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1 July 2010 Fire Ecology of the Florida Box Turtle (Terrapene Carolina Bauri Taylor) in Pine Rockland Forests of the Lower Florida Keys
Steven G. Platt, Hong Liu, Christopher K. Borg
Author Affiliations +
Abstract

Eastern box turtles (Terrapene Carolina) inhabit many fire-prone habitats in eastern North America and frequently succumb to natural and anthropogenic fires. However, little is known about the fire ecology of box turtles, and population-level effects of burning have yet to be quantified. We studied the effect of prescribed burns on a population of Florida box turtles (T. Carolina bauri) inhabiting National Key Deer Wildlife Refuge, Big Pine Key, Florida. A total of 27.4 ha were burned during seven prescribed fires (1998 to 2000). We found 14 fire-killed box turtles after four wet season burns (1.04 turtles/ha); no mortality was observed following three dry season burns. Multiple regression analysis indicated that season of burning had a significant effect on the occurrence of box turtle mortality. The effect of char height (used as a surrogate measure of fire intensity in our model) was only marginally significant. Our results suggested that between 10.2% and 21.6% of box turtles per ha perished during wet season fires. Fires appeared to affect male and female turtles equally. No juvenile mortality was observed, perhaps due to their apparent rarity in the study population. If fire mortality is a concern to land managers, we recommend restricting prescribed burns to periods of the year when box turtles are dormant (dry season in south Florida; late fall, winter, and early spring in temperate regions of North America). If prescribed burns must be conducted when turtles are active, survival might be enhanced by using slower moving, less intense backfires, and burning small areas. More frequent burning might reduce fuel loads and thus fire intensity, reducing the likelihood of turtle mortality.

INTRODUCTION

Disturbance is widely recognized as an important agent influencing the population dynamics of plants and animals (Sousa 1984; Pickett and White 1985). Fire is the most ubiquitous terrestrial disturbance after anthropogenic urban and agricultural activities, and studying the effects of fire on the biota of fire-prone ecosystems is an important topic in ecology and natural resource management (Sousa 1984; Pickett and White 1985; Whelan 1996; Russell et al. 1999). Moreover, restoring historical fire regimes in these ecosystems is desirable from the standpoint of conservation because many pyrogenic habitats have been significantly altered by decades of fire suppression or long-term changes in the seasonal timing of burning (Leach and Givnish 1996; Liu and Menges 2005). In some cases however, it is difficult to identify the historical fire regime (Snyder et al. 1990). Under such circumstances it is important to study the demographic response of native species to experimental fire regimes, as it is reasonable to expect a favorable response to the fire regime under which these taxa evolved (Menges and Kohfeldt 1995; Liu and Menges 2005). An experimental approach can, therefore, provide useful information on which to base land management decisions (Liu and Menges 2005).

The eastern box turtle, Terrapene carolina L., has an extensive distribution in the United States, ranging from southern Maine south to the Florida Keys, and west to Michigan, eastern Kansas, Oklahoma, and Texas (Ernst et al. 1994). In many areas of its geographic range, eastern box turtles occur in habitats that are prone to natural and anthropogenic wildfires or are subject to prescribed burning for land management purposes (Russell et al. 1999). Eastern box turtles have limited locomotor abilities and live among combustible leaf litter and woody debris, making them particularly vulnerable to fires (Dodd 2001). Consequently, reports of fire-related injuries and mortality are common in the literature (Babbitt and Babbitt 1951; Carr 1952; Rose 1986; Ernst et al. 1995; Dodd et al. 1997; Russell et al. 1999; Dodd 2001; Frese 2003). However, these reports consist largely of anecdotal observations that are difficult to interpret without associated estimates of population size and density (Russell et al. 1999). Indeed, the effect of fire on any eastern box turtle population has yet to be quantitatively assessed (Dodd 2001), Moreover, nothing is known about the seasonal effects of burning on fire mortality among box turtles. Dodd (2001) speculated that conducting prescribed burns during periods when box turtles were inactive might reduce fire mortality, but noted the lack of empirical data to test this hypothesis, Because long-term studies suggest that many box turtle populations are declining (Stickel 1978; Williams and Parker 1987; Schwartz and Schwartz 1991; Hall et al. 1999), understanding the various causes of mortality among populations has been accorded high priority (Maret et al. 2004).

The Florida box turtle, Terrapene Carolina bauri Taylor, one of six extant subspecies of T. Carolina in North America (Dodd 2001), is especially abundant in pine habitats of southern Florida (Carr 1952), where historically fires were a frequent form of disturbance (Taylor 1980; Snyder 1991; Carlson et al. 1993). The fire regime that existed in pine habitats of south Florida prior to Euro-American settlement is problematic and the seasonal timing of burning remains unclear (Robertson 1955; Alexander 1967; Snyder et al. 1990). Lightning was probably the primary ignition source before European settlement and lightning caused fires generally occurred during the wet season (Snyder 1991; Liu and Menges 2005). However, Native Americans inhabited the region for thousands of years (Carr and Beriault 1984), and may have burned extensively during the dry season (Snyder 1991; Spier and Snyder 1998). Studies of endemic pine rockland vegetation suggest a fire regime of lightning-caused early wet season burns and non-lightning fires during the dry season (Spier and Snyder 1998; Liu and Menges 2005).

Here, we compare fire mortality of Florida box turtles resulting from prescribed burns conducted during the wet and dry seasons in pine rockland forests on Big Pine Key, Florida. We relate our observations of fire mortality to a published estimate of population density (Verdon and Donnelly 2005), suggest management practices that might reduce fire mortality among Florida box turtles, and make inferences regarding the historic fire regimes in the lower Florida Keys.

Study Area

Our study was conducted on National Key Deer Wildlife Refuge (NKDWR; 24°42′N, 81°22′W), Big Pine Key, Monroe County, Florida. NKDWR (3110 ha) was established for the protection of the Florida Key deer, Odocoileus virginianus clavium Barbour and Allen, a federally endangered subspecies of white-tailed deer endemic to the lower Florida Keys. The refuge is characterized by extensive pine rockland forest with smaller areas of hardwood hammock and coastal mangrove forest, Pine rockland forest is a globally endangered ecosystem occurring on limestone outcrops in extreme southern Florida (Snyder et al. 1990). This ecosystem is dominated by a relatively open canopy of slash pine (Pinus elliottii Engelm.) with a diverse and often dense understory of shrubs, vines, herbs, and several species of palms (Snyder et al. 1990). The climate of the region is subtropical with a pronounced wet (May to October) and dry (November to April) season. Wet and dry season rainfall averages 743 and 265 mm, respectively, while the mean dry season temperature (22 °C) is considerably less than that of the wet season (29 °C) (Verdon and Donnelly 2005).

Pine rockland forest is a fire-dependent ecosystem; long periods (several decades) of fire exclusion reduce herb diversity and allow succession to proceed towards hardwood hammock vegetation (Robertson 1955; Alexander 1967; Snyder et al. 1990). Pine rockland forest on NKDWR consists of a mosaic of open and shrubby pinelands, probably resulting from the past occurrence of fires (Liu and Menges 2005). Open pinelands have a relatively sparse shrub layer and a well-developed herb layer, due to fairly frequent burning (Liu and Menges 2005). In contrast, shrubby pinelands have a dense shrub layer and poorly developed herb layer that results from less frequent burning (Liu and Menges 2005).

Field Sampling and Data Analysis

Our study of fire mortality among Florida box turtles took advantage of the experimental design of a larger investigation into the development of ecological criteria for prescribed burning in pine rockland forests on Big Pine Key (Liu et al. 2005a,b; Snyder et al. 2005). As part of that investigation, a block was established in each type of pineland (open and shrubby) on NKDWR during each of two years (1998 and 1999) for a total of four blocks (Dogwood, Iris, Orchid, and Poisonwood). The time elapsed since these blocks last burned ranged from 8 to 30 years (Table 1). Each of the four blocks was divided into three treatment units of 2 to 10 ha each: wet season burn, dry season burn, and unburned control. Wet season and dry season prescribed fires were replicated within and between years; however, for logistical reasons the dry season burn in Dogwood could not be conducted and the winter burn in Iris was delayed. The location and a detailed description of each block are provided elsewhere (Snyder et al. 2005). Prescribed fires occurred within 1 to 3 days of a significant rainfall event, during periods of little or no wind, and consisted of backfires and flanking fires with recorded flame lengths ranging from 0.3 to 4.5 m (Snyder et al. 2005). The maximum height of charred bark on the trunk of each canopy tree was measured in the seven treatment units following prescribed burns and used as a surrogate measure of fire intensity (Menges and Deyrup 2001; Snyder et al. 2005) in our analysis (see below).

We intensively searched seven burned areas for turtles during and immediately after prescribed fires, and conducted additional searches for dead turtles and shells during numerous post-burn visits to sample vegetation in the burned areas. Concurrent to this study, we searched control units and other unburned areas of NKDWR (1999–2000) for box turtles as part of a larger investigation into seed dispersal and wet season food habits (Liu et al. 2004; Platt et al. 2009). The straight-line carapace (CL) of each living and dead turtle was measured with calipers (± 0.1 cm). We followed Dodd et al. (1994) and considered turtles with a CL > 11.0 cm as adults. Sex was determined based on plastral morphology; males exhibit a deep concavity that is absent in females (Dodd 2001). Living turtles were permanently marked by notching a unique series of marginal scutes (Cagle 1939) and released at the capture site within 24 to 48 hours. Dead turtles were dissected and stomach contents removed as part of a concurrent study on seed dispersal (Liu et al. 2004) and wet season diet (Platt et al. 2009). If salvageable, dead turtles were deposited in the Campbell Museum, Clemson University, Clemson, South Carolina (CUSC 1750–51, 1783, 1785–86).

Table 1.

Treatment unit, area burned, date of burning, mean char height, month and year of most recent previous burn, years since last burn, and the number of fire-killed Florida box turtles (Terrapene Carolina bauri) recovered from wet and dry season burn units on National Key Deer Wildlife Refuge, Big Pine Key, Florida. Treatment units classified as open (OP) or shrubby (SH) pineland. Previous burns categorized as prescribed fire (Rx) or wildfire (WF). Fire history data are from Liu (2003).

t01_254.gif

We used a backward stepwise multiple regression model (SPSS 13.0, Chicago, Illinois) to determine if pineland type (open vs. shrubby), average char height on canopy trees, and fire season (wet season vs. dry season) had a significant effect on the occurrence of fire mortality among Florida box turtles, We did not include time elapsed since the most recent previous burn as an independent variable in our model because this variable is partially reflected in pineland type. The latter is a more comprehensive independent variable that results from both the time elapsed since the most recent previous burn and fire frequency over the last several decades. We square root transformed (√count + 1 √count + 1) the dependent variable (number of dead turtles on each burn unit) to meet the assumptions of parametric analysis (Littell et al. 2002).

RESULTS

A total of 27.4 ha were burned during four wet season and three dry season prescribed fires on NKDWR (Table 1). We found 14 fire-killed Florida box turtles (five males, nine females) after wet season burns on 13.4 ha (1.04 turtles/ha), but no mortality was observed following the dry season fires (Table 1). We expended approximately 78 person-hours searching for turtles in burned areas. The final model generated by backward stepwise multiple regression included both fire season and mean char height as explanatory variables (adjusted r2 = 0.73, F2,4 = 9.45, P = 0.03), Among these explanatory variables, coefficient of fire season was statistically significant (t = 3.7, P = 0.02), while mean char height was only marginally so (t = -2.4, P = 0.07), The number of fire-killed turtles found on each wet season treatment unit ranged from one to eight (Table 1). Eleven fire-killed turtles were recovered on the same day as the prescribed burn, and two and one were recovered two and five days, respectively, post-burn. All fire-killed turtles were adults (mean CL =14.2 cm; SE = 0.2; range =13.0 to 15.4 cm). During wet season fires, we observed four box turtles moving across fuel breaks to escape approaching flames and three others sheltering in small wetlands within the treatment unit as it was being burned, We found 217 adult and a single juvenile box turtle during approximately 68 person-hours of searching control units and other unburned habitat on NKDWR (Liu et al. 2004; Platt et al. 2009; S. Platt and H, Liu, unpubl. data).

DISCUSSION

There is little doubt that dead box turtles we recovered following prescribed burns were killed by fire. Turtle carcasses were intact, fresh, and heavily charred, strongly suggesting that burning was the cause of death. Moreover, with the exception of several weathered shells, dead box turtles were not found during an intensive search of unburned habitat on NKDWR (Platt et al. 2009; S. Platt and H. Liu, unpubl. data). Fire-killed box turtles were generally quite obvious during our immediate post-burn searches because fires removed ground vegetation, leaf litter, and clumps of dead palm fronds where turtles often shelter. Owing to the time spent intensively searching each plot immediately after burning, coupled with our later visits to sample vegetation, we believe that few if any fire-killed turtles escaped detection. Although not subjected to fire, the shells of several species of turtles remained intact for over two years under similar environmental conditions in north-central Florida (Dodd 1995). Therefore, it is likely that any fire-killed turtle we initially overlooked would have been found during our many subsequent visits to the burn units.

Our results indicate a strong seasonal effect on fire mortality among Florida box turtles; mortality only resulted from wet season fires, and dry season fires seemingly posed minimal risk to turtles. This is likely due to the seasonal activity patterns of Florida box turtles, which are most active from late April through October, but dormant and rarely found above-ground during other times of the year (Verdon and Donnelly 2005). Almost nothing is known concerning the dry season refugia used by Florida box turtles on NKDWR; but in central Florida, Dodd (2001) found dormant box turtles buried beneath leaf litter and in depressions created by decaying palm stumps. In addition to affording protection from fires, dry season retreats provide cool, moist microsites that prevent box turtles from desiccating during prolonged periods of drought (Dodd et al. 1994; Dodd 2001). Burn characteristics probably also influence turtle mortality on NKDWR; dry season burns are less intense than wet season burns, and consume less of the surface litter layer (Liu et al. 2005b; Snyder et al. 2005) where turtles are buried (Verdon and Donnelly 2005).

The estimated density of Florida box turtles on NKDWR ranges from 4.8 to 10.1 turtles/ha (Verdon and Donnelly 2005). A comparison of these densities with our estimate of fire mortality (1.04 turtles/ha) suggests that between 10.2% and 21.6% of Florida box turtles per ha perished during wet season burns. To our knowledge, this is the only population-level estimate of fire mortality available for T. Carolina. However, our estimate of fire mortality must be interpreted with caution because of the large variance associated with the population estimate of Verdon and Donnelly (2005). Owing to the small number of recaptures and the sensitivity of their model to this parameter, Verdon and Donnelly (2005) concluded the actual population size of box turtles on NKDWR may be somewhat larger than estimated. In that case, our estimate of population-level fire mortality would be correspondingly lower.

Given the current lack of demographic models (Seigel 2004), it cannot be determined how these estimated levels of mortality influence the long-term viability of Florida box turtle populations on NKDWR. Owing to a suite of life history characteristics such as delayed sexual maturity, high adult survivorship, long reproductive lifespan, and low fecundity, the viability of most turtle populations is sensitive to even small increases in mortality among older juveniles and adults (Congdon et al. 1993). Furthermore, fire losses are additive to other causes of adult mortality (e.g., roadkill, predation) and the apparently low annual recruitment of smaller size classes into this population (Verdon and Donnelly 2005), For example, our anecdotal observations suggest that the annual roadkill of adult Florida box turtles on Big Pine Key constitutes a chronic, additive, and probably significant source of mortality. Obviously, developing viability models for Florida box turtle populations is an urgent and necessary prerequisite for evaluating prescribed burning strategies on NKDWR and elsewhere, Finally, it is important to note that the prescribed burns we studied on NKDWR were limited in scale; larger burns might result in a greater proportion of the population succumbing to fire if the distance to a fuel break exceeds the locomotor abilities of turtles. Likewise, low intensity fires often produce a mosaic of burned and unburned patches, and the latter could serve as easily accessible refugia for box turtles during prescribed burns.

Previous studies of the incidence of fire-scarring among box turtle populations (Ernst et al. 1995; Dodd et al. 1997) suggest that fires do not randomly affect adults with respect to sex. Dodd et al. (1997) found fire-scarred males were larger (and probably older) than fire-scarred females, and speculated males have an increased likelihood of being burned during their longer lifespan. Female box turtles were more likely to exhibit fire-scarring than males, although the sex ratio of burned turtles did not significantly differ from the population sex ratio (Dodd et al. 1997). In contrast, Ernst et al. (1995) found no significant differences in the size of burned males and females; however, this study was based on museum specimens and may not be representative of wild populations (Dodd et al. 1997; Dodd 2001). The sex ratio of our sample of fire-killed turtles (1 male: 1.80 females) was similar to the moderately female-biased sex ratio (1 male: 1.74 females) reported for Florida box turtles on NKDWR (Verdon and Donnelly 2005), suggesting that fires affected male and female turtles equally.

Ernst et al. (1995) stated that hatchling and juvenile box turtles are more likely than adults to perish due to their habit of hiding among leaf litter, but we found no fire-killed juveniles on NKDWR. While it is possible that smaller turtles were overlooked during post-burn searching, we attribute this finding to the apparent rarity of juveniles in our study population. Verdon and Donnelly (2005) reported only six juveniles (7.5%) in a sample of 80 Florida box turtles marked on NKDWR, and there was only one juvenile among the 217 turtles (0.004%) we found on the refuge (S. Platt and H. Liu, unpubl. data).

Fire avoidance behaviors among eastern box turtles are poorly documented, Carr (1952) stated that box turtles “appear to have no notion of how to escape an advancing blaze”, Others suggest that box turtles passively escape fires by burrowing into shallow forms, leaving the carapace partially exposed above-ground (Babbitt and Babbitt 1951 ; Frese 2003), a behavior that might explain the prevalence of fire damage on neural and costal bones (Ernst et al. 1995). Florida box turtles occasionally inhabit gopher tortoise (Gopherus polyphemus) burrows (Jackson and Milstrey 1989; Dodd 2001), but it is unclear if these subterranean retreats are used to escape fires. Our observations of box turtles moving across fuel breaks and sheltering in small wetlands appear to be the only examples of active fire avoidance behaviors yet reported. These observations are not unexpected; because fire mortality is a potentially strong selective force, animals associated with pyrogenic vegetation will almost certainly evolve a suite of adaptive fire avoidance behaviors (Means and Campbell 1981; Russell et al. 1999).

Based on studies of endemic pine rockland plants, Liu and Menges (2005) suggested that the biota of the lower Florida Keys evolved under a regime of early wet season burns caused by lightning strikes coupled with non-lightning caused dry season fires. Both pineland clustervine (Jacquemontia curtisii Peter ex Hallier f.) and pineland partridge pea (Chaemacrista keyensis Pennell) exhibited a positive demographic response to dry season, but not wet season burning that was attributed to their dormancy during the dry season (Spier and Snyder 1998; Liu and Menges 2005). Furthermore, Chaemacrista keyensis exhibited a more favorable response to early wet season fires (prior to July) when compared to mid- or late-wet season fires (Liu and Menges 2005). In contrast, annual box turtle activity peaks early in the wet season (Verdon and Donnelly 2005) and, although only one of our burns occurred prior to July, fires during this period are expected to result in high levels of fire mortality. Clearly, research on additional taxa inhabiting these fire-prone ecosystems is needed to further elucidate the historic fire regime of the lower Florida Keys.

MANAGEMENT IMPLICATIONS

The results of our study lend support to an earlier recommendation for reducing fire mortality by conducting prescribed burns during periods when box turtles are inactive (Dodd 2001). Throughout most of their range in temperate North America, eastern box turtles are active only during the warmer months, generally from mid-March to early November (Neill 1948; Schwartz and Schwartz 1974; Dodd 2001). In Florida, ambient temperature is less important in determining seasonal activity patterns, which instead coincide with the annual cycle of wet and dry seasons (Dodd 2001; Verdon and Donnelly 2005). During inactive periods, eastern box turtles occupy a variety of retreats that are partially or wholly underground (Dodd 2001) and provide protection from fires. Therefore, if fire mortality of box turtles is a concern to land managers in south Florida, we recommend conducting prescribed burns during the dry season; elsewhere in eastern North America burning in the late fall, winter, and early spring would probably result in little mortality. Dry season prescribed burning is also consistent with management objectives on NKDWR that favor endemic understory herbs and forage production for Key deer (Taylor 1980; Carlson et al. 1993; Liu and Menges 2005; Liu et al. 2005a, b). Whereas the mixed fire regime of dry season and early wet season burns recommended by Liu and Menges (2005) will maintain populations of endangered endemic plants, the effects on box turtle populations are less clear and considerable mortality could result.

If burning is to be conducted during the active season, box turtle survival might be enhanced through careful fire management. For instance, our behavioral observations of box turtles during prescribed burns suggest that slower moving, less intense backfires are preferable to faster moving, more intense head fires because backfires give turtles time to move away from advancing flames and are more likely to leave unburned patches that could serve as refuges for fleeing turtles. Fast moving head fires may overtake and kill box turtles before they are able to reach safety. Furthermore, because prescribed burns reduce fuel loads and thus fire intensity (Wright and Bailey 1982; Wade and Lewis 1987), more frequent burning might reduce fire mortality among turtles, Prescribed burning can also affect wildlife abundance by altering the availability of preferred food plants (Leopold 1933; Perkins 1968; Russell et al. 1999). However, box turtles on NKDWR consume a diversity of plants (Platt et al. 2009) that respond differently to burning (Snyder et al. 2005), making generalizations regarding fire management practices difficult.

ACKNOWLEDGMENTS

Support for SGP was provided by Wildlife Conservation Society and Sul Ross State University, HL and CKB were supported by Florida International University. We are grateful for the field assistance of Joanne Springfield Globe and Elizabeth Berg. The National Key Deer Wildlife Refuge kindly provided lab space and logistic assistance. Early drafts of this manuscript benefited from the comments of Thomas Rainwater and Lewis Medlock.

LITERATURE CITED

1.

T.R. Alexander 1967. A tropical hammock on the Miami (Florida) limestone — a twentyfive-year study. Ecology 48:863–867. Google Scholar

2.

L.H. Babbitt , and C.H. Babbitt . 1951. A herpetological study in burned-over areas in Dade County, Florida. Copeia 1951:79. Google Scholar

3.

F.R. Cagle 1939. A system of marking turtles for future recognition. Copeia 1939:170–173. Google Scholar

4.

P.C. Carlson , G.W. Tanner , J.M. Wood , and S.R. Humphrey . 1993. Fire in Key Deer habitat improves browse, prevents succession, and preserves endemic herbs. Journal of Wildlife Management 57:914–928. Google Scholar

5.

A.F. Carr 1952. Handbook of Turtles: the Turtles of the United States, Canada, and Baja California. Cornell University Press, Ithaca, N.Y. Google Scholar

6.

R.S. Carr , and J.G. Beriault . 1984. Prehistoric man in south Florida. Pp. 1–14 in P.J. Gleason , ed., Environments of South Florida: Present and Past II. Miami Geological Society, Coral Gables, Fla. Google Scholar

7.

J.D. Congdon , A.E. Dunham , and R.C. van Loben Sels . 1993. Delayed sexual maturity and demographics of Blandings turtles (Emydoidea blandingii): implications for conservation and management of longlived organisms. Conservation Biology 7:826–833. Google Scholar

8.

C.K. Dodd Jr. 1995. Disarticulation of turtle shells in north-central Florida: how long does a shell remain in the woods? American Midland Naturalist 134:378–387. Google Scholar

9.

C.K. Dodd Jr. 2001. North American Box Turtles: a Natural History. University of Oklahoma Press, Norman. Google Scholar

10.

C.K. Dodd Jr. , R. Franz , and S.A. Johnson . 1997. Shell injuries and anomalies in an insular population of Florida box turtles (Terrapene Carolina bauri). Herpetological Natural History 5:66–72. Google Scholar

11.

C.K. Dodd Jr. , R. Franz , and L.L. Smith . 1994. Activity patterns and habitat use of box turtles (Terrapene Carolina bauri) on a Florida island, with recommendations for management. Chelonian Conservation and Biology 1:97–106. Google Scholar

12.

C.H. Ernst , T.P. Boucher , S.W. Sekscienski , and J.C. Wilgenbusch . 1995. Fire ecology of the Florida box turtle, Terrapene Carolina bauri. Herpetological Review 26:185–187. Google Scholar

13.

C.H. Ernst , J.E. Lovich , and R.W. Barbour . 1994. Turtles of the United States and Canada. Smithsonian Institution Press, Washington, D.C. Google Scholar

14.

P.W. Frese 2003. Tallgrass prairie amphibian and reptile assemblage. Fire mortality. Herpetological Review 34:159–160. Google Scholar

15.

R.J. Hall , P.F.P. Henry , and C.M. Bunck . 1999. Fifty-year trends in a box turtle population in Maryland. Biological Conservation 88:165–172. Google Scholar

16.

D. Jackson , and E.G. Milstrey . 1989. The fauna of gopher tortoise burrows. Pp. 86–98 in J. Diemer , D. Jackson , L. Landers , J. Layne , and D. Woods , eds., Proceedings of the Gopher Tortoise Relocation Symposium. Technical Report No. 5, Florida Game and Freshwater Fish Commission, Nongame Wildlife Program, Tallahassee. Google Scholar

17.

M.K. Leach , and T.J. Givnish . 1996. Ecological determinants of species loss in remnant prairies. Science 273:1555–1588. Google Scholar

18.

A. Leopold 1933. Game Management. Charles Scribner's Sons, New York. Google Scholar

19.

R.C. Littell , W.W. Stroup , and R.J. Freund . 2002. SAS for linear models, 4th ed. SAS Institute, Cary, N.C. Google Scholar

20.

H. Liu 2003. Population viability analyses of Chamaecrista keyensis (Leguminosae: Caesalpinioidae), a narrowly endemic herb of the Lower Florida Keys: effects of seasonal timing of fires and the urban-wildland interface., Ph.D. diss. Florida International University, Miami. Google Scholar

21.

H. Liu , and E.S. Menges . 2005. Winter fires promote greater vital rates in the Florida Keys than summer fires. Ecology 86:1483– 1495. Google Scholar

22.

H. Liu , E. Menges , and P.F. Quintata-Ascencio . 2005a. Population viability analysis of Chamaecrista keyensis — stochastic model simulations on the effect of different fire regimes. Ecological Applications 15:210–221. Google Scholar

23.

H. Liu , E. Menges , J. Snyder , S. Koptur , and M. Ross . 2005b. Effect of fire intensity on population vital rates of Chamaecrista keyensis. Natural Areas Journal 25:71–76. Google Scholar

24.

H. Liu , S.G. Platt , and C.K. Borg . 2004. Seed dispersal by the Florida box turtle (Terrapene Carolina bauri) in pine rockland forests of the lower Florida Keys, United States. Oecologia 138:539–546. Google Scholar

25.

T. Maret , J. Mitchell Jr. , and I.L. Brisbin . 2004. Research needs. Pp. 13–14 in C. Swarth and S. Hagood , eds., Summary of the Eastern Box Turtle Regional Workshop. Humane Society of the United States, Washington, D.C. Google Scholar

26.

D.B. Means , and H.W. Campbell . 1981. Effects of prescribed fire on amphibians and reptiles. Pp. 89–96 in G.W. Wood , ed., Prescribed fire and wildlife in southern forests. Belle Baruch Forest Science Institute, Clemson University, Georgetown, S.C. Google Scholar

27.

E.S. Menges , and M.A. Dreyup . 2001. Postfire survival in south Florida slash pine: interacting effects of fire intensity, fire season, vegetation, burn size, and bark beetles. International Journal of Wildland Fire 10:53–63. Google Scholar

28.

E.S. Menges , and N.M. Kohfeldt . 1995. Life history strategies of Florida scrub plants in relation to fire. Bulletin of the Torrey Botanical Club 122:282–297. Google Scholar

29.

W.T. Neill 1948. Hibernation of amphibians and reptiles in Richmond County, Georgia. Herpetologica 4:107–114. Google Scholar

30.

C.J. Perkins 1968. Controlled burning in the management of muskrats and waterfowl in Louisiana coastal marshes. Proceedings Tall Timbers Fire Ecology Conference 8:269–280. Google Scholar

31.

S.T.A. Pickett , and P.S. White . 1985. The Ecology of Natural Disturbance. Academic Press, New York. Google Scholar

32.

S.G. Platt , C. Hall , H. Liu , and C.K. Borg . 2009. Wet season food habits and intersexual dietary overlap of Florida box turtles (Terrapene Carolina bauri) on National Key Deer Wildlife Refuge, Florida. Southeastern Naturalist 8:335–346. Google Scholar

33.

W.B. Robertson 1955. An analysis of breeding bird populations of tropical Florida in relation to the vegetation., Ph.D. diss. University of Illinois, Champaign. Google Scholar

34.

F.L. Rose 1986. Carapace regeneration in Terrapene (Chelonia: Testudinidae). Southwestern Naturalist 31:131–134. Google Scholar

35.

K.B. Russell , D.H. van Lear, and D.C. Guynn Jr. 1999. Prescribed fire effects on herpetofauna: review and management implications. Wildlife Society Bulletin 27:374–384. Google Scholar

36.

E.R. Schwartz , and C.W. Schwartz . 1974. The three-toed box turtle in central Missouri: its population, home range, and movements. Missouri Department of Conservation, Terrestrial Series 5:1–28. Google Scholar

37.

E.R. Schwartz , and C.W. Schwartz . 1991. A quarter-century study of survivorship in a population of three-toed Box Turtles in Missouri. Copeia 1991:1120–1123. Google Scholar

38.

R.A. Seigel 2004. The importance of population demography in the conservation of box turtles: what do we know and what do we need to learn? Pp. 6–7 in C. Swarth and S. Hagood , eds., Summary of the Eastern Box Turtle Regional Workshop. Humane Society of the United States, Washington, D.C. Google Scholar

39.

J.R. Snyder 1991. Fire regimes in subtropical south Florida. Proceedings Tall Timbers Fire Ecology Conference 17:303–319. Google Scholar

40.

J.R. Snyder , A. Herndon , and W.B. Robertson . 1990. South Florida rockland. Pp. 230–277 in R.L. Myers and J.J. Ewel , eds., Ecosystems of Florida. University of Central Florida Press, Orlando. Google Scholar

41.

J.R. Snyder , M.S. Ross , S. Koptur , and J.P. Sah . 2005. Developing ecological criteria for prescribed fire in south Florida pine rockland ecosystems. Open File Report: OF 2006-1062, U.S. Geological Survey, Washington, D.C. Google Scholar

42.

W.P. Sousa 1984. The role of disturbance in natural communities. Annual Review of Ecology and Systematics 15:353–391. Google Scholar

43.

L.P. Spier , and J.R. Snyder . 1998. Effects of wet- and dry-season fires on Jacquemontia curtisii, a south Florida pine forest endemic. Natural Areas Journal 18:350–357. Google Scholar

44.

L.F. Stickel 1978. Changes in a box turtle population during three decades. Copeia 1978:221–225. Google Scholar

45.

D.L. Taylor Fire history and man-induced fire problems in subtropical south Florida. Pp. 63–68 in M. A. Stokes and J.H. Dietrich , eds., Proceedings of the Fire History Workshop. General Technical Report RM-81, U.S. Department of Agriculture, Forest Service, Ft. Collins, Colo. Google Scholar

46.

E. Verdon , and M.A. Donnelly . 2005. Population structure of Florida box turtles (Terrapene Carolina bauri) at the southernmost limit of their range. Journal of Herpetology 39:572–577. Google Scholar

47.

D.D. Wade , and C.E. Lewis . 1987. Managing southern grazing ecosystems with fire. Rangelands 9:115–119. Google Scholar

48.

R.J. Whelan 1996. The Ecology of Fire. Cambridge University Press, Cambridge, U.K. Google Scholar

49.

E.C. Williams , and W.S. Parker . 1987. A longterm study of a box turtle (Terrapene Carolina) population at Allee Memorial Woods, Indiana, with emphasis on survivorship. Herpetologica 43:328–335. Google Scholar

50.

H.A. Wright , and A.W. Bailey . 1982. Fire Ecology. J. Wiley, New York. Google Scholar
Steven G. Platt, Hong Liu, and Christopher K. Borg "Fire Ecology of the Florida Box Turtle (Terrapene Carolina Bauri Taylor) in Pine Rockland Forests of the Lower Florida Keys," Natural Areas Journal 30(3), 254-260, (1 July 2010). https://doi.org/10.3375/043.030.0301
Published: 1 July 2010
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