]
46. Diemer, Joan E. 1992. Demography of the tortoise Gopherus polyphemus in northern Florida. Journal of Herpetology. 26: 281-289. [74070]
47. Diemer, Joan E. 1992. Home range and movements of the tortoise Gopherus polyphemus in northern Florida. Journal of Herpetology. 26: 158-165. [74636]
48. Diemer, Joan E.; Moler, Paul E. 1982. Gopher tortoise response to site preparation in northern Florida. Proceedings, Annual Conference of Southeastern Area Fish and Wildlife Agencies. 36 :634-637. [74131]
49. Diemer, Joan E.; Moore, Clinton T. 1994. Reproduction of gopher tortoises in north-central Florida. In: Bury, R. B.; Germano, D. J., eds. Biology of North American tortoises. Fish and Wildlife Research Report 13. Washington, DC: U.S. Department of the Interior, National Geological Survey: 129-137. [75024]
50. Doonan, Terry J.; Stout, I. Jack. 1994. Effects of gopher tortoise (Gopherus polyphemus) body size on burrow structure. The American Midland Naturalist. 131: 273-280. [75304]
51. Douglass, John F.; Layne, James N. 1978. Activity and thermoregulation of the gopher tortoise (Gopherus polyphemus) in southern Florida. Herpetologica. 34: 359-374. [74069]
52. Eisenberg, John. 1983. The gopher tortoise as a keystone species. In: Bryant, Rhoda J.; Franz, Richard, eds. The gopher tortoise: a keystone species: Proceedings of the 4th annual meeting of the Gopher Tortoise Council; 1983 October 29; Valdosta, GA. Gainesville, FL: Florida Museum of Natural History; Gopher Tortoise Council: 1-4. [74638]
53. Epperson, Deborah M.; Heise, Colleen D. 2000. Nesting and hatchling ecology of gopher tortoises (Gopherus polyphemus) in southern Mississippi. Journal of Herpetology. 39: 315-324. [74147]
54. Ernst, Carl H.; Lovich, Jeffrey E.; Barbour, Roger W. 1994. Turtles of the United States and Canada. Washington, DC: Smithsonian Institution Press. 578 p. [74176]
55. Eubanks, Jeannie Ott; Michener, William K.; Guyer, Craig. 2003. Patterns of movement and burrow use in a population of gopher tortoises (Gopherus polyphemus). Herpetologica. 59: 311-321. [74720]
56. Eubanks, Jennine Ott; Hollister, Jeff W.; Guyer, Craig; Michener, William K. 2002. Reserve area requirements for gopher tortoises (Gopherus polyphemus). Chelonian Conservation and Biology. 4(2): 464-471. [74719]
57. Farrar, Robert M., Jr. 1998. Prescribed burning in selection stands of southern pine: current practice and future promise. In: Pruden, Teresa L.; Brennan, Leonard A., eds. Fire in ecosystem management: shifting the paradigm from suppression to prescription: Proceedings, Tall Timbers fire ecology conference; 1996 May 7-10; Boise, ID. No. 20. Tallahassee, FL: Tall Timbers Research Station: 151-160. [35625]
58. Florida Natural Areas Inventory. 1990. Guide to the natural communities of Florida, [Online]. In: Species and communities. Tallahassee, FL: Florida Natural Areas Inventory; Florida State University (Producer). Available: http://www.fnai.org/PDF/Natural_Communities_Guide.pdf [2009, June 2]. [74325]
59. Garner, James A.; Landers, J. Larry. 1981. Foods and habitat of the gopher tortoise in southwestern Georgia. Proceedings, Annual Conference of Southeastern Association of Fish and Wildlife Agencies. 35:120-134. [74124]
60. Gianopulos, Kristie D. 2001. Response of the threatened sand skink (Neoseps reynoldsi) and other herpetofaunal species to burning and clearcutting in the Florida sand pine scrub habitat. Tampa, FL: University of South Florida. 91 p. Thesis. [62064]
61. Gibbons, J. Whitfield. 1987. Why do turtles live so long? BioScience. 37(4): 292-269. [75620]
62. Gibson, David J.; Hartnett, David C.; Merrill, Gary L. S. 1990. Fire temperature heterogeneity in contrasting fire prone habitats: Kansas tallgrass prairie and Florida sandhill. Bulletin of the Torrey Botanical Club. 117(4): 348-356. [14138]
63. Glitzenstein, Jeff S.; Platt, William J.; Streng, Donna R. 1995. Effects of fire regime and habitat on tree dynamics in north Florida longleaf pine savannas. Ecological Monographs. 65(4): 441-476. [26727]
64. Godley, J. Steve. 1989. Comparison of gopher tortoise populations relocated onto reclaimed phosphate-mined sites in Florida. In: Diemer, Joan E.; Jackson, Dale R.; Landers, J. Larry; Layne, James N.; Wood, Don A., eds. Gopher tortoise relocation symposium: Proceedings; 1987 June 27; Gainesville, FL. Nongame Wildlife Program Technical Report #5. Tallahassee, FL: Florida Game and Fresh Water Fish Commission: 43-58. [49252]
65. Greenberg, Cathryn H. 2002. Fire, habitat structure and herpetofauna in the Southeast. In: Ford, W. Mark; Russell, Kevin R.; Moorman, Christopher E., eds. The role of fire in nongame wildlife management and community restoration: traditional uses and new directions: Proceedings of a special workshop; 2000 December 15; Nashville, TN. Gen. Tech. Rep. NE-288. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northeastern Research Station: 91-98. [41565]
66. Greenberg, Cathryn H.; Neary, Daniel G.; Harris, Larry D. 1994. Effect of high-intensity wildfire and silvicultural treatments on reptile communities in sand-pine scrub. Conservation Biology. 8(4): 1047-1057. [34511]
67. Guyer, Craig; Bailey, Mark A. 1993. Amphibians and reptiles of longleaf pine communities. In: Hermann, Sharon M., ed. The longleaf pine ecosystem: ecology, restoration, and management: Proceedings, 18th Tall Timbers fire ecology conference; 1991 May 30 - June 2; Tallahassee, FL. Tallahassee, FL: Tall Timbers Research, Inc: 139-158. [28421]
68. Guyer, Craig; Hermann, Sharon M. 1997. Patterns of size and longevity of gopher tortoise (Gopherus polyphemus) burrows: implications for the longleaf pine ecosystem. Chelonian Conservation and Biology. 2(4): 507-513. [74142]
69. Hainds, Mark J.; Mitchell, Robert J.; Palik, Brian J.; Boring, Lindsay R.; Gjerstad, Dean H. 1999. Distribution of native legumes (Leguminoseae) in frequently burned longleaf pine (Pinaceae)--wiregrass (Poaceae) ecosystems. American Journal of Botany. 86(11): 1606-1614. [35495]
70. Hann, Wendel; Havlina, Doug; Shlisky, Ayn; [and others]. 2008. Interagency fire regime condition class guidebook. Version 1.3, [Online]. In: Interagency fire regime condition class website. U.S. Department of Agriculture, Forest Service; U.S. Department of the Interior; The Nature Conservancy; Systems for Environmental Management (Producer). 119 p. Available: http://frames.nbii.gov/frcc/documents/FRCC_Guidebook_2008.07.10.pdf [2008, September 03]. [70966]
71. Hermann, Sharon M. 1993. Small-scale disturbances in longleaf pine forests. In: Hermann, Sharon M., ed. The longleaf pine ecosystem: ecology, restoration and management: Proceedings, 18th Tall Timbers fire ecology conference; 1991 May 30 - June 2; Tallahassee, FL. Tallahassee, FL: Tall Timbers Research, Inc: 265-274. [28373]
72. Hermann, Sharon M.; Guyer, Craig; Waddle, J. Hardin; Nelms, M. Greg. 2002. Sampling on private property to evaluate population status and effects of land use practices on the gopher tortoise, Gopherus polyphemus. Biological Conservation. 108: 289-298. [74721]
73. Herrington, Marilyn Davis. 1996. The effects of stand thinning on gopher tortoises, Gopherus polyphemus (Testudines: Testudinidae), in the Conecuh National Forest. Auburn, AL: Auburn University. 47 p. Thesis. [73508]
74. Humphrey, Stephen R.; Eisenberg, John F.; Franz, Richard. 1985. Possibilities for restoring wildlife of a longleaf pine savanna in an abandoned citrus grove. Wildlife Society Bulletin. 13: 487-496. [16416]
75. Jackson, Dale R. 1989. The fauna of gopher tortoise burrows. In: Diemer, Joan E.; Jackson, Dale R.; Landers, J. Larry; Layne, James N.; Wood, Don A., eds. Gopher tortoise relocation symposium: Proceedings; 1987 June 27; Gainesville, FL. Nongame Wildlife Program Technical Report #5. Tallahassee, FL: Florida Game and Fresh Water Fish Commission: 86-88. [49259]
76. Jones, David D.; Conner, Mike; Storey, Theresa H.; Warren, Robert J. 2004. Prescribed fire and raccoon use of longleaf pine forests: implications for managing nest predation? Wildlife Society Bulletin. 32(4): 1255-1259. [54693]
77. Kaczor, Sue A.; Hartnett, David C. 1990. Gopher tortoise (Gopherus polyphemus) effects on soils and vegetation in a Florida sandhill community. The American Midland Naturalist. 123: 100-111. [74143]
78. Kantola, Angela Torres; Humphrey, Stephen R. 1990. Habitat use by Sherman's fox squirrel (Sciurus niger shermani) in Florida. Journal of Mammalogy. 71(3): 411-419. [26449]
79. Kush, John S.; Meldahl, Ralph S. 2000. Composition of a virgin stand of longleaf pine in south Alabama. Castenea. 65(1): 56-63. [38811]
80. Kushlan, James A.; Mazzitti, Frank J. 1984. Environmental effects on a coastal population of gopher tortoise. Journal of Herpetology. 18: 231-239. [74598]
81. Landers, J. Larry. 1980. Recent research on the gopher tortoise and its implications. In: Franz, R.; Bryant, R. J., eds. The dilemma of the gopher tortoise -- is there a solution? Proceedings of the 1st annual meeting of the Gopher Tortoise Council; [Date unknown]; Auburn, AL. Gainesville, FL: Florida Museum of Natural History; Gopher Tortoise Council: 8-14. [74160]
82. Landers, J. Larry. 1987. Prescribed burning for managing wildlife in southeastern pine forests. In: Dickson, James G.; Maughan, O. Eugene, eds. Managing southern forests for wildlife and fish: a proceedings; 1986 October 5-8; Birmingham, AL. Gen. Tech. Rep. SO-65. New Orleans, LA: U.S. Department of Agriculture, Forest Service, Southern Forest Experimental Station: 19-27. [Proceedings of the Wildlife and Fish Ecology Technical Session, 1986 Society of American Foresters National Convention]. [25968]
83. Landers, J. Larry; Buckner, James L. 1981. The gopher tortoise: effects of forest management and critical aspects of its ecology. Forest Productivity and Research: Technical Note No. 56. Bainbridge, GA: International Paper Company, Wood Products and Resources Group, Southlands Experiment Forest. 7 p. [75641]
84. Landers, J. Larry; Byrd, Nathan A.; Komarek, Roy. 1990. A holistic approach to managing longleaf pine communities. In: Farrar, Robert M., Jr., ed. Proceedings of the symposium on the management of longleaf pine; 1989 April 4-6; Long Beach, MS. Gen. Tech. Rep. SO-75. New Orleans, LA: U.S. Department of Agriculture, Forest Service, Southern Forest Experiment Station: 135-167. [46849]
85. Landers, J. Larry; Garner, James A.; McRae, W. Alan. 1980. Reproduction of gopher tortoises (Gopherus polyphemus) in southwestern Georgia. Herpetologica. 36(4): 353-361. [16419]
86. Landers, J. Larry; Speake, Dan W. 1980. Management needs of sandhill reptiles in southern Georgia. Proceedings, Annual Conference of Southeast Association Fish & Wildlife Agencies. 34: 515-529. [21092]
87. Landers, Larry; Wade, Dale. 1994. Disturbance, persistence and diversity of the longleaf pine - bunchgrass ecosystem. In: Forests together: meeting tomorrow's challenges: Proceedings of the 1993 Society of American Foresters national convention; 1993 November 7-10; Indianapolis, IN. SAF Publication 94-01. Bethesda, MD: Society of American Foresters: 182-188. [23976]
88. LANDFIRE Rapid Assessment. 2005. Reference condition modeling manual (Version 2.1), [Online]. In: LANDFIRE. Cooperative Agreement 04-CA-11132543-189. Boulder, CO: The Nature Conservancy; U.S. Department of Agriculture, Forest Service; U.S. Department of the Interior (Producers). 72 p. Available: http://www.landfire.gov/downloadfile.php?file=RA_Modeling_Manual_v2_1.pdf [2007, May 24]. [66741]
89. LANDFIRE Rapid Assessment. 2007. Rapid assessment reference condition models, [Online]. In: LANDFIRE. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Lab; U.S. Geological Survey; The Nature Conservancy (Producers). Available: http://www.landfire.gov/models_EW.php [2008, April 18] [66533]
90. Layne, James N. 1989. Comparison of survival rates and movements of relocated and resident gopher tortoises in a south-central Florida population. In: Diemer, Joan E.; Jackson, Dale R.; Landers, J. Larry; Layne, James N.; Wood, Don A., eds. Gopher tortoise relocation symposium: Proceedings; 1987 June 27; Gainesville, FL. Nongame Wildlife Program Technical Report #5. Tallahassee, FL: Florida Game and Fresh Water Fish Commission: 73-83. [49254]
91. Lemon, Paul C. 1949. Successional responses of herbs in the longleaf-slash pine forest after fire. Ecology. 30(2): 135-145. [10133]
92. Lippincott, Carol L. 1997. Ecological consequences of Imperata cylindrica (cogongrass) invasion in Florida sandhill. Gainesville, FL: University of Florida. 165 p. Dissertation. [48904]
93. Lohoefener, Ren. 1982. Gopher tortoise ecology and land-use practices in southern DeSoto National Forest, Harrison County, Mississippi. In: Franz, R.; Bryandt, R. J., eds. The gopher tortoise and its sandhill habitat: Proceedings of the 3rd annual meeting of the Gopher Tortoise Council; 1982 October 2; [Tallahassee, FL]. Gainesville, FL: Florida Museum of Natural History; Gopher Tortoise Council: 50-74. [74655]
94. Lohoefener, Ren; Lohmeier, Lynne. 1981. Comparison of gopher tortoise (Gopherus polyphemus) habitats in young slash pine and old longleaf pine areas of southern Mississippi. Journal of Herpetology. 15: 239-242. [74125]
95. Lohoefener, Ren; Lohmeier, Lynne. 1986. Experiments with gopher tortoise (Gopherus polyphemus) relocation in southern Mississippi. Herpetological Review. 17(2): 37, 39-40. [74135]
96. Long, A. J.; Behm, A.; Cassidy, L.; DiMartino, J.; Doran, D.; Freeman, D.; Helmers, J.; Keefe, K.; Miller, A.; Ranasinghe, S.; Randall, C.; Rasser, M.; Ruth, A.; Shipley, D.; Van Loan, A. 2004. Prescribed fire and slash pine. In: Dickens, E. D.; Barnett, J. P.; Hubbard, W. G.; Jokela, E. J., eds. Slash pine: still growing and growing! Proceedings of the slash pine symposium; 2002 April 23-25; Jekyll Island, GA. Gen. Tech. Rep. SRS-76. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southern Research Station: 66-78. [55324]
97. Lovich, Jeffrey E.; Daniels, Ramona. 2000. Environmental characteristics of desert tortoise (Gopherus agassizii) burrow locations in an altered industrial landscape. Chelonian Conservation and Biology. 3(4): 714-721. [67616]
98. MacDonald, L. A.; Mushinsky, Henry R. 1988. Foraging ecology of the gopher tortoise, Gopherus polyphemus, in a sandhill habitat. Herpetologica. 44(3): 345-353. [16422]
99. Maehr, David S.; Brady, James R. 1984. Food habits of Florida black bears. Journal of Wildlife Management. 48(1): 230-235. [67705]
100. Marshall, John E. 1987. The effects of nest predation on hatching success in gopher tortoises (Gopherus polyphemus Daudin 1802). Mobile, AL: University of South Alabama. 67 p. Thesis. [74128]
101. Martin, Paige L.; Layne, James N. 1987. Relationship of gopher tortoise body size to burrow size in a south-central Florida population. Florida Scientist. 50(4): 264-267. [75305]
102. McCoy, Earl D.; Mushinsky, Henry R. 1995. The demography of Gopherus polyphemus (Daudin) in relation to size of available habitat. Project report: Nongame Wildlife Program--Project GFC-86-013. Tallahassee, FL: Florida Game and Fresh Water Fish Commission, Nongame Wildlife Program. 71 p. [73127]
103. McRae, W. Alan; Landers, J. Larry; Garner, James A. 1981. Movement patterns and home range of the gopher tortoise. The American Midland Naturalist. 106: 165-179. [74072]
104. Means, D. Bruce. 1982. Responses to winter burrow flooding of the gopher tortoise (Gopherus polyphemus Daudin). Herpetologica. 38: 521-525. [74071]
105. Means, D. Bruce. 1985. Management and recommendations for the gopher tortoise in longleaf pine ecosystems. In: Dodd, C. Kenneth, Jr., ed. Gopher tortoise habitat management--strategies and options: Proceedings of the 6th annual meeting of the Gopher Tortoise Council; 1985 November 15-17; Dixie, AL. Gainesville, FL: Florida Museum of Natural History; Gopher Tortoise Council: 41-56. [74132]
106. Means, D. Bruce; Grow, Gerald. 1985. The endangered longleaf pine community. ENFO. Winter Park, FL: Florida Conservation Foundation. 85(4): 1-12. [15894]
107. Mengak, Michael T.; Castleberry, Steven B. 2004. Wildlife management issues and opportunities in slash pine forests. In: Dickens, E. D.; Barnett, J. P.; Hubbard, W. G.; Jokela, E. J., eds. Slash pine: still growing and growing! Proceedings of the slash pine symposium; 2002 April 23-25; Jekyll Island, GA. Gen. Tech. Rep. SRS-76. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southern Research Station: 79-83. [55325]
108. Miller, Darren A.; Leopold, Bruce D.; Conner, L. Mike. 1999. Effects of pine and hardwood basal areas after uneven-aged silvicultural treatments on wildlife habitat. Southern Journal of Applied Forestry. 23(3): 151-157. [38474]
109. Miller, Melanie. 2000. Fire autecology. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 9-34. [36981]
110. Mitchell, Maggie. 2005. Home range, reproduction, and habitat characteristics of the female gopher tortoise (Gopherus polyphemus) in southeast Georgia. Statesboro, GA: Georgia Southern University. 97 p. Thesis. [73504]
111. Moore, William H.; Swindel, Benee F.; Terry, W. Stephen. 1982. Vegetative response to prescribed fire in a north Florida flatwoods forest. Journal of Range Management. 35(3): 386-389. [9783]
112. Morafka, David J. 1994. Neonates: missing links in the life histories of North American tortoises. In: Bury, R. B.; Germano, D. J., eds. Biology of North American tortoises. Fish and Wildlife Research Report 13. Washington, DC: U.S. Department of the Interior, National Geological Survey: 161-173. [75023]
113. Mushinsky, H. R.; Gibson, D. J. 1991. The influence of fire periodicity on habitat structure. In: Bell, S. S.; McCoy, Earl D.; Mushinsky, H. R., eds. Habitat structure: the physical arrangement of objects in space. Population and Human Biology Series. New York: Springer: 237-259. [74889]
114. Mushinsky, Henry R. 1985. Fire and the Florida sandhill herpetofaunal community: with special attention to responses of Cnemidophorus sexlineatus. Herpetologica. 41(3): 333-342. [28578]
115. Mushinsky, Henry R.; McCoy, Earl D. 1994. Comparison of tortoise populations on islands and on the mainland in Florida. In: Bury, R. B.; Germano, D. J., eds. Biology of North American tortoises. Fish and Wildlife Research Report 13. Washington, DC: U.S. Department of the Interior, National Geological Survey: 39-47. [74139]
116. Mushinsky, Henry R.; Stilson, Terri A.; McCoy, Earl D. 2003. Diet and dietary preference of the juvenile gopher tortoise (Gopherus polyphemus). Herpetologica. 59: 475-483. [74890]
117. Myers, Ronald L. 1990. Scrub and high pine. In: Myers, Ronald L.; Ewel, John J., eds. Ecosystems of Florida. Orlando, FL: University of Central Florida Press: 150-193. [17389]
118. Myers, Ronald L.; Ewel, John J, eds. 1990. Ecosystems of Florida. Orlando, FL: University of Central Florida Press. 765 p. [17384]
119. Noss, Reed F. 1988. The longleaf pine landscape of the Southeast: almost gone and almost forgotten. Endangered Species UPDATE. 5(5): 1-5. [17077]
120. Noss, Reed F.; LaRoe, Edward T., III; Scott, J. Michael. 1995. Endangered ecosystems of the United States: a preliminary assessment of loss and degradation. Biological Report 28. Washington, DC: U.S. Department of the Interior, National Biological Services. 58 p. [50483]
121. O'Meara, Timothy E.; Abbott, Michael J. 1987. Gopher tortoise response to summer burning in longleaf pine/turkey oak sandhills. Annual performance report: 1 May 1987 - 4 December 1987. Tallahassee, FL: Florida Game and Fresh Water Fish Commission, Division of Wildlife, Nongame Wildlife Section. 8 p. [62080]
122. Osentoski, M. F.; Lamb, T. 1995. Intraspecific phylogeography of the gopher tortoise, Gopherus polyphemus: RFLP analysis of amplified mtDNA segments. Molecular Ecology. 4: 709-718. [74886]
123. Ostertag, Thomas E.; Robertson, Kevin M. 2007. A comparison of native versus old-field vegetation in upland pinelands managed with frequent fire, South Georgia, USA. In: Masters, Ronald E.; Galley, Krista E. M., eds. Fire in grassland and shrubland ecosystems: Proceedings of the 23rd Tall Timbers fire ecology conference; 2005 October 17-20; Bartlesville, OK. Tallahassee, FL: Tall Timbers Research Station: 109-120. [69864]
124. Parr, Catherine L.; Andersen, Alan N. 2006. Patch mosaic burning for biodiversity conservation: a critique of the pyrodiveristy paradigm. Conservation Biology. 20(6): 1610-1619. [65366]
125. Pearson, Henry A.; Lohoefener, Renne R.; Wolfe, James L. 1987. Amphibians and reptiles on longleaf-slash pine forests in southern Mississippi. In: Pearson, Henry A.; Smeins, Fred E.; Thill, Ronald E., compilers. Ecological, physical, and socioeconomic relationships within southern national forests: Proceedings, Southern Evaluation Project workshop; 1987 May 26-27; Longbeach, MS. Gen. Tech. Rep. SO-68. New Orleans, LA: U.S. Department of Agriculture, Forest Service, Southern Forest Experiment Station: 157-165. [14733]
126. Platt, William J. 1999. Southeastern pine savannas. In: Anderson, Roger C.; Fralish, James S.; Baskin, Jerry M., eds. Savannas, barrens, and rock outcrop plant communities of North America. Cambridge; New York: Cambridge University Press: 23-51. [52459]
127. Platt, William J.; Glitzenstein, Jeff S.; Streng, Donna R. 1991. Evaluating pyrogenicity and its effects on vegetation in longleaf pine savannas. In: Proceedings, 17th Tall Timbers fire ecology conference; 1989 May 18-21; Tallahassee, FL. Tallahassee, FL: Tall Timbers Research Station: 143-161. [17606]
128. Ravdal, Stig. 2000. The effects of silviculture and prescribed burning on herpetofauna in Florida sand-pine scrub. Tampa, FL: University of South Florida. 40 p. Thesis. [62085]
129. Robbins, Louise E.; Myers, Ronald L. 1992. Seasonal effects of prescribed burning in Florida: a review. Misc. Publ. No. 8. Tallahassee, FL: Tall Timbers Research, Inc. 96 p. [21094]
130. Rostal, David C.; Jones, Douglas N., Jr. 2002. Population biology of the gopher tortoise (Gopherus polyphemus) in southeast Georgia. Chelonian Conservation and Biology. 4(2): 479-487. [74723]
131. Russell, Kevin R.; Van Lear, David H.; Guynn, David C., Jr. 1999. Prescribed fire effects on herpetofauna: review and management implications. Wildlife Society Bulletin. 27(2): 374-384. [33086]
132. Schmalzer, Paul A.; Hinkle, C. Ross. 1992. Species composition and structure of oak--saw palmetto scrub vegetation. Castanea. 57(4): 220-251. [21018]
133. Schmidt, Karl P. 1953. A checklist of North American amphibians and reptiles. 6th ed. Chicago, IL: American Society of Icthyologists and Herpetologists. 280 p. [24389]
134. Smith, Jane Kapler; Zouhar, Kristin; Sutherland, Steve; Brooks, Matthew L. 2008. Fire and nonnative invasive plants--introduction. In: Zouhar, Kristin; Smith, Jane Kapler; Sutherland, Steve; Brooks, Matthew L., eds. Wildland fire in ecosystems: fire and nonnative invasive plants. Gen. Tech. Rep. RMRS-GTR-42-vol. 6. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 1-6. [70898]
135. Smith, Lora L. 1995. Nesting ecology, female home range and activity, and population size class structure of the gopher tortoise, Gopherus polyphemus, on the Katherine Ordway Preserve, Putnam County, Florida. Bulletin of the Florida Museum of Natural History. 38 PT. I(4): 97-126. [74146]
136. Smith, Rebecca B.; Breininger, David R,; Larson, Vickie L. 1997. Home range characteristics of radiotagged gopher tortoises on Kennedy Space Center, Florida. Chelonian Conservation and Biology. 2(3): 407-417. [74140]
137. Spillers, Daniel M.; Speake, Dan W. 1988. A survey method for measuring gopher tortoise density and habitat distribution. In: Szaro, Robert C.; Severson, Kieth E.; Patton, David R., technical coordinators. Management of amphibians, reptiles, and small mammals in North America: Proceedings of the symposium; 1988 July 19-21; Flagstaff, AZ. Gen. Tech. Rep. RM-166. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 199-204. [16260]
138. Stiles, Edmund W. 1992. Animals as seed dispersers. In: Fenner, Michael, ed. Seeds: The ecology of regeneration in plant communities. 1st edition. Wallingford, UK: C.A.B. International: 87-104. [60798]
139. Stocker, Randall; Hupp, Karen V. S. 2008. Fire and nonnative invasive plants in the Southeast bioregion. In: Zouhar, Kristin; Smith, Jane Kapler; Sutherland, Steve; Brooks, Matthew L., eds. Wildland fire in ecosystems: fire and nonnative invasive plants. Gen. Tech. Rep. RMRS-GTR-42-vol. 6. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 91-112. [70903]
140. Stout, I. Jack; Marion, Wayne R. 1993. Pine flatwoods and xeric pine forests of the southern (lower) Coastal Plain. In: Martin, William H.; Boyce, Stephen G.; Echternacht, Arthur C., eds. Biodiversity of the southeastern United States: Lowland terrestrial communities. New York: John Wiley & Sons, Inc.: 373-446. [22015]
141. Stout, I. Jack; Richardson, Donald R.; Roberts, Richard E. 1989. Response of resident and relocated gopher tortoises to a prescribed burn in a sand pine scrub community. In: Diemer, Joan E.; Jackson, Dale R.; Landers, J. Larry; Layne, James N.; Wood, Don A., eds. Gopher tortoise relocation symposium: Proceedings; 1987 June 27; Gainesville, FL. Nongame Wildlife Program Technical Report #5. Tallahassee, FL: Florida Game and Fresh Water Fish Commission: 84-85. [49256]
142. Tanner, George W.; Terry, W. Steve. 1981. Effect of roller chopping and web plowing on gopher tortoise burrows in southern Florida. In: Lohoefener, Ren; Lohmeier, Lynne; Johnston, Gail, eds. The future of gopher tortoise habitats: Proceedings of the Gopher Tortoise Council; 1981 November 14; Jackson, MS. Gainesville, FL: Florida State Museum; Gopher Tortoise Council: 66-73. [74344]
143. Taylor, Robert W., Jr. 1982. Human predation on the gopher tortoise (Gopherus polyphemus) in north-central Florida. Bulletin of the Florida State Museum: Biological Sciences. 28: 79-102. [74126]
144. Tuberville, Tracey Dianne. 1998. Effects of soil disturbance by gopher tortoises (Gopherus polyphemus) on vegetation structure and composition in a pine-oak sandhills community. Athens, GA: University of Georgia. 94 p. Thesis. [73510]
145. U.S. Fish and Wildlife Service. 2013. Listed animals. In: Environmental Conservation Online System, [Online]. In: Species reports. Available: http://ecos.fws.gov/tess_public/pub/listedAnimals.jsp. [86534]
146. Van Horne, B. 1983. Density as a misleading indicator of habitat quality. Journal of Wildlife Management. 47(4): 893-901. [6902]
147. Van Lear, D. H.; Harlow, R. F. 2002. Fire in the eastern United States: influence on wildlife habitat. In: Ford, W. Mark; Russell, Kevin R.; Moorman, Christopher E., eds. The role of fire in nongame wildlife management and community restoration: traditional uses and new directions: Proceedings of a special workshop; 2000 December 15; Nashville, TN. Gen. Tech. Rep. NE-288. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northeastern Research Station: 2-10. [41533]
148. Van Lear, David H. 2000. Fire and silviculture: Recent advances in the silvicultural use of prescribed fire. In: Moser, W. Keith; Moser, Cynthia F., eds. Fire and forest ecology: innovative silviculture and vegetation management: Proceedings of the 21st Tall Timbers fire ecology conference: an international symposium; 1998 April 14-16; Tallahassee, FL. No. 21. Tallahassee, FL: Tall Timbers Research, Inc: 183-189. [37663]
149. Van Lear, David H.; Carroll, W. D.; Kapeluck, P. R.; Johnson, Rhett. 2005. History and restoration of the longleaf pine-grassland ecosystem: implications for species at risk. Forest Ecology and Management. 211(1-2): 150-165. [54058]
150. Wade, Dale D.; Brock, Brent L.; Brose, Patrick H.; Grace, James B.; Hoch, Greg A.; Patterson, William A., III. 2000. Fire in eastern ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 53-96. [36983]
151. Ware, Stewart; Frost, Cecil; Doerr, Phillip D. 1993. Southern mixed hardwood forest: the former longleaf pine forest. In: Martin, William H.; Boyce, Stephen G.; Echternacht, Arthur C., eds. Biodiversity of the southeastern United States: Lowland terrestrial communities. New York: John Wiley & Sons, Inc: 447-493. [22016]
152. Wilson, Dawn S. 1990. Home range, activity, and burrow use of juvenile gopher tortoises (Gopherus polyphemus) in a central Florida population. Tampa, FL: University of South Florida. 43 p. Thesis. [73128]
153. Wilson, Dawn S. 1991. Estimates of survival for juvenile gopher tortoises, Gopherus polyphemus. Journal of Herpetology. 25(3): 376-379. [74891]
154. Wilson, Dawn S.; Muchinsky, Henry R.; McCoy, Earl, D. 1994. Home range, activity, and use of burrows of juvenile gopher tortoises in central Florida. In: Bury, R. B.; Germano, D. J., eds. Biology of North American tortoises. Fish and Wildlife Research Report 13. Washington, DC: U.S. Department of the Interior, National Geological Survey: 147-160. [75025]
155. Wilson, Dawn S.; Mushinksy, Henry R.; McCoy, Earl D. 1991. Relationship between gopher tortoise body size and burrow width. Herpetological Review. 22(4): 122-124. [75306]
156. Wilson, Dawn S.; Mushinsky, Henry R.; Fischer, Richard A. 1997. Species profile: gopher tortoise (Gopherus polyphemus) on military installations in the southeastern United States. Strategic Environmental Research and Development Program: Final Report. Technical Report SERDP-97-10. Vicksburg, MS: U.S. Army Corps of Engineers, Waterways Experiment Station. 35 p. [73125]
157. Woodruff, Robert E. 1982. Arthropods of gopher tortoise burrows. In: Franz, Richard; Bryant, Rhoda J., eds. The gopher tortoise and its sandhill habitat: Proceedings of the third annual meeting of the Gopher Tortoise Council; 1982, October 31; Tallahassee, FL. Gainesville, FL: Florida Museum of Natural History; Gopher Tortoise Council: 24-48. [74197]
158. Wright, John Stephen. 1982. Distribution and population biology of the gopher tortoise, Gopherus polyphemus, in South Carolina. Clemson, SC: Clemson University. 74 p. Thesis. [73132]
159. Yager, Lisa Y.; Hinderliter, Matthew G.; Heise, Colleen D.; Epperson, Deborah M. 2007. Gopher tortoise response to habitat management by prescribed burning. The Journal of Wildlife Management. 71(2): 428-434. [66815]

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