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Vegetative Characteristics of Gopher Tortoise (Gopherus polyphemus) Habitat on the Lower Suwannee National Wildlife Refu...

University of Florida Institutional Repository

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VEGETATIVE CHARACTERISTICS OF GOPHER TORTOISE ( Gopherus polyphemus) HABITAT ON THE LOWER SU WANNEE NATIONAL WILDLIFE REFUGE: IMPLICATIONS FOR RESTOR ATION AND MANAGEMENT OF PINE COMMUNITIES By STEPHEN E. BARLOW A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLOR IDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2004

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Copyright 2004 by Stephen E. Barlow

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This document is dedicated to my father the late Douglas Alvor d Barlow, Dad, mentor and friend.

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ACKNOWLEDGMENTS I wish to thank the members of my graduate supervisory committee for their invaluable input and guidance: Dr. Mark W. Clark, chairman and research assistant professor, Soil and Water Sciences Department; Dr. George W. Tanner, co-chairman and professor, Department of Wildlife Ecology and Conservation; and Mr. Kenneth Litzenberger, Refuge Manager, Lower Suwannee National Wildlife Refuge. I also would like to thank the following individuals: my wife, Elizabeth, for her strong support, patience and encouragement; my Mom, Kay, for instilling within me a love of nature; my in-laws, Joe and Jane Works, for constant support and encouragement; Russ Singleton for helping with gopher tortoise burrow surveys; Vivian R. Soriero, Linda Casey and John C. Jones for assisting with vegetation surveys; Joan Berish with the Florida Fish and Wildlife Conservation Commission for assistance throughout the project; Dr. Lori Wendland, University of Florida College of Veterinary Medicine, for testing tortoise blood samples; Daniel Barrand for assistance with GIS analysis; Kenneth W. McCain for providing fire and management history information, and all employees of the Lower Suwannee National Wildlife Refuge. iv

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TABLE OF CONTENTS page ACKNOWLEDGMENTS .................................................................................................iv LIST OF TABLES .............................................................................................................vi LIST OF FIGURES .........................................................................................................viii ABSTRACT .......................................................................................................................ix INTRODUCTION ...............................................................................................................1 MATERIAL AND METHODS ...........................................................................................7 Study Area ....................................................................................................................7 Survey Compartments ..................................................................................................7 Vegetation Surveys .....................................................................................................15 Statistical Analysis ......................................................................................................19 RESULTS..........................................................................................................................20 Vegetation Surveys.....................................................................................................20 Chi-Square Analysis...................................................................................................25 DISCUSSION....................................................................................................................40 High Pine Habitats......................................................................................................40 Pine Flatwoods Habitats.............................................................................................42 MANAGEMENT IMPLICATIONS.................................................................................44 APPENDIX LITERATURE CITED......................................................................................................62 BIOGRAPHICAL SKETCH.............................................................................................67 v

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LIST OF TABLES Table page 1. Habitats and relative burrow densities within burrow survey compartments on the Lower Suwannee NWR, Florida..............................................................................18 2. Descriptive statistics of vegetation cover for each habitat and burrow density category on the Lower Suwannee NWR, Florida...................................................................21 3. Chi-square tests of independence between canopy cover and shrub cover within the 2 habitat categories on the Lower Suwannee NWR, Florida......................................30 4. Chi-square tests of independence between canopy cover and herbaceous ground cover within the 2 habitat categories on the Lower Suwannee NWR, Florida..................31 5. Chi-square tests of independence between canopy cover and woody ground cover within the 2 habitat categories on the Lower Suwannee NWR, Florida..................32 6. Chi-square tests of independence between canopy cover and bare ground cover within the 2 habitat categories on the Lower Suwannee NWR, Florida.............................33 7. Chi-square tests of independence between canopy cover and detritus ground cover within the 2 habitat categories on the Lower Suwannee NWR, Florida..................34 8. Chi-square tests of independence between shrub cover and herbaceous ground cover within the 2 habitat categories on the Lower Suwannee NWR, Florida..................35 9. Chi-square tests of independence between shrub cover and woody ground cover within the 2 habitat categories on the Lower Suwannee NWR, Florida..................37 10. Chi-square tests of independence between shrub cover and bare ground cover within the 2 habitat categories on the Lower Suwannee NWR, Florida.............................38 11. Chi-square tests of independence between shrub cover and detritus ground cover within the 2 habitat categories on the Lower Suwannee NWR, Florida..................39 12. Analysis of variance results for high pine canopy cover comparisons......................50 13. Analysis of variance results for high pine shrub cover comparisons.........................51 14. Analysis of variance results for high pine woody ground cover comparisons..........52 vi

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16. Analysis of variance results for high pine detritus ground cover comparisons.........54 17. Analysis of variance results for high pine herbaceous ground cover comparisons...55 18. Analysis of variance results for pine flatwoods canopy cover comparisons.............56 19. Analysis of variance results for pine flatwoods shrub cover comparisons................57 20. Analysis of variance results for pine flatwoods woody ground cover comparisons..58 21. Analysis of variance results for pine flatwoods bare mineral soil cover comparisons.59 22. Analysis of variance results for pine flatwoods detritus ground cover comparisons.60 23. Analysis of variance results for pine flatwoods herbaceous ground cover comparisons..............................................................................................................61 vii

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LIST OF FIGURES Figure page 1. Location of the Lower Suwannee NWR, Florida..........................................................8 2. Land cover forms on the Lower Suwannee NWR, Florida...........................................9 3. Location of tortoise burrow and vegetation survey compartments, Lower Suwannee NWR, Florida...........................................................................................................10 4. Soil composition of high pine survey compartment categories on the Lower Suwannee NWR, Florida..........................................................................................12 5. Soil composition of pine flatwoods survey compartment categories on the Lower Suwannee NWR, Florida..........................................................................................14 6. Representative pictures of each survey compartment category...................................16 7. Comparisons of canopy cover between burrow densities within each of the pine community types on the Lower Suwannee NWR, Florida.......................................22 8. Comparisons of shrub cover between burrow densities within each of the pine community types on the Lower Suwannee NWR, Florida.......................................23 9. Comparisons of herbaceous ground cover between burrow densities within each of the pine community types on the Lower Suwannee NWR, Florida.........................24 10. Comparisons of woody ground cover between burrow densities within each of the pine community types on the Lower Suwannee NWR, Florida...............................26 11. Comparisons of bare soil ground cover between burrow densities within each of the pine community types on the Lower Suwannee NWR, Florida...............................27 12. Comparisons of detritus ground cover between burrow densities within each of the pine community types on the Lower Suwannee NWR, Florida...............................28 viii

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Abstract of Thesis Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science VEGETATIVE CHARACTERISTICS OF GOPHER TORTOISE (Gopherus polyphemus) HABITAT ON THE LOWER SUWANNEE NATIONAL WILDLIFE REFUGE: IMPLICATIONS FOR RESTORATION AND MANAGEMENT OF PINE COMMUNITIES By Stephen E. Barlow December 2004 Chair: Mark W. Clark Major Department: Soil and Water Science Previous land use practices on the Lower Suwannee National Wildlife Refuge altered natural fire conditions resulting in habitat degradation, especially within the high pine areas, allowing succession to begin shifting these habitats to mesic hammocks. In addition, many pine flatwoods sites exist as densely planted slash pine plantations, which have been relegated to an infrequent winter fire regime, leading to a shrub level monoculture of gallberry (Ilex glabra) and saw palmetto (Serenoa repens). Changes in vegetative structure of these prominent pine communities are thought to have lowered the quality of gopher tortoise habitat. This investigation evaluates the relationship between vegetative cover of different community strata and the density of gopher tortoise burrows. Vegetation cover was measured on sites with high and low gopher tortoise burrow densities in high pine and pine flatwoods habitats on the refuge from April to July 2004. ix

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The line intercept technique was used to measure canopy cover, shrub cover and ground cover. Survey compartment soil compositions were compared using Geographic Information Systems (GIS) data and the county soil surveys. It was predicted that compartments with high tortoise burrow densities would have a relatively open canopy and shrub layers, with relatively high herbaceous and bare soil ground covers. Analysis of variance on vegetation cover data revealed that density of the canopy, shrub layer, and ground cover types were significantly different between high pine sites with different tortoise burrow densities. While the pine flatwoods sites exhibited significant differences in all categories except bare soil, due to a lack of bare soil within these habitats. Lower canopy cover, shrub cover and higher herbaceous ground cover characterized sites with high tortoise burrow densities. Soil analysis revealed higher soil series diversity within the pine flatwoods sites. This soil series diversity is not represented vegetatively within the current slash pine plantation monoculture. The lack of relationship between soil characteristics and vegetation community composition is most likely due to past timber practices that converted all usable land to pine production with little regard for preserving small pockets of habitat. Close analysis of soil series should guide land managers efforts in restoration of habitats in order to regain pre-European development habitat diversity. Aggressive techniques to reverse succession on high pine sites are also recommended. Restoration of the herbaceous ground layer and a reintroduction to summer fires could then be employed. Timber thinning and a more varied fire regime emphasizing summer burning would benefit pine flatwoods sites; otherwise continued habitat succession and degradation will significantly inhibit tortoise populations. x

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INTRODUCTION Once the most extensive terrestrial ecosystems in Florida, pine flatwoods and high pine communities have been heavily influenced by humans, therefore many characteristics of these communities have changed markedly since European settlement. Pine communities were extensively timbered starting during the Civil War through today (Abrahamson and Harnett 1990). Concurrently, human population growth and the concomitant fragmentation of these vast pine stands have led to a decrease in the extent and frequency of natural fires. In their natural state, these communities are characterized by their openness and frequent occurrence of fires (Laessle 1942, Ober 1954, Edmisten 1963, Platt et al. 1988). Though few natural stands closely resemble pre-settlement pine communities, the consensus is that present stands differ from pre-settlement stands by having lower fire frequencies, more even age structure, and a denser under-story with greater shrub cover and less herbaceous cover (Abrahamson and Hartnett 1990, Noel et al. 1998). In recent times many of these areas have been owned and managed by timber companies that practice intense plantation production of short rotation pine trees for the paper industry. Thus, managers of many public lands in Florida are often faced with restoring and managing pine communities that have been severely altered. Characteristics of pine communities under timber production management and fire exclusion include densely planted even-aged stands of slash pine (Pinus eliotti) often planted in raised, 1

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2 mechanically bedded rows, with intense herbicide use. Forest stands are also often prepared using heavy machinery, they have a monotypic shrub under story, raised road beds, windrows, borrow pits, rail tramways and logging roads crisscrossing the forest. An understanding of the impacts of these former land use practices is necessary for the formulation of management strategies concerned with restoring habitats to their pre-European settlement state. Numerous researchers have described the gradual transition high pine communities make to mesic hardwood forests and the invasion of non fire-adapted species in the absence of frequent fire (Veno 1976, Givens et al. 1984, Myers 1985, Myers and White 1987). With the exclusion of fire, fire-adapted species also decline and since they often serve as the primary fuel source for frequent fires frequency and intensity of natural, lightning-ignited fires is reduced. Under natural conditions pine flatwoods are stable and essentially nonsuccessional due to fire (Abrahamson and Hartnett 1990). When fire is removed from pine flatwoods, or if the natural frequency or seasonality of fire is altered, flatwoods can succeed to a variety of vegetation types. Human modifications to the landscape, such as certain silvicultural practices (logging, clearing and drainage), can also stimulate successional change. It has been noted by numerous authors that disturbance of the natural fire frequency is the most common cause of successional changes in pine flatwoods (Monk 1968, Robbins and Myers 1992, Peroni and Abrahamson 1986). Loss of pine communities along with fire exclusion and certain silvicultural practices have negatively impacted numerous wildlife species that characterize pine communities. Bird species that require open pine forests such as the red-headed

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3 woodpecker (Melanerpes erythrocephalus), brown-headed nuthatch (Sitta pusilla), loggerhead shrike (Lanius ludovicianus) and eastern bluebird (Sialia sialis) have experienced declining populations throughout the southeast (Cox 1987). Within high pine communities, population size and species richness of birds decline noticeably when these areas are converted to timber production (Repenning and Labisky 1985). Silviculture practices across the United States have been implicated in the decline or elimination of at least 26 species of salamanders including the flatwoods salamander (Ambystoma cingulatum) in Florida (Bury 1983, Brode and Bury 1984, Herrington and Larsen 1985, Corn and Bury 1989, Welsh 1990, Blymer and McGinnes 1977, Ash 1988, Buhlmann et al. 1988, Pentranka et al. 1993, Jordan and Mount 1975, Dodd 1991, Dodd 1993, Means et al. 1996). Due to loss and degradation of habitat, numerous species found within pine communities are currently listed as species of special concern, threatened or endangered. One such species is the gopher tortoise (Gopherus polyphemus), which is a state listed species of special concern that has experienced a population reduction through loss of habitat and a lack of fire in existing habitats. Changes in fire frequency in pine communities are thought to have decreased herbaceous food plants, thereby negatively influencing gopher tortoise habitat (Landers and Speake 1980, Cox et al. 1987). Since grasses and forbs constitute the bulk of the gopher tortoises diet, increased shading and detritus buildup associated with fire exclusion lead to reduced productivity of these plants and a decline in tortoise numbers (Garner and Landers 1979, Franz and Auffenberg 1974).

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4 The gopher tortoise is a keystone speciesone that holds a critical role in the ecosystem (Campbell and Christman 1982, Jackson and Milstrey 1988). Their burrows provide shelter for more than 300 species of obligate and facultative commensals, including arachnids, insects, reptiles, amphibians, birds, and mammals (Jackson and Milstrey 1988). Several of these species such as the southeastern indigo snake (Drymarchon couperi), the gopher frog (Rana capito) and the Florida mouse (Podomys floridanus) are threatened species or species of special concern. Decline of the gopher tortoise has negatively affected these secondary burrow users because there are fewer burrows available (Berish 2001). A primary factor affecting density of gopher tortoises is habitat quality, particularly as it relates to food availability as influenced by fire and primary succession (OMeara and Abbott 1987). However, caution should be used when comparing tortoise densities to habitat variables as dispersion of gopher tortoise burrows within available habitats is poorly understood (Cox et al. 1987). To more accurately determine the quality of habitat measures of mean reproductive success, survival and number of individuals in each age class are needed to determine quality of the habitat (Van Horne 1983). Poor habitat quality may cause tortoises to form dense colonies in small patches of suitable habitat, thus a survey which is conducted at too small of a scale, or multiple scales may over estimate tortoise densities across the habitat or flaw site comparisons (McCoy and Mushinsky 1995). To better understand the status of tortoise populations on the Lower Suwannee National Wildlife Refuge Refuge (LSNWR) and their habitat preferences, we conducted tortoise burrow surveys in 2002 in a manner similar to that of Auffenberg and Franz

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5 (1982) and in Cox et al. (1987). In this survey, burrows were recorded as active, inactive and old. Active burrows were marked using a handheld GPS unit. Marked burrows were then projected as a layer file using Arcview 8.3 geographic information systems (GIS) software to compare burrow colony locations with habitat and soil features. Through GIS analysis it appeared refuge tortoises were congregated in colonies within various survey compartments. This aggregating of tortoises into pockets suggests preferential habitat within the broader survey compartment possibly representing remnants of formerly large areas of habitat (McCoy and Mushinsky 1995). This clumping of tortoise burrows makes the task of density estimation across refuge habitats imperfect at best, unless additional environmental variables influencing tortoise distribution can be identified. Furthermore, tortoise burrow congregations did not appear to relate to particular GIS habitat or soil layers, thus the level of detail within the GIS data layers were lacking for discernment of these apparent site choices by tortoises. The goal of this study was to investigate what additional environmental variables might be influencing tortoise distribution and what changes in management might be implemented to enhance tortoise habitat in pine communities on the LSNWR. Specific hypothesis to be addressed under this goal include the following: 1) To determine if differences are present between vegetation structural characteristics (ground cover, shrub cover and canopy cover) among tortoise burrow survey compartments that exhibited relatively high or low burrow densities. 2) To determine if a relationship is present between the amount of canopy and shrub cover to ground cover forms or if these variables are independent. It was predicted that areas with low tortoise burrow density would exhibit high canopy and shrub cover with low herbaceous ground coverage, while areas with high tortoise burrow densities would contain structural characteristics of low canopy and shrub

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6 cover with higher herbaceous ground coverage. It was further predicted that the amount of ground cover forms observed would be dependent upon the amount of shrub or canopy cover present.

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MATERIAL AND METHODS Study Area The study was conducted on the Lower Suwannee NWR, a 21,500-hectare refuge located in the heart of north Floridas Big Bend region in Dixie and Levy counties (Figure1). Situated along the Gulf of Mexico the refuge is bisected by the Suwannee River, which flows southward through the refuge for 32 kilometers before emptying into the Gulf of Mexico. Purchased by the federal government in 1979 from various timber companies, the area still bears signs from decades of timber production management. The topography of the study area is relatively flat, with dominant habitats including salt marsh, southern hydric hardwood, bottomland forest, and pine plantation (Figure 2) (Sykes et al. 1999). This diversity of habitats on the area is matched by the variety of soil orders found on the refuge. Of the seven soil orders in Florida, the refuge contains six, with Histosols being the most common (Slabaugh et al. 1996, Liudahl et al. 2003). The refuge contains 43 individual soil series, many of which are poorly drained or very poorly drained and strongly acid (pH 5.1-5.5) to extremely acid (pH 4.0-4.4) (Slabaugh et al. 1996, Liudahl et al. 2003). Survey Compartments Survey transects selected for this study were evenly divided between high pine habitats and pine flatwood habitats as well as by tortoise burrow density (Figure 3). Ten transects were randomly selected from high pine habitats: 5 transects in high tortoise 7

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8 Figure 1. Location of the Lower Suwannee NWR, Florida.

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9 Figure 2. Land cover forms on the Lower Suwannee NWR, Florida.

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10 Figure 3. Location of tortoise burrow and vegetation survey compartments, Lower Suwannee NWR, Florida.

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11 burrow density compartments and 5 transects within low tortoise burrow density compartments. Similarily10 transects were randomly selected from pine flatwoods compartments, also equally divided by tortoise burrow density. High pine survey compartments consisted of even-aged mechanically planted long-leaf pines, planted in 1993 (Figure 6). Thus, the species, age, and basal area of the planted pine canopy component were controlled for between burrow density compartments. Interspersed within the planted long-leaf pines were turkey oak (Quercus laevis), live oak (Quercus virginiana) and a shrub layer containing immature oaks (Quercus spp.), sparkleberry (Vaccinium arboreum) and dog fennel (Eupatorium compositifolium). Herbaceous ground cover within the compartments contained deers tongue (Carphephorus odoratissimus), greenbriar (Smilax spp.), partridge pea (Chamaecrista pilosa) wire grass and associated forbs and grasses. The refuge practices an aggressive prescribed burning program, introducing fire to approximately 600 hectares annually (pers. comm., K. McCain Chiefland, FL). With the majority of these being dormant season burns, within densely planted flatwoods habitats. Though occasional attempts have been made to burn within the high pine habitats, lack of a uniformly distributed fuel source throughout has made most of theses burns spotty and anemic at best (pers. comm., K. McCain Chiefland, FL). Following the planting of long-leaf pines on these areas, a lack of fire for at least 10 years has led to the aforementioned encroachment of non-pyric species such as oaks. Needle cast from the planted pine component within these areas currently does not provide adequate fuels to carry fire under safe weather conditions. Also, with the current tree species composition

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12 HIGH PINEHIGH BURROW DENSITY SOILS31%69% Ridgewood Ortega A HIGH PINE LOW BURROW DENSITY SOILS29%35%36% Ridgewood Leon (Spodosol) Clara (Spodosol) B Figure 4. Soil composition of high pine survey compartment categories on the Lower Suwannee NWR, Florida. A) Soil composition of high pine compartments with high tortoise burrow densities. B) Soil composition of high pine compartments with low tortoise burrow densities.

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13 and densities, the chances of an intense, unmanageable scrub fire killing all the trees is a serious management concern. Soils within high pine survey compartments were represented by the Clara and Leon series (Figure 4). All soils are very deep, moderately well drained soils of upland sites (Liudahl et al. 2003). Pine flatwoods survey compartments consisted of even-aged slash pines, planted in 1978 by Buckeye Cellulose Inc., (Figure 6). Thus, the species, age, and basal area of the planted pine canopy component were controlled for between burrow density compartments. Isolated cypress (Taxodium distichum) wetlands are widely interspersed within these habitats but were not included within survey compartments. Years of fire exclusion followed by only dormant season fires has helped to foster the dense stands of gallberry and saw-palmetto common within refuge pine flatwoods communities, inhibiting production of herbaceous ground covers (forbs and grasses). A lack of growing season burns could facilitate the establishment of dense gallberry stands, since growing season burns appear to hinder gallberrys resprouting ability, while time since the last fire has been shown to increase the density of saw palmetto (Hughes and Knox 1964, Maliakal et al. 2000). The sites were bedded for planting, but other management techniques conducted on these sites by the timber company is unknown. The U.S. Fish and Wildlife Service assumed management of the stands in 1985, since refuge acquisition the sites have received several dormant season prescribed burns. The plantations also received third row thinnings in 1997-98.

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14 Pine Flatwoods High Burrow Density Soils 34%2%9%49%6% Smyrna (Spodosol) Placid (Inceptisol) Adamsville (Entisol) Ridgewood (Entisol) Leon (Spodosols) A Pine Flatwoods Low Burrow Density Soils34%10%26%0%30% Chaires (Spodosol) Resota (Entisol) Ridgewood (Entisol) Bodiford (Alfisol) Leon (Spodosol) B Figure 5. Soil composition of pine flatwoods survey compartment categories on the Lower Suwannee NWR, Florida. A) Soil composition of pine flatwoods compartments with high tortoise burrow densities. B) Soil composition of pine flatwoods compartments with low tortoise burrow densities.

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15 Soil composition of pine flatwoods survey compartments contained primarily Spodosols, though Entisols, Inceptisols, and Alfisols were also present (Figure 5). The soils of flatwoods survey compartments were less uniform and more interspersed than high pine compartments. Spodosols were represented by Smyrna, Chaires and Leon series, which have similar characteristics of being deep to very deep and poorly drained. While Entisols were represented by Adamsville, Ridgewood and Resota series which are very deep, moderately drained psamments. Inceptisols and Alfisols occupied only 0.33 hectares within the flatwoods compartments and were represented by Bodiford and Placid soil series (Slabaugh et al. 1996, Liudahl et al. 2003). Vegetation Surveys Vegetation surveys were conducted from April through July 2004. Canopy cover, shrub cover and ground cover were determined within survey compartments that had exhibited relatively high or low tortoise burrow densities during the 2002 tortoise burrow surveys. A line-intercept technique was used to estimate percent canopy cover and shrub cover along each transect (Higgins et al. 1994). A 10-meter line was stretched between two stakes at a height of 1-meter. Percent canopy cover was measured by observing the total length of line intercepted by vertical projections of the canopy. Percent shrub cover was measured by observing the total length of line intercepted by plants touching the line. Only shrubs touching the nylon line were included, whereas the entire plant intercepting the vertical line was counted as percent cover. Percent ground cover was obtained using the pen-drop technique, where a pen was dropped at 1-meter increments along the line and the dominant ground cover touching the pen was recorded. Ground cover forms were grouped at each sampling location into the following 4 categories: (1) Woody (i.e., woody shrubs, tree seedlings), (2) Herbaceous (i.e., non-woody plants, legumes, forbs

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16 B A C D Figure 6. Representative pictures of each survey compartment category: A) High pine with high tortoise burrow density, B) High pine with low tortoise burrow density, C) Pine flatwoods with high tortoise burrow density and D) Pine flatwoods with low tortoise burrow density.

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17 and grasses), (3) Detritus (i.e., non-living plant material, leaves, needles) and (4)Bare(i.e., mineral soil). Vegetation surveys were conducted along the same transects used during the 2002 tortoise burrow surveys. Tortoise burrow survey transects were 200-meters long by 20-meters wide, covering approximately .4 hectares. Survey transects were grouped in compartments, which were placed within suitable tortoise habitats using the refuges GIS habitat and soils layers (Cox et al. 1987, Breininger et al. 1988). Following the 2002 tortoise burrow survey, compartments were labeled either high burrow density or low burrow density; compartments were also arranged within 2 habitat categoriespine flatwoods or high pine, see Table 1 (Abrahamson and Hartnett 1990, Myers 1990). Burrow densities for each compartment were determined by dividing the compartment area by the total number of both active and inactive burrows observed, burrow densities >1burrow/hectare were designated as high density while those compartments with < 1burrow/hectare were identified as low density compartments. Using a random numbers table, survey transects within 8 distinct compartments were randomly selected for vegetation cover surveys (Ott 1983). Five survey transects were selected from each habitat and burrow density category, (i.e., 5 transects taken from high pine/high burrow density compartments were compared with 5 transects taken from high pine/low burrow density compartments, while 5 transects were taken from flatwoods pine/high burrow density compartments were compared with 5 transects taken from flatwoods pine/low burrow density compartments). Since all survey compartments had been mechanically planted as even-aged stands, I was able to control for timber basal area, age class and species. Compartments also contained

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18 Table 1. Habitats and relative burrow densities within burrow survey compartments on the Lower Suwannee NWR, Florida as determined during the 2002 survey. Compartment/Transects Habitat Burrows/Hectare Burrow Density 1/1-20* Pine Flatwoods 1.13 High 1/21-30 Pine Flatwoods 1.00 Low 1/31-40 Pine Flatwoods 0.50 Low 1/41-50 Pine Flatwoods 1.25 High 1/51-60 Pine Flatwoods 0.25 Low 1/61-70 Pine Flatwoods 0.00 Low 2/1-10 Pine Flatwoods 1.00 Low 2/11-17 Pine Flatwoods 0.00 Low 3/1-6 Pine Flatwoods 0.00 Low 4/1-16 Pine Flatwoods 0.94 Low 4/17-36 Pine Flatwoods 0.88 Low 4/37-53 Pine Flatwoods 0.74 Low 6/1-15 Pine Flatwoods 0.00 Low 6/16-25 Pine Flatwoods 0.00 Low 7/1-10* Pine Flatwoods 0.00 Low 7/11-16* Pine Flatwoods 0.00 Low 8/1-15* Pine Flatwoods 3.50 High 8/16-25* Pine Flatwoods 3.50 High 9/1-20* High Pine 4.25 High 9/21-35* High Pine 5.17 High 9/36-45 High Pine 1.00 Low 9/46-55 High Pine 2.50 High 9/56-61* High Pine 0.83 Low 9/62-70 High Pine 0.83 Low *Compartments containing randomly selected vegetation surveys.

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19 very similar burn histories; with no compartments receiving a growing season burn prior to this study. Vegetation cover percentages were recorded at 20-meter intervals along the 200-meter burrow survey transects. At 20-meter intervals a 10-meter vegetation cover transect was surveyed perpendicular to and on both sides of the burrow survey transect. This provided 20, 10-meter vegetation cover transects spaced throughout each burrow survey transect. Statistical Analysis Microsoft Excel 2000 was used for statistical analysis. I arcsine transformed percentages (i.e., percent canopy cover, shrub cover and ground cover) prior to analysis so they would better meet the normality assumption of analysis of variance (Ott 1983). The assumption of homogeneity of population variances was not considered critical since the sample sizes were equal. I considered a 0.05 probability level statistically significant for all tests. A chi-square test of independence was used to determine if the amount of the 4 ground cover forms were independent of the amount of shrub or canopy cover observed. For the chi-square test of independence canopy, shrub and the 4 ground cover forms data was grouped into 2 categories: High for values > 0.5, and Low for values < 0.5. Data represented in descriptive statistics or used for the chi-square test was not transformed.

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RESULTS Vegetation Surveys Analysis of canopy closure data indicates significant differences when comparing between burrow densities within each of the pine community types (Figure 6). Canopy cover differed significantly between high pine burrow density category sites ( P =0.000). Canopy cover also differed between pine flatwoods high and low tortoise burrow density sites ( P =0.000). Within both habitat categories, low tortoise burrow density sites had greater canopy closure than respective high tortoise burrow density sites. The mean values prior to transformation, for each habitat and tortoise burrow density category can be found in table 2. Shrub cover within the two pine communities also differed significantly between high pine tortoise burrow density category sites ( P =0.000). Shrub cover also differed between pine flatwoods high and low tortoise burrow density sites ( P =0.000). Within both habitat categories, low tortoise burrow density sites had greater shrub cover densities than respective high tortoise burrow density sites (Figure 7). Herbaceous ground cover differed significantly between high pine tortoise burrow density category sites ( P =0.000). Herbaceous ground cover also differed between pine flatwoods high and low tortoise burrow density sites ( P =0.000). Within both habitat categories high tortoise burrow density sites had greater herbaceous ground cover densities than respective low tortoise burrow density sites (Figure 8). 20

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21 Table 2. Descriptive statistics of vegetation cover for each habitat and burrow density category on the Lower Suwannee NWR, Florida. HIGH PINE Canopy Shrub Herb Woody Bare Detritus Burrow Cover Cover Cover Cover Soil Cover Density Mean SD Mean SD Mean SD Mean SD Mean SD Mean SD High(n=100) 0.12 0.13 0.10 0.12 0.43 0.23 0.13 0.14 0.26 0.19 0.17 0.21 95% CI 0.03 0.02 0.05 0.03 0.04 0.04 Low(n=100) 0.43 0.03 0.72 0.22 0.14 0.14 0.43 0.20 0.07 0.11 0.36 0.22 95% CI 0.05 0.04 0.03 0.04 0.02 0.04 PINE FLATWOODS High(n=100) 0.28 0.28 0.11 0.15 0.53 0.30 0.23 0.18 0.00 0.02 0.20 0.21 95% CI 0.05 0.03 0.06 0.03 0.00 0.04 Low (n=100) 0.52 0.30 0.81 0.12 0.19 0.16 0.51 0.22 0.00 0.00 0.30 0.22 95% CI 0.06 0.02 0.03 0.04 0.00 0.04

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22 High Pine High Pine Pine Flatwoods Pine FlatwoodsHigh Burrow Low Burrow High Burrow Low BurrowDensity Density Density Density Canopy Cover 0.00.20.40.60.81.01.2 Figure 7. Comparisons of canopy cover between burrow densities within each of the pine community types on the Lower Suwannee NWR, Florida. Box contains upper and lower quartiles, dotted line indicates mean, dash indicates median, whiskers connect box to largest and smallest values. Data was not transformed.

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23 High Pine High Pine Pine Flatwoods Pine FlatwoodsHigh Burrow Low Burrow High Burrow Low BurrowDensity Density Density Density Shrub Cover 0.00.20.40.60.81.01.2 Figure 8. Comparisons of shrub cover between burrow densities within each of the pine community types on the Lower Suwannee NWR, Florida. Box contains upper and lower quartiles, dotted line indicates mean, dash indicates median, whiskers connect box to largest and smallest values. Data was not transformed.

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24 High Pine High Pine Pine Flatwoods Pine FlatwoodsHigh Burrow Low Burrow High Burrow Low BurrowDensity Density Density Density Herb Cover 0.00.20.40.60.81.01.2 Figure 9. Comparisons of herbaceous ground cover between burrow densities within each of the pine community types on the Lower Suwannee NWR, Florida. Box contains upper and lower quartiles, dotted line indicates mean, dash indicates median, whiskers connect box to largest and smallest values. Data was not transformed.

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25 Woody ground cover differed between high pine tortoise burrow density category sites ( P =0.000). Woody ground cover also differed between pine flatwoods high and low tortoise burrow density sites (0 P =0.000). Within both habitat categories, low tortoise burrow density sites had greater woody ground cover densities than respective high burrow density sites (Figure 9). Amount of exposed bare mineral soil differed between high pine tortoise burrow density category sites ( P =0.000), as illustrated in Figure 10. However, amount of exposed bare mineral soil did not differ between pine flatwoods high and low tortoise burrow density sites ( P =0.320). Within high pine habitat, sites with high tortoise burrow densities had more exposed bare mineral soil than respective low burrow density sites. Within pine flatwoods sites, areas of bare mineral soil were infrequent, thus accounting for the lack of difference between sites. The amount of detritus ground cover differed between high pine tortoise burrow density category sites ( P =0.000). The amount of detritus ground cover also differed between pine flatwoods high and low tortoise burrow density sites ( P =0.004). Within both habitat categories low tortoise burrow density sites had more detritus ground cover than respective high burrow density sites (Figure 11). Chi-Square Analysis Chi-square tests of independence results for the association of canopy cover to shrub cover within both habitat categories are shown in table 3. Shrub cover was dependent upon canopy cover in both high pine and pine flatwoods categories respectively (=40.7, 24.7).

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26 High Pine High Pine Pine Flatwoods Pine FlatwoodsHigh Burrow Low Burrow High Burrow Low BurrowDensity Density Density Density Woody Cover 0.00.20.40.60.81.01.2 Figure 10. Comparisons of woody ground cover between burrow densities within each of the pine community types on the Lower Suwannee NWR, Florida. Box contains upper and lower quartiles, dotted line indicates mean, dash indicates median, whiskers connect box to largest and smallest values. Data was not transformed.

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27 High Pine High Pine Pine Flatwoods Pine FlatwoodsHigh Burrow Low Burrow High Burrow Low BurrowDensity Density Density Density Bare Soil Cover 0.00.20.40.60.81.0 Figure 11. Comparisons of bare soil ground cover between burrow densities within each of the pine community types on the Lower Suwannee NWR, Florida. Box contains upper and lower quartiles, dotted line indicates mean, dash indicates median, whiskers connect box to largest and smallest values. Data was not transformed.

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28 High Pine High Pine Pine Flatwoods Pine FlatwoodsHigh Burrow Low Burrow High Burrow Low BurrowDensity Density Density Density Detritus Cover 0.00.20.40.60.81.0 Figure 12. Comparisons of detritus ground cover between burrow densities within each of the pine community types on the Lower Suwannee NWR, Florida. Box contains upper and lower quartiles, dotted line indicates mean, dash indicates median, whiskers connect box to largest and smallest values. Data was not transformed.

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29 Chi-square tests of independence results for the association of canopy cover to herbaceous ground cover, within both habitat categories are shown in table 4. Notice within the pine flatwoods habitat category canopy cover and herbaceous ground cover are independent (=2). While herbaceous ground cover was not independent of canopy cover within the high pine category (=8.9). Chi-square tests of independence results for the association of canopy cover to woody ground cover, within both habitat categories are shown in table 5. High chi-square values for both high pine and pine flatwoods categories respectively (= 17.4, 17.8) indicate the amount of woody ground cover was not independent of the accompanying canopy cover. Chi-square tests of independence results for the association of canopy cover to bare ground cover, within both habitat categories are shown in table 6. The chi-square values suggest bare ground cover is not independent within the high pine habitats (= 5.6), however within the pine flatwoods category bare ground cover is independent of canopy cover (= 0.01). Chi-square tests of independence results for the association of canopy cover to detritus ground cover are shown in table 7. Within both habitat categories detritus ground cover was dependent upon the amount of canopy cover, with a high pine value of 7 and a pine flatwoods of 4.9. Chi-square tests of independence results for the association of shrub cover to herbaceous ground cover, within both habitat categories are shown in table 8. Within both habitat categories herbaceous ground cover was not independent of shrub cover (high pine = 37.9, pine flatwoods = 60.3). Chi-square tests of independence results for the association of shrub cover to woody ground cover, within both habitat categories are shown in table 9. Indicating the amount of woody ground cover was not independent

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30 Table 3. Chi-square tests of independence between canopy cover and shrub cover within the 2 habitat categories on the Lower Suwannee NWR, Florida. High Pine Vegetation > 50% Shrub <50% Shrub Cover Observed (expected) Observed (expected) Totals > 50% Canopy 39(20.7) 6(25.8) 45 <50% Canopy 50(70.2) 105(87.4) 155 Totals 89 111 200 = 40.7 df= (2-1)(2-1)= 1 P (=.05) = 3.84 Pine Flatwoods Vegetation > 50% Shrub <50% Shrub Cover Observed (expected) Observed (expected) Totals > 50% Canopy 58(41.6) 21(39.2) 79 <50% Canopy 45(63.4) 76(59.8) 121 Totals 103 97 200 = 24.7 df= (2-1)(2-1)= 1 P (=.05) = 3.84

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31 Table 4. Chi-square tests of independence between canopy cover and herbaceous ground cover within the 2 habitat categories on the Lower Suwannee NWR, Florida. High Pine Vegetation > 50% Herb <50% Herb Cover Observed (expected) Observed (expected) Totals > 50% Canopy 3(11) 42(35.4) 45 <50% Canopy 44(37.4) 111(120.1) 155 Totals 47 153 200 = 8.9 df= (2-1)(2-1)= 1 P (=.05) = 3.84 Pine Flatwoods Vegetation > 50% Herb <50% Herb Cover Observed (expected) Observed (expected) Totals > 50% Canopy 21(26.4) 58(54.4) 79 <50% Canopy 44(40.3) 77(83) 121 Totals 65 135 200 = 2 df= (2-1)(2-1)= 1 P (=.05) = 3.84

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32 Table 5. Chi-square tests of independence between canopy cover and woody ground cover within the 2 habitat categories on the Lower Suwannee NWR, Florida. High Pine Vegetation > 50% Woody <50% Woody Cover Observed (expected) Observed (expected) Totals > 50% Canopy 22(11.5) 23(34.5) 45 <50% Canopy 28(39) 127(117) 155 Totals 50 150 200 = 17.4 df= (2-1)(2-1)= 1 P (=.05) = 3.84 Pine Flatwoods Vegetation > 50% Woody <50% Woody Cover Observed (expected) Observed (expected) Totals > 50% Canopy 39(25.6) 40(54.4) 79 <50% Canopy 25(39) 96(83) 121 Totals 64 136 200 = 17.8 df= (2-1)(2-1)= 1 P (=.05) = 3.84

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33 Table 6. Chi-square tests of independence between canopy cover and bare ground cover within the 2 habitat categories on the Lower Suwannee NWR, Florida. High Pine Vegetation > 50% Bare <50% Bare Cover Observed (expected) Observed (expected) Totals > 50% Canopy 0(4.1) 45(41.9) 45 <50% Canopy 18(14) 137(142) 155 Totals 18 182 200 = 5.6 df= (2-1)(2-1)= 1 P (=.05) = 3.84 Pine Flatwoods Vegetation > 50% Bare <50% Bare Cover Observed (expected) Observed (expected) Totals > 50% Canopy 0(0) 79(80) 79 <50% Canopy 0(0) 121(122) 121 Totals 0 200 200 = 0.01 df= (2-1)(2-1)= 1 P (=.05) = 3.84

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34 Table 7. Chi-square tests of independence between canopy cover and detritus ground cover within the 2 habitat categories on the Lower Suwannee NWR, Florida. High Pine Vegetation > 50% Detritus <50% Detritus Cover Observed (expected) Observed (expected) Totals > 50% Canopy 17(10.6) 28(35.4) 45 <50% Canopy 29(35.9) 126(120) 155 Totals 46 154 200 = 7 df= (2-1)(2-1)= 1 P (=.05) = 3.84 Pine Flatwoods Vegetation > 50% Detritus <50% Detritus Cover Observed (expected) Observed (expected) Totals > 50% Canopy 11(17.6) 68(62.4) 79 <50% Canopy 33(26.8) 88(95.2) 121 Totals 44 156 200 = 4.9 df= (2-1)(2-1)= 1 P (=.05) = 3.84

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35 Table 8. Chi-square tests of independence between shrub cover and herbaceous ground cover within the 2 habitat categories on the Lower Suwannee NWR, Florida. High Pine Vegetation > 50% Herb <50% Herb Cover Observed (expected) Observed (expected) Totals > 50% Shrub 2(23.4) 87(66.6) 89 <50% Shrub 45(29.1) 66(82.9) 111 Totals 47 153 200 = 37.9 df= (2-1)(2-1)= 1 P (=.05) = 3.84 Pine Flatwoods Vegetation > 50% Herb <50% Herb Cover Observed (expected) Observed (expected) Totals > 50% Shrub 5(30.7) 98(73.3) 103 <50% Shrub 54(29.4) 43(69.1) 97 Totals 59 141 200 = 60.3 df= (2-1)(2-1)= 1 P (=.05) = 3.84

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36 the amount of shrub cover. With high chi-square values reached within both habitat categories (high pine = 60.3, pine flatwoods = 50.5). Chi-square tests of independence results for the association of shrub cover to bare ground cover, within both habitat categories are shown in table 10. Within the high pine habitats bare ground cover was not independent of shrub cover (= 15.9), while the pine flatwoods category was unable to show independence (= 0), most probably due to a lack of bare ground within this habitat. Chi-square tests of independence results for the association of shrub cover to detritus ground cover, within both habitat categories are shown in table 11. Within both high pine and pine flatwoods categories, respectively the amount of detritus ground cover was not independent from the amount of accompanying shrub cover (= 16.4 and 4.6).

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37 Table 9. Chi-square tests of independence between shrub cover and woody ground cover within the 2 habitat categories on the Lower Suwannee NWR, Florida. High Pine Vegetation > 50% Woody <50% Woody Cover Observed (expected) Observed (expected) Totals > 50% Shrub 46(22.5) 43(67.5) 89 <50% Shrub 4(28) 107(84) 111 Totals 50 150 200 = 60.3 df= (2-1)(2-1)= 1 P (=.05) = 3.84 Pine Flatwoods Vegetation > 50% Woody <50% Woody Cover Observed (expected) Observed (expected) Totals > 50% Shrub 56(33.3) 47(71.8) 103 <50% Shrub 7(31.4) 90(67.6) 97 Totals 63 137 200 = 50.5 df= (2-1)(2-1)= 1 P (=.05) = 3.84

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38 Table 10. Chi-square tests of independence between shrub cover and bare ground cover within the 2 habitat categories on the Lower Suwannee NWR, Florida. High Pine Vegetation > 50% Bare <50% Bare Cover Observed (expected) Observed (expected) Totals > 50% Shrub 0(8.1) 89(81.9) 89 <50% Shrub 18(10) 93(101.9) 111 Totals 18 182 200 = 15.9 df= (2-1)(2-1)= 1 P (=.05) = 3.84 Pine Flatwoods Vegetation > 50% Bare <50% Bare Cover Observed (expected) Observed (expected) Totals > 50% Shrub 0(0) 103(103) 103 <50% Shrub 0(0) 97(97) 97 Totals 0 200 200 = 0 df= (2-1)(2-1)= 1 P (=.05) = 3.84

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39 Table 11. Chi-square tests of independence between shrub cover and detritus ground cover within the 2 habitat categories on the Lower Suwannee NWR, Florida. High Pine Vegetation > 50% Detritus <50% Detritus Cover Observed (expected) Observed (expected) Totals > 50% Shrub 32(20.7) 57(70.2) 89 <50% Shrub 13(25.8) 98(87.4) 111 Totals 45 155 200 = 16.4 df= (2-1)(2-1)= 1 P (=.05) = 3.84 Pine Flatwoods Vegetation > 50% Detritus <50% Detritus Cover Observed (expected) Observed (expected) Totals > 50% Shrub 29(22.9) 74(81.1) 103 <50% Shrub 15(21.6) 82(76.4) 97 Totals 0 200 200 = 4.6 df= (2-1)(2-1)= 1 P (=.05) = 3.84

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DISCUSSION Vegetation cover survey results showed significant differences in all vegetation categories except the bare soil ground cover form, since bare soil was uncommon within the pine flatwoods compartments. These results supported the hypothesis that compartments with high herbaceous ground cover and low shrub/canopy covers supported higher densities of gopher tortoise burrows. While areas with low burrow densities had higher shrub and canopy covers with lower herbaceous ground cover. Chi-square results allowed the rejection of the null hypothesis that ground cover densities were independent of canopy and shrub cover densities in almost all categories. Though within the flatwoods habitats herbaceous and bare ground covers were not dependent upon canopy cover. Also, within the flatwoods habitats bare ground cover was not dependent upon the amount of shrub cover. The following discussion describes these findings in the context of the respective habitat categories. High Pine Habitats In high pine habitats on the Lower Suwannee NWR, tortoise burrow densities were higher within survey compartments with the most open shrub layers. A lack of burrows within a survey compartment was associated with both shrub and canopy closure. The shrub and canopy layers within low tortoise burrow density compartments were characterized by a dense covering of young oaks. These successional changes within the shrub and canopy layers appear to have led to lower herbaceous ground cover, less areas of exposed bare soil with higher woody and detritus ground covers observed. Within 40

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41 high pine habitat compartments paired comparisons indicate the strongest association between tortoise burrow density and vegetation cover was at the shrub layer, this is likely because of the rather young age of the planted pine component (11 years) where light limitation to understory vegetation due to canopy closure has not yet occurred. The vegetation cover differences, and therefore tortoise burrow density differences, between survey compartments cannot be fully explained by past management practices. The U. S. Fish and Wildlife Service assumed management of the lands containing the high pine survey compartments in 1990, from the Georgia Pacific timber company. Soon after refuge acquisition the area was completely clear-cut of planted slash pines by Georgia Pacific in accordance with a deed agreement. After clear cutting, refuge personnel began mechanical, V-blade planting of the sites to long leaf pine in 1993. Intensive site prepping was not conducted prior to the 1993 plantings. A detailed management history, including methods of site prep, vegetation control and planting techniques on these sites prior to 1990 is unknown. Though some site prep techniques could entomb tortoises, such as the piling of large windrows directly on a burrow, gopher tortoises are able to dig out of impacted burrows following certain types of site preparation on sandy soils (Landers and Buckner 1981, Diemer and Moler 1982). Thus, it is unlikely tortoises entombment can explain differences in tortoise density noted in the 2002 survey. One explanation for the observed vegetation cover differences between survey compartments is available soil moisture. The GIS soil analysis revealed that 29% of the two low tortoise burrow density compartments occurred in a Leon soil series. This soil series has a higher moisture content than other soils series present in this survey

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42 compartment. However this particular Spodosol soil is very deep and has high permeability in the A and E horizon with moderate permeability in the Bh horizon. Also, plant community composition observed did not reflect an extreme difference in available moisture compared to other high pine sites, as more mesic or flatwoods type vegetation such as gallberry (Ilex glabra) or ericaceous shrubs were not observed to dominate any of the high pine sites. A working hypothesis to be tested in the future is that higher soil moisture contents may have increased the rate of shrub and canopy closure within these sites, but that soil moisture differences were not dramatic enough to shift community composition towards flatwoods type plants. Pine Flatwoods Habitats Within pine flatwoods compartments surveyed, differences in vegetation cover and gopher tortoise burrow densities were most strongly associated with the amount of shrub cover. Observed shrub cover was very low in compartments characterized as high tortoise burrow density, while the opposite was observed in low tortoise burrow density compartments. Since all compartments contained planted slash pines of very similar ages and basal areas, canopy differences observed were related to canopy closure by plants within the shrub layer. This successional closure of the shrub and canopy layers presumably caused lower herbaceous ground cover along with higher woody and detritus ground covers observed within the low tortoise density compartments. The shrub cover observed in low tortoise burrow density compartments is characterized by dense stands of gallberry and saw-palmetto with intermixed oaks. Shifting to growing season burns may reduce the density of saw-palmetto and gallberry stands. It has been theorized that top kill of many species early in the growing season can halt carbohydrate production when carbohydrate reserves normally in the root system are

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43 at their lowest level, thereby increasing kill (Waldrop et al. 1987). However, other studies suggest that even after repeated, early growing season burns these very fire resilient species will not be eradicated (Hough 1968, Hughes and Knox 1964). Though growing season burns should decrease their densities due to the stress placed on carbohydrate reserves. Carbohydrate reserves in the rhizomes of gallberry and saw-palmetto have been found at their lowest in August (Hough 1968, Hughes and Knox 1964). Therefore, if the management goal is to decrease the densities of these two species late growing season burns may be most effective. As with the high pine survey compartments the observed differences in vegetation cover can not be fully explained by past management practices; most probably, slight differences in soil moisture may have accelerated succession within certain compartments.

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MANAGEMENT IMPLICATIONS Gopher tortoise burrow densities were highest in high pine and pine flatwoods communities on the Lower Suwannee NWR when shrub and canopy cover was relatively low and herbaceous ground cover, as well as areas of bare soil, were relatively high. Gopher tortoise densities are higher in open areas with herbaceous ground cover than in brushy, shaded sites; the former have patches of bare ground needed for nest excavation and also provide abundant herbaceous vegetation for feeding (Cox et al. 1987). This type of habitat can be promoted by growing season fires (Robbins and Meyers 1992). Furthermore, it has been suggested that growing season fires might increase the amount of food available in late summer when food quality is declining and would provide food conditions for new hatchlings, which emerge in late summer and early fall (Cox et al. 1987). Under current conditions burning is problematic, if not impossible within high pine areas on the refuge due to years of habitat degradation from a lack of fire. Fire suppression has allowed these areas to begin succession to a mesic hardwood forest. This process of succession has concentrated gopher tortoise populations to the higher, drier sites, where inevitable succession is lagging behind more moist sites. Mesic hardwood forest conditions are characterized by higher shading, greater detritus accumulation, and less herbaceous ground cover than natural high pine forests. Since these areas have been excluded from fire for at least 15 years, a lack of ground fuel greatly restricts a fires 44

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45 intensity and its ability to spread thereby rendering fire alone somewhat ineffective as a management tool for restoration of these habitats. Two alternate methods for restoring high pine habitats on the refuge include use of a selective herbicide to remove hardwoods and restoration plantings. The utility of a selective herbicide such as hexazinone, has been demonstrated for restoration of longleaf pine communities (Hay-Smith and Tanner 1994). Hexazinone can facilitate the release of wiregrass and aid in the reduction of scrub oak populations without damage to other woody and herbaceous vegetation. Current research calls for forest managers to use hexazinone rates ranging from 0.84 to 1.68Kg/ha, in liquid form applied in a grid pattern to the soil surface. Applications should be made prior to rainfall, as rain is required to distribute the herbicide through the root column. Broadcast applications of granular hexazinone are discouraged as this may damage the herbaceous plant layer. It should also be noted, the use of an herbicide is an important precursor to facilitate future prescribed fire, not to replace fire. After herbicide use restoration plantings of wiregrass should be pursued as aggressively as possible. Restoration plantings of wiregrass will reintroduce this fire-dependant species to these sites and further provide an ignition source to carry future prescribed fires. These plantings will also provide forage for resident gopher tortoises and other fauna. Depending upon the amount of site preparation needed on individual sites and costs, refuge management can choose between sowing wiregrass seeds or planting plugs. Wiregrass seeds do best when sown upon bare soil, while plugs, despite their higher costs are better suited for a wider array of planting sites.

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46 Following herbicide application and the establishment of wiregrass, prescribed fire should be returned to these sites. Initially emphasis should be placed on growing season burns to facilitate the further propagation of wiregrass. Subsequently, burning at various times of the year under varying conditions will encourage plant diversity without completely eliminating individual species. A fire frequency of 2 to 4 years would mimic the natural burning frequency, maintain pine dominance and promote reproduction in wiregrasses (Tanner et al. 1991). Pine flatwoods habitats on the refuge exist as slash pine plantations established for pulpwood production, therefore, these habitats differ markedly from natural flatwoods habitats. To bolster pulpwood production, timber companies established dense plantations of slash pine across landscapes. Currently slash pine plantations on the refuge are planted too densely and should be thinned. Timber thinning opens the canopy, which promotes herbaceous growth on the forest floor and aids in prescribed burning by allowing the upward release of heat, the latter especially important for the safe implementation of growing season burns. As with the high pine habitats, a return to growing season burns and a varied burn cycle with burns occurring every 2 to 4 years should be a management goal. Compartments surveyed for this study contained slash pine basal areas of approximately 21.75 sq. meters/hectare (94 sq. feet/acre) (pers. comm., D. Barrand, Chiefland, FL). The refuges habitat management plan appropriately calls for thinning of these stands down to 11.5-15 sq. meters/hectare (50-65 sq. feet/acre) during second entry thinnings (unpubl. Data, USFWS, LSNWR). This study highlights the importance of timber thinning and the need to restore diversity to areas indiscriminately planted to slash

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47 pine monocultures. Upon thinning, restoration plantings of longleaf pine and wiregrass should take place. Within the habitat management plan the refuge is divided into 9 compartments to facilitate habitat management actions and aid record keeping. Management compartment boundaries were established along major physiographical and legal features such as roads, streams and landowner boundaries. Some of these compartments are further compartmentalized into sub-compartments for more precise land management work. The current system of compartmentalization should be further delineated since the current compartments ignore or leave out habitats that were converted to slash pine stands by the previous timber company management. For example, areas within the compartments I sampled contained ridges of sandy, Entisol soils which would have supported either high pine or scrub habitats prior to timber management. Also, within these compartments, areas of wetter, Aquent soils were bedded and planted through. Spodosols would be the primary soils dominated by pine flatwoods, as they form under coniferous trees whose needles are low in base-forming cations and high in acid resins. These acids bind with iron and aluminum and are carried downward until they precipitate forming the characteristic spodic horizon (Brady and Weil 2002). A closer look at each management unit, noting these areas of soil diversity could benefit future habitat management activities on the refuge. For example, depending on area, higher ridges could be clear cut of planted slash pine, then restored to high pine or scrub habitats, while areas of spodosol soils could be maintained as pine flatwoods. Instead of looking at one block of slash pine timber, this shift in management approach would focus

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48 on restoring the natural interspersion of habitats that existed on the area prior to timber company acquisition, thereby greatly increasing habitat diversity on the refuge.

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APPENDIX ANALYSIS OF VARIANCE TABLES

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SAD Analysis of Variance Tables Table 12. Analysis of variance results for high pine canopy cover comparisons. Anova: Single Factor SUMMARY Groups Count Sum Average Variance High Burrow Density 100 29.53 0.295 0.047 Low Burrow Density 100 70.42 0.704 0.088 ANOVA Source of Variation SS df MS F P-value F crit Between Groups 8.359 1 8.359 122.700 1.675E-22 3.888 Within Groups 13.490 198 0.068 Total 21.850 199 50

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51 Table 13. Analysis of variance results for high pine shrub cover comparisons. Anova: Single Factor SUMMARY Groups Count Sum Average Variance High Burrow Density 100 26.01 0.260 0.042 Low Burrow Density 100 106.15 1.061 0.068 ANOVA Source of Variation SS df MS F P-value F crit Between Groups 32.112 1 32.112 575.799 1.629E-60 3.888 Within Groups 11.042 198 0.055 Total 43.154 199

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52 Table 14. Analysis of variance results for high pine woody ground cover comparisons. Anova: Single Factor SUMMARY Groups Count Sum Average Variance High Burrow Density 100 29.01 0.290 0.057 Low Burrow Density 100 70.77 0.707 0.057 ANOVA Source of Variation SS df MS F P-value F crit Between Groups 8.719 1 8.719 152.52 2.370E-26 3.888 Within Groups 11.319 198 0.057 Total 20.039 199

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53 Table 15. Analysis of variance results for high pine bare mineral soil cover comparisons. Anova: Single Factor SUMMARY Groups Count Sum Average Variance High Burrow Density 100 49.32 0.493 0.070 Low Burrow Density 100 18.2 0.182 0.043 ANOVA Source of Variation SS df MS F P-value F crit Between Groups 4.842 1 4.842 84.635 5.109E-17 3.888 Within Groups 11.328 198 0.057 Total 16.170 199

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54 Table 16. Analysis of variance results for high pine detritus ground cover comparisons. Anova: Single Factor SUMMARY Groups Count Sum Average Variance High Burrow Density 100 33.52 0.335 0.095 Low Burrow Density 100 53.3 0.533 0.107 ANOVA Source of Variation SS df MS F P-value F crit Between Groups 1.956 1 1.956 19.259 1.854E-05 3.888 Within Groups 20.111 198 0.101 Total 22.067 199

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55 Table 17. Analysis of variance results for high pine herbaceous ground cover comparisons. Anova: Single Factor SUMMARY Groups Count Sum Average Variance High Burrow Density 100 71.27 0.712 0.080 Low Burrow Density 100 31.31 0.313 0.056 ANOVA Source of Variation SS df MS F P-value F crit Between Groups 7.984 1 7.984 116.741 1.089E-21 3.888 Within Groups 13.541 198 0.068 Total 21.525 199

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56 Table 18. Analysis of variance results for pine flatwoods canopy cover comparisons. Anova: Single Factor SUMMARY Groups Count Sum Average Variance High Burrow Density 100 49.39 0.493 0.135 Low Burrow Density 100 79.48 0.794 0.133 ANOVA Source of Variation SS df MS F P-value F crit Between Groups 4.527 1 4.527 33.646 2.583E-08 3.888 Within Groups 26.640 198 0.134 Total 31.167 199

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57 Table 19. Analysis of variance results for pine flatwoods shrub cover comparisons. Anova: Single Factor SUMMARY Groups Count Sum Average Variance High Burrow Density 100 26.36 0.263 0.061 Low Burrow Density 100 113.48 1.134 0.033 ANOVA Source of Variation SS df MS F P-value F crit Between Groups 37.949 1 37.949 801.44 1.564E-71 3.888 Within Groups 9.375 198 0.047 Total 47.325 199

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58 Table 20. Analysis of variance results for pine flatwoods woody ground cover comparisons. Anova: Single Factor SUMMARY Groups Count Sum Average Variance High Burrow Density 100 45.52 0.455 0.068 Low Burrow Density 100 80.15 0.801 0.062 ANOVA Source of Variation SS df MS F P-value F crit Between Groups 5.996 1 5.996 91.798 4.173E-18 3.888 Within Groups 12.933 98 0.065 Total 18.929 199

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59 Table 21. Analysis of variance results for pine flatwoods bare mineral soil cover comparisons. Anova: Single Factor SUMMARY Groups Count Sum Average Variance High Burrow Density 100 3.29 0.032 0.000841 Low Burrow Density 100 3 0.03 1.822E-18 ANOVA Source of Variation SS df MS F P-value F crit Between Groups 0.00042 1 0.000420 1 0.318 3.888 Within Groups 0.083 198 0.0004205 Total 0.083 199

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60 Table 22. Analysis of variance results for pine flatwoods detritus ground cover comparisons. Anova: Single Factor SUMMARY Groups Count Sum Average Variance High Burrow Density 100 37.86 0.378 0.095 Low Burrow Density 100 53.23 0.532 0.084 ANOVA Source of Variation SS df MS F P-value F crit Between Groups 1.181 1 1.181 13.147 0.00036624 3.888 Within Groups 17.789 198 0.089 Total 18.970 199

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61 Table 23. Analysis of variance results for pine flatwoods herbaceous ground cover comparisons. Anova: Single Factor SUMMARY Groups Count Sum Average Variance High Burrow Density 100 81.06 0.810 0.165 Low Burrow Density 100 40.45 0.404 0.053 ANOVA Source of Variation SS df MS F P-value F crit Between Groups 8.245 1 8.245 75.187 1.541E-15 3.888 Within Groups 21.714 198 0.109 Total 29.960 199

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LITERATURE CITED Abrahamson, W. G., and D. C. Harnett. 1990. Pine flatwoods and dry prairies. Pages 103-149 in R. L. Myers and J. J. Ewel, eds. Ecosystems of Florida. Univ. of Central Florida Press, Orlando. 765 pp. Ash, A. N. 1988. Disappearance of salamanders from clearcut plots. The Journal of the Elisha Mitchell Scientific Society 104:116-122. Auffenberg, W. 1969. Tortoise behavior and survival. Rand McNally, Chicago, IL. 179 pp. Auffenberg, W. and R. Franz. 1982. The status and distribution of the gopher tortoise (Gopherus polyphemus). Pages 95-126 in North American Tortoises: conservation and ecology. Wildlife Research Report 12. U.S. Fish and Wildlife Service, Washington, D.C. Berish, J. E. 2001. Management considerations for the gopher tortoise in Florida. Final Report. Florida Fish and Wildlife Conservation Commission, Tallahassee, FL. Blymer, M. J., and B. S. McGinnes. 1977. Observations on possible detrimental effects of clearcutting on terrestrial amphibians. Bulletin of the Maryland Herpetological Society 13:167-178. Brady, N. C., and R. R. Weil. 2002. The nature and properties of soils. 13 th Ed. Pearson Education, Inc. Delhi, India. Breininger, D. R., P. A. Schmalzer, D. A. Rydene, and C. R. Hinkle. 1988. Burrow and habitat relationships of the gopher tortoise in coastal shrub and slash pine flatwoods on Merritt Island, Florida. Final Rep. Proj. No. GFC-84-016. Fla. Game and Fresh Water Fish Comm. Tallahassee, FL. 24pp. Brode, J. M., and R. B. Bury. 1984. The importance of riparian systems to amphibians and reptiles. Pages 30-36 in R. E. Warner and K. E. Hendrix, eds. Proceedings of the conference on California riparian systems, Univ. of California, Davis. Buhlmann, K. A., C. A. Pague, J.C. Mitchell, and R. B. Glasgow. 1988. Forestry operations and terrestrial salamanders: techniques in a study of the Cow Knob salamander, Plethodon punctatus. Pages 38-44 in R. C. Szaro, K. E. Severson and D. R. Patton eds. Management of amphibians, reptiles, and mammals in North America. Technical Report RM-166. U.S. Forest Service, Rocky Mountain Forest and Range Experiment Station, Fort Collins, CO. 62

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63 Bury, R. B. 1983. Differences in amphibian populations in logged old growth redwood forests. Northwest Science 57:167-178. Campbell, H. W., and S. P. Christman. 1982. The herpetological components of Florida sandhill and sand pine scrub associations. Pages 163-171 in N. J. Schott, Jr., ed. Herpetological communities. U.S. Fish and Wildl. Serv. Wild. Res. Rep. 13. Corn, P. S., and R. B. Bury. 1989. Logging in western Oregon: Responses of headwater habitats and stream amphibians. Forest Ecology and Management 29:39-57. Cox, J. 1987. The breeding bird survey in Florida: 1969-1983. Fla. Field Nat. 15, 29-56. Cox, J., D. Inkley, and R. Kautz. 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. Florida Game and Fresh Water Fish Commission. Tallahassee, Fl. 69pp. Diemer, J. E., and P. E. Moler. 1982. Gopher tortoise response to site preparation in northern Florida. Proceedings of the Annual Conference of the Southeastern Assoc. of Fish and Wildl. Agencies 36:634-637. Dodd, C. K. Jr. 1991. The status of the Red Hills salamander, Phaeognathus bubrichtt, Alabama, U.S.A., 1976-1988. Biological Conservation 55:57-75. _____. 1993. Distribution of striped newts (Notophthahnus perstriatus) in Georgia. U. S. Fish and Wildlife Service. Jacksonville, FL. Edmisten, J. E. 1963. The ecology of the Florida pine flatwoods. Ph.D. Dissertation, Univ. of Florida, Gainesville. Franz, R., and W. Auffenberg. 1974. The gopher tortoise in Georgia. Herpetological Rev. 5:74-75. Garner, J. A., and J. L. Landers. 1979. Foods and habitat of the gopher tortoise in south-western Georgia. Poc. Ann. Conf. South-eastern Assoc. Fish and Wildl. Agencies. 35:120-134. Givens, K. T., J. N. Layne, W. G. Abrahamson, and S. C. White-Schuler. 1984. Structural changes and successional relationships of five Florida Lake Wales Ridge plant communities. Bull. Torrey Bot. Club 63, 8-18. Hay-Smith, L., and G. W. Tanner. 1999. Restoring longleaf pine sandhill communities with an herbicide. Florida Cooperative Extension Service, Unv. of Florida. WEC-131. 4pp.

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64 Herrington, R. E., and J. H. Larsen. 1985. Current status, habitat requirements and management of the Larch mountain salamander Plethodon larselli (Burns). Biological Conservation 34:169-179. Higgins, K. F., J. L. Oldemeyer, K. J. Jenkins, G. K. Clambey, and R. F. Harlow. 1994. Vegetation sampling and measurement in T. A. Bookhout ed. Research and management techniques for wildlife and habitats. 5 th edition. The Wildlife Society, Bethesda, Md. 740 pp. Hough, W. A. 1968. Carbohydrate reserves of saw-palmetto: seasonal variation and effects of burning. Forest Science 14:399-405. Hughes, R. H., and F. E. Knox. 1964. Response of gallberry to seasonal burning. U.S. Department of Agriculture, Forest Service, Southeastern Forest Experiment Station, Research Note SE-21. Jackson, D. R., and E. R. Milstrey. 1988. The fauna of gopher tortoise burrows. Proc. Gopher Tortoise Relocation Symp. J. E. Diemer, D. R. Jackson, J. L. Landers, J. N. Layne, and D. A. Woods, eds. Nongame Wildlife Program Technical Report. Florida Game and Fresh Water Fish Commission, Tallahassee, FL. Jordan, R. Jr., and R. H. Mount. 1975. The status of the Red Hills Salamander, Phaeognathus bubrichtt, Highton. Journal of Herpetology 9:211-215. Laessle, A. M. 1942. The plant communities of the Welaka area with special reference to correlation between soils and vegetational succession. Biol. Sci. Ser. 4. Univ. of Florida Pub., Gainesville. Landers, J. L., and D. W. Speake. 1980. Management needs of sandhill reptiles in southern Georgia. Ann. Conf. Southeast. Assoc. Fish and Wildl. Agencies 34:515-529. _____. and J. L. Buckner. 1981. The gopher tortoise: effects of forest management and critical aspects of its ecology. Technical Note No.56. Southlands Experimental Forest, Bainbridge, Georgia. Liudahl, K., R. L. Weatherspoon, and E. L. Readle. 2003. Soil survey of Dixie County, Florida. United States Department of Agriculture, Natural Resources Conservation Service. Maliakal, S. T., E. S. Menges, and J. S. Denslow. 2000. Community composition and regeneration of Lake Wales ridge wiregrass flatwoods in relation to time-since-fire. Journal of the Torr. Bot. Soc., Vol. 127, No. 2. pp. 125-138. McCoy E. D., and H. R. Mushinsky. 1995. The demography of Gopherus polyphemus (Daudin) in relation to size of available habitat. Nongame Wildl. Program Project Report. 77 pp. Florida Game and Fresh Water Fish Commission, Tallahassee, FL.

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65 Means, D. B., J. G. Palis, and M. Baggett. 1996. Effects of slash pine silviculture on a Florida population of flatwoods salamander. Conservation Biology Volume 10, No. 2: 426-437. Monk, C. D. 1968. Successional and environmental relationships of the forest vegetation of north-central Florida. Am. Midl. Nat. 79, 441-457. Myers, R. L. 1985. Fire and the dynamic relationship between Florida sandhill and sand pine scrub vegetation. Bull. Torrey Bot. Club 112, 241-252. Myers, R. L. and D. L. White. 1987. Landscape history and changes in sandhill vegetation in north-central and south-central Florida. Bull. Torrey bot. Club 114, 21-32. _____. 1990. Scrub and high pine. Pages 150-193 in R. L. Myers and J. J. Ewel, eds. Ecosystems of Florida. Univ. of Central Florida Press, Orlando. 765 pp. Noel, J. M., W. J. Platt and E. B. Moser. 1998. Structural characteristics of oldand second-growth stands of longleaf pine (Pinus palustris) in the gulf coastal region of the U.S.A. Conservation Biology. Vol. 12. Num. 3, 533-548. Ober, L. D. 1954. Plant communities of the flatwood forests in Austin Cary memorial forest. M.S. Thesis, Univ. of Florida, Gainesville. OMeara, T. E., and M. J. Abbott. 1987. Gopher tortoise response to summer burning in longleaf pine/turkey oak sandhills. Annual performance report. Nongame Wildlife Section. Florida Game and Fresh Water Fish Commission. 8pp. Ott, R. L. 1993. An introduction to statistical methods and data analysis. 4 th edition. Duxbury Press. 1051 pp. Belmont, California. Pentranka, J. W., M. E. Eldridge and K. E. Haley. 1993. Effects of timber harvesting on southern Appalachian salamanders. Conservation Biology 7:363-370. Peroni, P. A., and W. G. Abrahamson. 1986. Succession in Florida sandridge vegetation: a retrospective study. Fla. Sci. 49, 176-191. Platt, W. J., G. W. Evans, and S. L. Rathbun. 1988. The populations dynamics of a long-lived conifer (Pinus palustris). Am. Nat. 131, 491-525. Repenning, R. W., and R. F. Labisky. 1985. Effects of even-age timber management on bird communities of the longleaf pine forest in northern Florida. J. Wildl. Manage. 49, 1088-1098. Robbins, L. E., and R. L. Myers. 1992. Seasonal effects of prescribed burning in Florida: A review. Tall Timbers Res. Sta. Misc. Publ. No. 8.

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66 Seigel, R. A., R. B. Smith, and N. A. Seigel. 2003. Swine Flu or 1918 pandemic? Upper respiratory tract disease and the sudden mortality of gopher tortoises (Gopherus polyphemus) on a protected habitat in Florida. J. Herpetol. 37:137-144. Slabaugh, J. D., A. O. Jones, W. E. Puckett, and J. N. Schuster. 1996. Soil survey of Levy County, Florida. United States Department of Agriculture, Natural Resources Conservation Service. Sykes, P. W., C. B. Kepler, K. L. Litzenberger, H. R. Sansing, E. T. Lewis, and J. S. Hatfield. 1999. Density and habitat of breeding swallow-tailed kites in the Lower Suwannee ecosystem, Florida. Journal of Field Ornithology, Vol. 70, No. 3 pages 321-336. Tanner, G. W., W. R. Marion, and J. J. Mullahey. 2002. Understanding fire: natures land management tool. Florida Cooperative Extension Service, Unv. of Florida. CIR-1018. 4pp. Van Horne, B. 1983. Density as a misleading indicator of habitat quality. Journal of Wildl. Manag. 47:893-901 Veno, P. A. 1976. Successional relationships of five Florida plant communities. Ecology 57, 498-508. Welsh, H.H. Jr. 1990. Relictual amphibians and old-growth forests. Conservation Biology 4:309-319. Waldrop, T. A., D. H. Van Lear, R. T. Lloyd, and W. R. Harms. 1987. Long-term studies of prescribed burning in loblolly pine forests of the southeastern coastal plain. U.S. Department of Agriculture, Forest Service, Southeastern Forest Experiment Station, General Technical Report SE-45.

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BIOGRAPHICAL SKETCH Stephen E. Barlow was born on 25 July 1969 in Plant City, Florida. Soon after, his family moved to Chiefland, Florida, where he was raised; he graduated from Chiefland High School in May 1987. After serving five years in the U. S. Army and three years in the U.S. Army Reserve he attended Pittsburg State University in Pittsburg, Kansas, from which he received his Bachelor of Science degree in biology in December 1997. He enrolled at the University of Florida in January 1998, majoring in environmental science with a minor in wildlife ecology, and graduated with the Master of Science degree in December 2004. During graduate school he began his professional career as a wildlife technician with the Florida Fish and Wildlife Conservation Commission in January 1999. He was soon promoted to biological scientist and after 3 years of service with the state of Florida, he accepted his current position with the U.S. Fish and Wildlife Service as the wildlife biologist on the Lower Suwannee National Wildlife Refuge. He and Elizabeth Jane Works, of Humboldt, Kansas, were married on 21 December 1992. They have one son, Seth Douglas Barlow, born in Gainesville, Florida, on 21 August 2001. 67


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Permanent Link: http://ufdc.ufl.edu/UFE0008342/00001

Material Information

Title: Vegetative Characteristics of Gopher Tortoise (Gopherus polyphemus) Habitat on the Lower Suwannee National Wildlife Refuge: Implications for Restoration and Management of Pine Communities
Physical Description: x, 67 p.
Language: English
Creator: Barlow, Stephen E. ( Dissertant )
Clark, Mark W. ( Thesis advisor )
Tanner, George W. ( Thesis advisor )
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 2004
Copyright Date: 2004

Subjects

Subjects / Keywords: Soil and Water Science thesis, M.S
Dissertations, Academic -- UF -- Soil and Water Science

Notes

Abstract: Previous land use practices on the Lower Suwannee National Wildlife Refuge altered natural fire conditions resulting in habitat degradation, especially within the high pine areas, allowing succession to begin shifting these habitats to mesic hammocks. In addition, many pine flatwoods sites exist as densely planted slash pine plantations, which have been relegated to an infrequent winter fire regime, leading to a shrub level monoculture of gallberry (Ilex glabra) and saw palmetto (Serenoa repens). Changes in vegetative structure of these prominent pine communities are thought to have lowered the quality of gopher tortoise habitat. This investigation evaluates the relationship between vegetative cover of different community strata and the density of gopher tortoise burrows. Vegetation cover was measured on sites with high and low gopher tortoise burrow densities in high pine and pine flatwoods habitats on the refuge from April to July 2004. The line intercept technique was used to measure canopy cover, shrub cover and ground cover. Survey compartment soil compositions were compared using Geographic Information Systems (GIS) data and the county soil surveys. It was predicted that compartments with high tortoise burrow densities would have a relatively open canopy and shrub layers, with relatively high herbaceous and bare soil ground covers. Analysis of variance on vegetation cover data revealed that density of the canopy, shrub layer, and ground cover types were significantly different between high pine sites with different tortoise burrow densities. While the pine flatwoods sites exhibited significant differences in all categories except bare soil, due to a lack of bare soil within these habitats. Lower canopy cover, shrub cover and higher herbaceous ground cover characterized sites with high tortoise burrow densities. Soil analysis revealed higher soil series diversity within the pine flatwoods sites. This soil series diversity is not represented vegetatively within the current slash pine plantation monoculture. The lack of relationship between soil characteristics and vegetation community composition is most likely due to past timber practices that converted all usable land to pine production with little regard for preserving small pockets of habitat. Close analysis of soil series should guide land managers? efforts in restoration of habitats in order to regain pre-European development habitat diversity. Aggressive techniques to reverse succession on high pine sites are also recommended. Restoration of the herbaceous ground layer and a reintroduction to summer fires could then be employed. Timber thinning and a more varied fire regime emphasizing summer burning would benefit pine flatwoods sites; otherwise continued habitat succession and degradation will significantly inhibit tortoise populations.
Subject: Flatwoods, gopher, habitat, longleaf, pine, restoration, tortoise
General Note: Title from title page of source document.
General Note: Document formatted into pages; contains 77 pages.
General Note: Includes vita.
Thesis: Thesis (M.S.)--University of Florida, 2004.
Bibliography: Includes bibliographical references.
Original Version: Text (Electronic thesis) in PDF format.

Record Information

Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
System ID: UFE0008342:00001

Permanent Link: http://ufdc.ufl.edu/UFE0008342/00001

Material Information

Title: Vegetative Characteristics of Gopher Tortoise (Gopherus polyphemus) Habitat on the Lower Suwannee National Wildlife Refuge: Implications for Restoration and Management of Pine Communities
Physical Description: x, 67 p.
Language: English
Creator: Barlow, Stephen E. ( Dissertant )
Clark, Mark W. ( Thesis advisor )
Tanner, George W. ( Thesis advisor )
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 2004
Copyright Date: 2004

Subjects

Subjects / Keywords: Soil and Water Science thesis, M.S
Dissertations, Academic -- UF -- Soil and Water Science

Notes

Abstract: Previous land use practices on the Lower Suwannee National Wildlife Refuge altered natural fire conditions resulting in habitat degradation, especially within the high pine areas, allowing succession to begin shifting these habitats to mesic hammocks. In addition, many pine flatwoods sites exist as densely planted slash pine plantations, which have been relegated to an infrequent winter fire regime, leading to a shrub level monoculture of gallberry (Ilex glabra) and saw palmetto (Serenoa repens). Changes in vegetative structure of these prominent pine communities are thought to have lowered the quality of gopher tortoise habitat. This investigation evaluates the relationship between vegetative cover of different community strata and the density of gopher tortoise burrows. Vegetation cover was measured on sites with high and low gopher tortoise burrow densities in high pine and pine flatwoods habitats on the refuge from April to July 2004. The line intercept technique was used to measure canopy cover, shrub cover and ground cover. Survey compartment soil compositions were compared using Geographic Information Systems (GIS) data and the county soil surveys. It was predicted that compartments with high tortoise burrow densities would have a relatively open canopy and shrub layers, with relatively high herbaceous and bare soil ground covers. Analysis of variance on vegetation cover data revealed that density of the canopy, shrub layer, and ground cover types were significantly different between high pine sites with different tortoise burrow densities. While the pine flatwoods sites exhibited significant differences in all categories except bare soil, due to a lack of bare soil within these habitats. Lower canopy cover, shrub cover and higher herbaceous ground cover characterized sites with high tortoise burrow densities. Soil analysis revealed higher soil series diversity within the pine flatwoods sites. This soil series diversity is not represented vegetatively within the current slash pine plantation monoculture. The lack of relationship between soil characteristics and vegetation community composition is most likely due to past timber practices that converted all usable land to pine production with little regard for preserving small pockets of habitat. Close analysis of soil series should guide land managers? efforts in restoration of habitats in order to regain pre-European development habitat diversity. Aggressive techniques to reverse succession on high pine sites are also recommended. Restoration of the herbaceous ground layer and a reintroduction to summer fires could then be employed. Timber thinning and a more varied fire regime emphasizing summer burning would benefit pine flatwoods sites; otherwise continued habitat succession and degradation will significantly inhibit tortoise populations.
Subject: Flatwoods, gopher, habitat, longleaf, pine, restoration, tortoise
General Note: Title from title page of source document.
General Note: Document formatted into pages; contains 77 pages.
General Note: Includes vita.
Thesis: Thesis (M.S.)--University of Florida, 2004.
Bibliography: Includes bibliographical references.
Original Version: Text (Electronic thesis) in PDF format.

Record Information

Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
System ID: UFE0008342:00001


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VEGETATIVE CHARACTERISTICS OF GOPHER TORTOISE (Gopherus
polyphemus) HABITAT ON THE LOWER SUWANNEE NATIONAL WILDLIFE
REFUGE: IMPLICATIONS FOR RESTORATION AND MANAGEMENT OF PINE
COMMUNITIES
















By

STEPHEN E. BARLOW


A THESIS PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
MASTER OF SCIENCE

UNIVERSITY OF FLORIDA


2004



























Copyright 2004

by

Stephen E. Barlow

































This document is dedicated to my father the late Douglas Alvord Barlow, Dad, mentor
and friend.















ACKNOWLEDGMENTS

I wish to thank the members of my graduate supervisory committee for their

invaluable input and guidance: Dr. Mark W. Clark, chairman and research assistant

professor, Soil and Water Sciences Department; Dr. George W. Tanner, co-chairman and

professor, Department of Wildlife Ecology and Conservation; and Mr. Kenneth

Litzenberger, Refuge Manager, Lower Suwannee National Wildlife Refuge.

I also would like to thank the following individuals: my wife, Elizabeth, for her

strong support, patience and encouragement; my Mom, Kay, for instilling within me a

love of nature; my in-laws, Joe and Jane Works, for constant support and encouragement;

Russ Singleton for helping with gopher tortoise burrow surveys; Vivian R. Soriero, Linda

Casey and John C. Jones for assisting with vegetation surveys; Joan Berish with the

Florida Fish and Wildlife Conservation Commission for assistance throughout the

project; Dr. Lori Wendland, University of Florida College of Veterinary Medicine, for

testing tortoise blood samples; Daniel Barrand for assistance with GIS analysis;

Kenneth W. McCain for providing fire and management history information, and all

employees of the Lower Suwannee National Wildlife Refuge.















TABLE OF CONTENTS

page

A C K N O W L E D G M E N T S ......... ................................................................................... iv

L IST O F T A B L E S ................................................................... .............. vi

LIST OF FIGURE S ................ ...... ......... ............ ............... viii

ABSTRACT .............. .......................................... ix

INTRODUCTION .............. ................................... ..............

MATERIAL AND METHODS ............................................................. ..................7

Study A rea ....................................................................... . 7
Survey C om part ents .................................................................. ........................ 7
V vegetation Surveys .................. ...................................................... .. 15
Statistical A n aly sis........... .... ............................................................ .......... .. ..... .. 19

R E S U L T S ................................................................................2 0

V eg station Su rv ey s ........................................................................... ..........2 0
C hi-Squ are A naly sis ............................................................... .. ....... ... ...... 2 5

D IS C U S S IO N ........................................................................................4 0

H igh P ine H habitats ................................................................. .............. .... 40
P ine F latw oods H habitats ...................................................................... ..................42

MANAGEMENT IMPLICATIONS ............................................. .......................... 44

APPENDIX................... .. .............. ................... ............. 50

L IT E R A T U R E C IT E D ............................................................................. ....................62

B IO G R A PH IC A L SK E TCH ...................................................................... ..................67








v
















LIST OF TABLES


Table page

1. Habitats and relative burrow densities within burrow survey compartments on the
Low er Suw annee NW R Florida .................................. ............................... ....... 18

2. Descriptive statistics of vegetation cover for each habitat and burrow density category
on the Lower Suwannee NW R, Florida. ...................................... ............... 21

3. Chi-square tests of independence between canopy cover and shrub cover within the 2
habitat categories on the Lower Suwannee NWR, Florida .................................30

4. Chi-square tests of independence between canopy cover and herbaceous ground cover
within the 2 habitat categories on the Lower Suwannee NWR, Florida ................31

5. Chi-square tests of independence between canopy cover and woody ground cover
within the 2 habitat categories on the Lower Suwannee NWR, Florida ................32

6. Chi-square tests of independence between canopy cover and bare ground cover within
the 2 habitat categories on the Lower Suwannee NWR, Florida ...........................33

7. Chi-square tests of independence between canopy cover and detritus ground cover
within the 2 habitat categories on the Lower Suwannee NWR, Florida ................34

8. Chi-square tests of independence between shrub cover and herbaceous ground cover
within the 2 habitat categories on the Lower Suwannee NWR, Florida ................35

9. Chi-square tests of independence between shrub cover and woody ground cover
within the 2 habitat categories on the Lower Suwannee NWR, Florida ................37

10. Chi-square tests of independence between shrub cover and bare ground cover within
the 2 habitat categories on the Lower Suwannee NWR, Florida ...........................38

11. Chi-square tests of independence between shrub cover and detritus ground cover
within the 2 habitat categories on the Lower Suwannee NWR, Florida ................39

12. Analysis of variance results for high pine canopy cover comparisons....................50

13. Analysis of variance results for high pine shrub cover comparisons.......................51

14. Analysis of variance results for high pine woody ground cover comparisons. .........52









16. Analysis of variance results for high pine detritus ground cover comparisons.........54

17. Analysis of variance results for high pine herbaceous ground cover comparisons...55

18. Analysis of variance results for pine flatwoods canopy cover comparisons ............56

19. Analysis of variance results for pine flatwoods shrub cover comparisons ..............57

20. Analysis of variance results for pine flatwoods woody ground cover comparisons..58

21. Analysis of variance results for pine flatwoods bare mineral soil cover comparisons.59

22. Analysis of variance results for pine flatwoods detritus ground cover comparisons.60

23. Analysis of variance results for pine flatwoods herbaceous ground cover
co m p ariso n s............................................................................................ 6 1
















LIST OF FIGURES


Figure p

1. Location of the Lower Suwannee NW R, Florida. ........................................ ..............8

2. Land cover forms on the Lower Suwannee NWR, Florida. ........................................9

3. Location of tortoise burrow and vegetation survey compartments, Lower Suwannee
N W R Florida. .........................................................................10

4. Soil composition of high pine survey compartment categories on the Lower
Suw annee N W R F lorida ............................................................................ ... .... 12

5. Soil composition of pine flatwoods survey compartment categories on the Lower
Suw annee N W R F lorida ............................................................................ ... .... 14

6. Representative pictures of each survey compartment category...............................16

7. Comparisons of canopy cover between burrow densities within each of the pine
community types on the Lower Suwannee NWR, Florida.................. .............22

8. Comparisons of shrub cover between burrow densities within each of the pine
community types on the Lower Suwannee NWR, Florida.................. .............23

9. Comparisons of herbaceous ground cover between burrow densities within each of
the pine community types on the Lower Suwannee NWR, Florida.......................24

10. Comparisons of woody ground cover between burrow densities within each of the
pine community types on the Lower Suwannee NWR, Florida..............................26

11. Comparisons of bare soil ground cover between burrow densities within each of the
pine community types on the Lower Suwannee NWR, Florida..............................27

12. Comparisons of detritus ground cover between burrow densities within each of the
pine community types on the Lower Suwannee NWR, Florida..............................28















Abstract of Thesis Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Master of Science

VEGETATIVE CHARACTERISTICS OF GOPHER TORTOISE (Gopherus
polyphemus) HABITAT ON THE LOWER SUWANNEE NATIONAL WILDLIFE
REFUGE: IMPLICATIONS FOR RESTORATION AND MANAGEMENT OF PINE
COMMUNITIES
By

Stephen E. Barlow

December 2004

Chair: Mark W. Clark
Major Department: Soil and Water Science

Previous land use practices on the Lower Suwannee National Wildlife Refuge

altered natural fire conditions resulting in habitat degradation, especially within the high

pine areas, allowing succession to begin shifting these habitats to mesic hammocks. In

addition, many pine flatwoods sites exist as densely planted slash pine plantations, which

have been relegated to an infrequent winter fire regime, leading to a shrub level

monoculture of gallberry (Ilex glabra) and saw palmetto (Serenoa repens). Changes in

vegetative structure of these prominent pine communities are thought to have lowered the

quality of gopher tortoise habitat. This investigation evaluates the relationship between

vegetative cover of different community strata and the density of gopher tortoise

burrows.

Vegetation cover was measured on sites with high and low gopher tortoise burrow

densities in high pine and pine flatwoods habitats on the refuge from April to July 2004.









The line intercept technique was used to measure canopy cover, shrub cover and ground

cover. Survey compartment soil compositions were compared using Geographic

Information Systems (GIS) data and the county soil surveys. It was predicted that

compartments with high tortoise burrow densities would have a relatively open canopy

and shrub layers, with relatively high herbaceous and bare soil ground covers.

Analysis of variance on vegetation cover data revealed that density of the canopy,

shrub layer, and ground cover types were significantly different between high pine sites

with different tortoise burrow densities. While the pine flatwoods sites exhibited

significant differences in all categories except bare soil, due to a lack of bare soil within

these habitats. Lower canopy cover, shrub cover and higher herbaceous ground cover

characterized sites with high tortoise burrow densities.

Soil analysis revealed higher soil series diversity within the pine flatwoods sites.

This soil series diversity is not represented vegetatively within the current slash pine

plantation monoculture. The lack of relationship between soil characteristics and

vegetation community composition is most likely due to past timber practices that

converted all usable land to pine production with little regard for preserving small

pockets of habitat. Close analysis of soil series should guide land managers' efforts in

restoration of habitats in order to regain pre-European development habitat diversity.

Aggressive techniques to reverse succession on high pine sites are also

recommended. Restoration of the herbaceous ground layer and a reintroduction to

summer fires could then be employed. Timber thinning and a more varied fire regime

emphasizing summer burning would benefit pine flatwoods sites; otherwise continued

habitat succession and degradation will significantly inhibit tortoise populations.















INTRODUCTION


Once the most extensive terrestrial ecosystems in Florida, pine flatwoods and high

pine communities have been heavily influenced by humans, therefore many

characteristics of these communities have changed markedly since European settlement.

Pine communities were extensively timbered starting during the Civil War through today

(Abrahamson and Harnett 1990). Concurrently, human population growth and the

concomitant fragmentation of these vast pine stands have led to a decrease in the extent

and frequency of natural fires. In their natural state, these communities are characterized

by their openness and frequent occurrence of fires (Laessle 1942, Ober 1954, Edmisten

1963, Platt et al. 1988). Though few natural stands closely resemble pre-settlement pine

communities, the consensus is that present stands differ from pre-settlement stands by

having lower fire frequencies, more even age structure, and a denser under-story with

greater shrub cover and less herbaceous cover (Abrahamson and Hartnett 1990, Noel et

al. 1998). In recent times many of these areas have been owned and managed by timber

companies that practice intense plantation production of short rotation pine trees for the

paper industry.

Thus, managers of many public lands in Florida are often faced with restoring and

managing pine communities that have been severely altered. Characteristics of pine

communities under timber production management and fire exclusion include densely

planted even-aged stands of slash pine (Pinus eliotti) often planted in raised,









mechanically bedded rows, with intense herbicide use. Forest stands are also often

prepared using heavy machinery, they have a monotypic shrub under story, raised road

beds, windows, borrow pits, rail tramways and logging roads crisscrossing the forest.

An understanding of the impacts of these former land use practices is necessary for the

formulation of management strategies concerned with restoring habitats to their pre-

European settlement state.

Numerous researchers have described the gradual transition high pine communities

make to mesic hardwood forests and the invasion of non fire-adapted species in the

absence of frequent fire (Veno 1976, Givens et al. 1984, Myers 1985, Myers and White

1987). With the exclusion of fire, fire-adapted species also decline and since they often

serve as the primary fuel source for frequent fires frequency and intensity of natural,

lightning-ignited fires is reduced.

Under natural conditions pine flatwoods are stable and essentially

nonsuccessional due to fire (Abrahamson and Hartnett 1990). When fire is removed from

pine flatwoods, or if the natural frequency or seasonality of fire is altered, flatwoods can

succeed to a variety of vegetation types. Human modifications to the landscape, such as

certain silvicultural practices (logging, clearing and drainage), can also stimulate

successional change. It has been noted by numerous authors that disturbance of the

natural fire frequency is the most common cause of successional changes in pine

flatwoods (Monk 1968, Robbins and Myers 1992, Peroni and Abrahamson 1986).

Loss of pine communities along with fire exclusion and certain silvicultural

practices have negatively impacted numerous wildlife species that characterize pine

communities. Bird species that require open pine forests such as the red-headed









woodpecker (Melanerpes erythrocephalus), brown-headed nuthatch (Sitta pusilla),

loggerhead shrike (Lanius ludovicianus) and eastern bluebird (Sialia sialis) have

experienced declining populations throughout the southeast (Cox 1987). Within high

pine communities, population size and species richness of birds decline noticeably when

these areas are converted to timber production (Repenning and Labisky 1985).

Silviculture practices across the United States have been implicated in the decline or

elimination of at least 26 species of salamanders including the flatwoods salamander

(Ambystoma cingulatum) in Florida (Bury 1983, Brode and Bury 1984, Herrington and

Larsen 1985, Corn and Bury 1989, Welsh 1990, Blymer and McGinnes 1977, Ash 1988,

Buhlmann et al. 1988, Pentranka et al. 1993, Jordan and Mount 1975, Dodd 1991, Dodd

1993, Means et al. 1996).

Due to loss and degradation of habitat, numerous species found within pine

communities are currently listed as species of special concern, threatened or endangered.

One such species is the gopher tortoise (Gopheruspolyphemus), which is a state listed

species of special concern that has experienced a population reduction through loss of

habitat and a lack of fire in existing habitats. Changes in fire frequency in pine

communities are thought to have decreased herbaceous food plants, thereby negatively

influencing gopher tortoise habitat (Landers and Speake 1980, Cox et al. 1987). Since

grasses and forbs constitute the bulk of the gopher tortoise's diet, increased shading and

detritus buildup associated with fire exclusion lead to reduced productivity of these plants

and a decline in tortoise numbers (Garner and Landers 1979, Franz and Auffenberg

1974).









The gopher tortoise is a keystone species-one that holds a critical role in the

ecosystem (Campbell and Christman 1982, Jackson and Milstrey 1988). Their burrows

provide shelter for more than 300 species of obligate and facultative commensals,

including arachnids, insects, reptiles, amphibians, birds, and mammals (Jackson and

Milstrey 1988). Several of these species such as the southeastern indigo snake

(Drymarchon couperi), the gopher frog (Rana capitol) and the Florida mouse (Podomys

floridanus) are threatened species or species of special concern. Decline of the gopher

tortoise has negatively affected these secondary burrow users because there are fewer

burrows available (Berish 2001).

A primary factor affecting density of gopher tortoises is habitat quality, particularly

as it relates to food availability as influenced by fire and primary succession (O'Meara

and Abbott 1987). However, caution should be used when comparing tortoise densities

to habitat variables as dispersion of gopher tortoise burrows within available habitats is

poorly understood (Cox et al. 1987). To more accurately determine the quality of habitat

measures of mean reproductive success, survival and number of individuals in each age

class are needed to determine quality of the habitat (Van Home 1983). Poor habitat

quality may cause tortoises to form dense colonies in small patches of suitable habitat,

thus a survey which is conducted at too small of a scale, or multiple scales may over

estimate tortoise densities across the habitat or flaw site comparisons (McCoy and

Mushinsky 1995).

To better understand the status of tortoise populations on the Lower Suwannee

National Wildlife Refuge Refuge (LSNWR) and their habitat preferences, we conducted

tortoise burrow surveys in 2002 in a manner similar to that of Auffenberg and Franz









(1982) and in Cox et al. (1987). In this survey, burrows were recorded as active, inactive

and old. Active burrows were marked using a handheld GPS unit. Marked burrows were

then projected as a layer file using Arcview 8.3 geographic information systems (GIS)

software to compare burrow colony locations with habitat and soil features.

Through GIS analysis it appeared refuge tortoises were congregated in colonies

within various survey compartments. This aggregating of tortoises into pockets suggests

preferential habitat within the broader survey compartment possibly representing

remnants of formerly large areas of habitat (McCoy and Mushinsky 1995). This

clumping of tortoise burrows makes the task of density estimation across refuge habitats

imperfect at best, unless additional environmental variables influencing tortoise

distribution can be identified. Furthermore, tortoise burrow congregations did not appear

to relate to particular GIS habitat or soil layers, thus the level of detail within the GIS

data layers were lacking for discernment of these apparent site choices by tortoises.

The goal of this study was to investigate what additional environmental variables

might be influencing tortoise distribution and what changes in management might be

implemented to enhance tortoise habitat in pine communities on the LSNWR. Specific

hypothesis to be addressed under this goal include the following:

1) To determine if differences are present between vegetation structural
characteristics (ground cover, shrub cover and canopy cover) among
tortoise burrow survey compartments that exhibited relatively high or low
burrow densities.
2) To determine if a relationship is present between the amount of canopy and
shrub cover to ground cover forms or if these variables are independent.

It was predicted that areas with low tortoise burrow density would exhibit high

canopy and shrub cover with low herbaceous ground coverage, while areas with high

tortoise burrow densities would contain structural characteristics of low canopy and shrub






6


cover with higher herbaceous ground coverage. It was further predicted that the amount

of ground cover forms observed would be dependent upon the amount of shrub or canopy

cover present.















MATERIAL AND METHODS

Study Area

The study was conducted on the Lower Suwannee NWR, a 21,500-hectare refuge

located in the heart of north Florida's Big Bend region in Dixie and Levy counties

(Figurel). Situated along the Gulf of Mexico the refuge is bisected by the Suwannee

River, which flows southward through the refuge for 32 kilometers before emptying into

the Gulf of Mexico. Purchased by the federal government in 1979 from various timber

companies, the area still bears signs from decades of timber production management.

The topography of the study area is relatively flat, with dominant habitats including

salt marsh, southern hydric hardwood, bottomland forest, and pine plantation (Figure 2)

(Sykes et al. 1999). This diversity of habitats on the area is matched by the variety of soil

orders found on the refuge. Of the seven soil orders in Florida, the refuge contains six,

with Histosols being the most common (Slabaugh et al. 1996, Liudahl et al. 2003). The

refuge contains 43 individual soil series, many of which are poorly drained or very poorly

drained and strongly acid (pH 5.1-5.5) to extremely acid (pH 4.0-4.4) (Slabaugh et al.

1996, Liudahl et al. 2003).

Survey Compartments

Survey transects selected for this study were evenly divided between high pine

habitats and pine flatwood habitats as well as by tortoise burrow density (Figure 3). Ten

transects were randomly selected from high pine habitats: 5 transects in high tortoise


















































I0 I I 1 0 I
0 37,5 75 150 Kilometers


S 3.75 7.5 15 Kileters
0 3.75 7T5 15 Kilometers


Figure 1. Location of the Lower Suwannee NWR, Florida.






















































Figure 2. Land cover forms on the Lower Suwannee NWR, Florida.

















HIGH PIE~
mHIGjaH BURRO
DNITY^J- ^
HIGH PIN


r


^Imie-
^*T1ura.; ''f -


PIN LTOD


Figure 3. Location of tortoise burrow and vegetation survey compartments, Lower
Suwannee NWR, Florida.









burrow density compartments and 5 transects within low tortoise burrow density

compartments. SimilarilylO transects were randomly selected from pine flatwoods

compartments, also equally divided by tortoise burrow density.

High pine survey compartments consisted of even-aged mechanically planted long-

leaf pines, planted in 1993 (Figure 6). Thus, the species, age, and basal area of the

planted pine canopy component were controlled for between burrow density

compartments. Interspersed within the planted long-leaf pines were turkey oak (Quercus

laevis), live oak (Quercus virginiana) and a shrub layer containing immature oaks

(Quercus spp.), sparkleberry (Vaccinium arboreum) and dog fennel (Eupatorium

compositifolium). Herbaceous ground cover within the compartments contained deer's

tongue (Carphephorus odoratissimus), greenbriar (Smilax spp.), partridge pea

(Chamaecristapilosa) wire grass and associated forbs and grasses.

The refuge practices an aggressive prescribed burning program, introducing fire to

approximately 600 hectares annually (pers. comm., K. McCain Chiefland, FL). With the

majority of these being dormant season burns, within densely planted flatwoods habitats.

Though occasional attempts have been made to burn within the high pine habitats, lack of

a uniformly distributed fuel source throughout has made most of theses burns "spotty"

and anemic at best (pers. comm., K. McCain Chiefland, FL). Following the planting of

long-leaf pines on these areas, a lack of fire for at least 10 years has led to the

aforementioned encroachment of non-pyric species such as oaks. Needle cast from the

planted pine component within these areas currently does not provide adequate fuels to

carry fire under safe weather conditions. Also, with the current tree species composition





































35%


HIGH PINE
LOW BURROW DENSITY
SOILS




36%
SRidgewood
3 Leon (Spodosol)
4 r Clara (Spodosol)


29%
B


Figure 4. Soil composition of high pine survey compartment categories on the Lower
Suwannee NWR, Florida. A) Soil composition of high pine compartments
with high tortoise burrow densities. B) Soil composition of high pine
compartments with low tortoise burrow densities.


HIGH PINE
HIGH BURROW DENSITY
SOILS




--- L 31%


* Ridgewood
* Ortega











and densities, the chances of an intense, unmanageable scrub fire killing all the trees is a

serious management concern. Soils within high pine survey compartments were

represented by the Clara and Leon series (Figure 4). All soils are very deep, moderately

well drained soils of upland sites (Liudahl et al. 2003).

Pine flatwoods survey compartments consisted of even-aged slash pines, planted

in 1978 by Buckeye Cellulose Inc., (Figure 6). Thus, the species, age, and basal area of

the planted pine canopy component were controlled for between burrow density

compartments. Isolated cypress (Taxodium distichum) wetlands are widely interspersed

within these habitats but were not included within survey compartments. Years of fire

exclusion followed by only dormant season fires has helped to foster the dense stands of

gallberry and saw-palmetto common within refuge pine flatwoods communities,

inhibiting production of herbaceous ground covers forbss and grasses). A lack of

growing season burns could facilitate the establishment of dense gallberry stands, since

growing season burns appear to hinder gallberry's resprouting ability, while time since

the last fire has been shown to increase the density of saw palmetto (Hughes and Knox

1964, Maliakal et al. 2000).

The sites were bedded for planting, but other management techniques conducted

on these sites by the timber company is unknown. The U.S. Fish and Wildlife Service

assumed management of the stands in 1985, since refuge acquisition the sites have

received several dormant season prescribed burs. The plantations also received third

row thinnings in 1997-98.































Pine Flatwoods
Low Burrow Density
Soils


OU /0 34%




0% 111o %
26%


MChaires (Spodosol)
* Resota (Entisol)
ORidgewood (Entisol)
OBodiford (Alfisol)
* Leon (Spodosol)

B


Figure 5. Soil composition of pine flatwoods survey compartment categories on the
Lower Suwannee NWR, Florida. A) Soil composition of pine flatwoods
compartments with high tortoise burrow densities. B) Soil composition of
pine flatwoods compartments with low tortoise burrow densities.


Pine Flatwoods
High Burrow Density
Soils


6%

34% m Smyrna (Spodosol)
Placid (Inceptisol)
E Adamsville (Entisol)
49% O] Ridgewood (Entisol)
22% m Leon (Spodosols)
S9%
A









Soil composition of pine flatwoods survey compartments contained primarily Spodosols,

though Entisols, Inceptisols, and Alfisols were also present (Figure 5). The soils of

flatwoods survey compartments were less uniform and more interspersed than high pine

compartments. Spodosol's were represented by Smyrna, Chaires and Leon series, which

have similar characteristics of being deep to very deep and poorly drained. While

Entisol's were represented by Adamsville, Ridgewood and Resota series which are very

deep, moderately drained psamments. Inceptisols and Alfisols occupied only 0.33

hectares within the flatwoods compartments and were represented by Bodiford and Placid

soil series (Slabaugh et al. 1996, Liudahl et al. 2003).

Vegetation Surveys

Vegetation surveys were conducted from April through July 2004. Canopy cover,

shrub cover and ground cover were determined within survey compartments that had

exhibited relatively high or low tortoise burrow densities during the 2002 tortoise burrow

surveys. A line-intercept technique was used to estimate percent canopy cover and shrub

cover along each transect (Higgins et al. 1994). A 10-meter line was stretched between

two stakes at a height of 1-meter. Percent canopy cover was measured by observing the

total length of line intercepted by vertical projections of the canopy. Percent shrub cover

was measured by observing the total length of line intercepted by plants touching the line.

Only shrubs touching the nylon line were included, whereas the entire plant intercepting

the vertical line was counted as percent cover. Percent ground cover was obtained using

the pen-drop technique, where a pen was dropped at 1-meter increments along the line

and the dominant ground cover touching the pen was recorded. Ground cover forms

were grouped at each sampling location into the following 4 categories: (1) Woody (i.e.,

woody shrubs, tree seedlings), (2) Herbaceous (i.e., non-woody plants, legumes, forbs







































Figure 6. Representative pictures of each survey compartment category: A) High pine
with high tortoise burrow density, B) High pine with low tortoise burrow
density, C) Pine flatwoods with high tortoise burrow density and D) Pine
flatwoods with low tortoise burrow density.









and grasses), (3) Detritus (i.e., non-living plant material, leaves, needles) and (4)Bare(i.e.,

mineral soil). Vegetation surveys were conducted along the same transects used during

the 2002 tortoise burrow surveys. Tortoise burrow survey transects were 200-meters

long by 20-meters wide, covering approximately .4 hectares. Survey transects were

grouped in compartments, which were placed within suitable tortoise habitats using the

refuge's GIS habitat and soils layers (Cox et al. 1987, Breininger et al. 1988). Following

the 2002 tortoise burrow survey, compartments were labeled either high burrow density

or low burrow density; compartments were also arranged within 2 habitat categories-pine

flatwoods or high pine, see Table 1 (Abrahamson and Hartnett 1990, Myers 1990).

Burrow densities for each compartment were determined by dividing the

compartment area by the total number of both active and inactive burrows observed,

burrow densities >lburrow/hectare were designated as high density while those

compartments with
Using a random numbers table, survey transects within 8 distinct compartments were

randomly selected for vegetation cover surveys (Ott 1983).

Five survey transects were selected from each habitat and burrow density category,

(i.e., 5 transects taken from high pine/high burrow density compartments were compared

with 5 transects taken from high pine/low burrow density compartments, while 5

transects were taken from flatwoods pine/high burrow density compartments were

compared with 5 transects taken from flatwoods pine/low burrow density compartments).

Since all survey compartments had been mechanically planted as even-aged stands, I was

able to control for timber basal area, age class and species. Compartments also contained




















Table 1. Habitats and relative burrow densities within burrow survey compartments on
the Lower Suwannee NWR, Florida as determined during the 2002 survey.


Compartment/Transects Habitat Burrows/Hectare
1/1-20* Pine Flatwoods 1.13
1/21-30 Pine Flatwoods 1.00
1/31-40 Pine Flatwoods 0.50
1/41-50 Pine Flatwoods 1.25
1/51-60 Pine Flatwoods 0.25
1/61-70 Pine Flatwoods 0.00
2/1-10 Pine Flatwoods 1.00
2/11-17 Pine Flatwoods 0.00
3/1-6 Pine Flatwoods 0.00
4/1-16 Pine Flatwoods 0.94
4/17-36 Pine Flatwoods 0.88
4/37-53 Pine Flatwoods 0.74
6/1-15 Pine Flatwoods 0.00
6/16-25 Pine Flatwoods 0.00
7/1-10* Pine Flatwoods 0.00
7/11-16* Pine Flatwoods 0.00
8/1-15* Pine Flatwoods 3.50
8/16-25* Pine Flatwoods 3.50
9/1-20* High Pine 4.25
9/21-35* High Pine 5.17
9/36-45 High Pine 1.00
9/46-55 High Pine 2.50
9/56-61* High Pine 0.83
9/62-70 High Pine 0.83
*Compartments containing randomly selected vegetation surveys.


Burrow Density
High
Low
Low
High
Low
Low
Low
Low
Low
Low
Low
Low
Low
Low
Low
Low
High
High
High
High
Low
High
Low
Low









very similar bum histories; with no compartments receiving a growing season bum prior

to this study. Vegetation cover percentages were recorded at 20-meter intervals along the

200-meter burrow survey transects. At 20-meter intervals a 10-meter vegetation cover

transect was surveyed perpendicular to and on both sides of the burrow survey transect.

This provided 20, 10-meter vegetation cover transects spaced throughout each burrow

survey transect.

Statistical Analysis

Microsoft Excel 2000 was used for statistical analysis. I arcsine transformed

percentages (i.e., percent canopy cover, shrub cover and ground cover) prior to analysis

so they would better meet the normality assumption of analysis of variance (Ott 1983).

The assumption of homogeneity of population variances was not considered critical since

the sample sizes were equal. I considered a 0.05 probability level statistically significant

for all tests. A chi-square test of independence was used to determine if the amount of

the 4 ground cover forms were independent of the amount of shrub or canopy cover

observed. For the chi-square test of independence canopy, shrub and the 4 ground cover

forms data was grouped into 2 categories: High for values > 0.5, and Low for values <

0.5. Data represented in descriptive statistics or used for the chi-square test was not

transformed.















RESULTS


Vegetation Surveys

Analysis of canopy closure data indicates significant differences when comparing

between burrow densities within each of the pine community types (Figure 6). Canopy

cover differed significantly between high pine burrow density category sites (P=0.000).

Canopy cover also differed between pine flatwoods high and low tortoise burrow density

sites (P=0.000). Within both habitat categories, low tortoise burrow density sites had

greater canopy closure than respective high tortoise burrow density sites. The mean

values prior to transformation, for each habitat and tortoise burrow density category can

be found in table 2.

Shrub cover within the two pine communities also differed significantly between

high pine tortoise burrow density category sites (P=0.000). Shrub cover also differed

between pine flatwoods high and low tortoise burrow density sites (P=0.000). Within

both habitat categories, low tortoise burrow density sites had greater shrub cover

densities than respective high tortoise burrow density sites (Figure 7).

Herbaceous ground cover differed significantly between high pine tortoise burrow

density category sites (P=0.000). Herbaceous ground cover also differed between pine

flatwoods high and low tortoise burrow density sites (P=0.000). Within both habitat

categories high tortoise burrow density sites had greater herbaceous ground cover

densities than respective low tortoise burrow density sites (Figure 8).





















Table 2. Descriptive statistics of vegetation cover for each habitat and burrow density
category on the Lower Suwannee NWR, Florida.


HIGH PINE
Canopy
Burrow Cover
Density Mean SD


Shrub
Cover
Mean SD


Herb
Cover
Mean SD


Woody
Cover
Mean SD


Bare
Soil
Mean SD


Detritus
Cover
Mean SD


High(n= 100) 0.12
95% CI 0.03

Low(n=100) 0.43
95% CI 0.05


PINE FLATWOODS

High(n=100) 0.28 0.28
95% CI 0.05

Low (n=100) 0.52 0.30
95% CI 0.06


0.13 0.10 0.12
0.02

0.03 0.72 0.22
0.04


0.43 0.23 0.13 0.14 0.26 0.19 0.17 0.21
0.05 0.03 0.04 0.04

0.14 0.14 0.43 0.20 0.07 0.11 0.36 0.22
0.03 0.04 0.02 0.04


0.11 0.15 0.53 0.30 0.23
0.03 0.06 0.03


0.00 0.02 0.20 0.21
0.00 0.04


0.81 0.12 0.19 0.16 0.51 0.22 0.00 0.00 0.30 0.22
0.02 0.03 0.04 0.00 0.04



































High Pine
High Burrow
Density


High Pine
Low Burrow
Density


Pine Flatwoods
High Burrow
Density


Pine Flatwoods
Low Burrow
Density


Figure 7. Comparisons of canopy cover between burrow densities within each of the pine
community types on the Lower Suwannee NWR, Florida. Box contains upper
and lower quartiles, dotted line indicates mean, dash indicates median,
whiskers connect box to largest and smallest values. Data was not
transformed.


































High Pine
High Burrow
Density


High Pine
Low Burrow
Density


Pine Flatwoods
High Burrow
Density


Pine Flatwoods
Low Burrow
Density


Figure 8. Comparisons of shrub cover between burrow densities within each of the pine
community types on the Lower Suwannee NWR, Florida. Box contains upper
and lower quartiles, dotted line indicates mean, dash indicates median,
whiskers connect box to largest and smallest values. Data was not
transformed.


1.2


1.0


0.8 -


0.6 -


0.4 -


0.2 -


0.0 -


































High Pine
High Burrow
Density


High Pine
Low Burrow
Density


Pine Flatwoods
High Burrow
Density


Pine Flatwoods
Low Burrow
Density


Figure 9. Comparisons of herbaceous ground cover between burrow densities within
each of the pine community types on the Lower Suwannee NWR, Florida.
Box contains upper and lower quartiles, dotted line indicates mean, dash
indicates median, whiskers connect box to largest and smallest values. Data
was not transformed.


I I I I











Woody ground cover differed between high pine tortoise burrow density category

sites (P=0.000). Woody ground cover also differed between pine flatwoods high and low

tortoise burrow density sites (0 P=0.000). Within both habitat categories, low tortoise

burrow density sites had greater woody ground cover densities than respective high

burrow density sites (Figure 9).

Amount of exposed bare mineral soil differed between high pine tortoise burrow

density category sites (P=0.000), as illustrated in Figure 10. However, amount of

exposed bare mineral soil did not differ between pine flatwoods high and low tortoise

burrow density sites (P=0.320). Within high pine habitat, sites with high tortoise burrow

densities had more exposed bare mineral soil than respective low burrow density sites.

Within pine flatwoods sites, areas of bare mineral soil were infrequent, thus accounting

for the lack of difference between sites.

The amount of detritus ground cover differed between high pine tortoise burrow

density category sites (P=0.000). The amount of detritus ground cover also differed

between pine flatwoods high and low tortoise burrow density sites (P=0.004). Within

both habitat categories low tortoise burrow density sites had more detritus ground cover

than respective high burrow density sites (Figure 11).

Chi-Square Analysis

Chi-square tests of independence results for the association of canopy cover to

shrub cover within both habitat categories are shown in table 3. Shrub cover was

dependent upon canopy cover in both high pine and pine flatwoods categories

respectively (2=40.7, 24.7).



































High Pine
High Burrow
Density


High Pine
Low Burrow
Density


Pine Flatwoods
High Burrow
Density


Pine Flatwoods
Low Burrow
Density


Figure 10. Comparisons of woody ground cover between burrow densities within each of
the pine community types on the Lower Suwannee NWR, Florida. Box
contains upper and lower quartiles, dotted line indicates mean, dash indicates
median, whiskers connect box to largest and smallest values. Data was not
transformed.


I I I I

































High Pine
High Burrow
Density


High Pine
Low Burrow
Density


Pine Flatwoods Pine Flatwoods
High Burrow Low Burrow
Density Density


Figure 11. Comparisons of bare soil ground cover between burrow densities within each
of the pine community types on the Lower Suwannee NWR, Florida. Box
contains upper and lower quartiles, dotted line indicates mean, dash indicates
median, whiskers connect box to largest and smallest values. Data was not
transformed.

































High Pine
High Burrow
Density


High Pine
Low Burrow
Density


Pine Flatwoods
High Burrow
Density


Pine Flatwoods
Low Burrow
Density


Figure 12. Comparisons of detritus ground cover between burrow densities within each
of the pine community types on the Lower Suwannee NWR, Florida. Box
contains upper and lower quartiles, dotted line indicates mean, dash indicates
median, whiskers connect box to largest and smallest values. Data was not
transformed.









Chi-square tests of independence results for the association of canopy cover to

herbaceous ground cover, within both habitat categories are shown in table 4. Notice

within the pine flatwoods habitat category canopy cover and herbaceous ground cover are

independent (x=2). While herbaceous ground cover was not independent of canopy

cover within the high pine category (2=8.9).

Chi-square tests of independence results for the association of canopy cover to

woody ground cover, within both habitat categories are shown in table 5. High chi-

square values for both high pine and pine flatwoods categories respectively (X2= 17.4,

17.8) indicate the amount of woody ground cover was not independent of the

accompanying canopy cover. Chi-square tests of independence results for the association

of canopy cover to bare ground cover, within both habitat categories are shown in table 6.

The chi-square values suggest bare ground cover is not independent within the high pine

habitats (x= 5.6), however within the pine flatwoods category bare ground cover is

independent of canopy cover (x2= 0.01). Chi-square tests of independence results for the

association of canopy cover to detritus ground cover are shown in table 7. Within both

habitat categories detritus ground cover was dependent upon the amount of canopy cover,

with a high pine X2 value of 7 and a pine flatwoods of 4.9.

Chi-square tests of independence results for the association of shrub cover to

herbaceous ground cover, within both habitat categories are shown in table 8. Within

both habitat categories herbaceous ground cover was not independent of shrub cover

(high pine x= 37.9, pine flatwoods x2= 60.3). Chi-square tests of independence results

for the association of shrub cover to woody ground cover, within both habitat categories

are shown in table 9. Indicating the amount of woody ground cover was not independent












Table 3. Chi-square tests of independence between canopy cover and shrub cover within
the 2 habitat categories on the Lower Suwannee NWR, Florida.


High Pine
Vegetation
Cover


>50% Canopy

<50% Canopy


Totals

X2= 40.7

df= (2-1)(2-1)=

P (a=.05)= 3.84


Pine Flatwoods
Vegetation
Cover

>50% Canopy

<50% Canopy


Totals

X2= 24.7

df= (2-1)(2-1)=


>50% Shrub
Observed (expected)


39(20.7)

50(70.2)


>50% Shrub
Observed (expected)


58(41.6)

45(63.4)


<50% Shrub
Observed (expected)


6(25.8)

105(87.4)


200


<50% Shrub
Observed (expected)


21(39.2)

76(59.8)


200


P (a=.05)= 3.84


Totals


Totals









Table 4. Chi-square tests of independence between canopy cover and herbaceous ground
cover within the 2 habitat categories on the Lower Suwannee NWR, Florida.


High Pine
Vegetation
Cover


>50% Canopy

<50% Canopy


Totals


>50% Herb
Observed (expected)


3(11)


44(37.4)


<50% Herb
Observed (expected) Totals


42(35.4)

111(120.1)


200


X2= 8.9


df= (2-1)(2-1)= 1

P (a=.05)= 3.84


Pine Flatwoods
Vegetation
Cover

>50% Canopy

<50% Canopy


Totals

X2= 2

df= (2-1)(2-1)=


>50% Herb
Observed (expected)


21(26.4)

44(40.3)


<50% Herb
Observed (expected)


58(54.4)

77(83)


200


P (a=.05)= 3.84


Totals









Table 5. Chi-square tests of independence between canopy cover and woody ground
cover within the 2 habitat categories on the Lower Suwannee NWR, Florida.


>50% Woody
Observed (expected)


<50% Woody
Observed (expected)


>50% Canopy

<50% Canopy


Totals

X2= 17.4

df= (2-1)(2-1)= 1

P (a=.05)= 3.84


Pine Flatwoods


Vegetation
Cover


22(11.5)

28(39)


>50% Woody
Observed (expected)


23(34.5)

127(117)


<50% Woody
Observed (expected)


>50% Canopy

<50% Canopy


Totals

X2= 17.8

df= (2-1)(2-1)=

P (a=.05)= 3.84


High Pine
Vegetation
Cover


Totals


45

155


200


39(25.6)

25(39)


Totals


40(54.4)

96(83)


200









Table 6. Chi-square tests of independence between canopy cover and bare ground cover
within the 2 habitat categories on the Lower Suwannee NWR, Florida.


>50% Bare
Observed (expected)


<50% Bare
Observed (expected)


>50% Canopy

<50% Canopy


Totals

x2= 5.6

df= (2-1)(2-1)= 1

P (a=.05)= 3.84


Pine Flatwoods


Vegetation
Cover


>50% Canopy

<50% Canopy


Totals

X2= 0.01

df= (2-1)(2-1)=

P (a=.05)= 3.84


0(4.1)

18(14)


45(41.9)

137(142)


>50% Bare <50% Bare
Observed (expected) Observed (expected)


0(0)

0(0)


79(80)


121(122)


200


High Pine
Vegetation
Cover


Totals


45

155


200


Totals


200









Table 7. Chi-square tests of independence between canopy cover and detritus ground
cover within the 2 habitat categories on the Lower Suwannee NWR, Florida.


>50% Detritus
Observed (expected)


<50% Detritus
Observed (expected)


>50% Canopy

<50% Canopy


Totals

2= 7

df= (2-1)(2-1)= 1

P (a=.05)= 3.84


Pine Flatwoods


Vegetation
Cover


>50% Canopy

<50% Canopy


Totals

x2= 4.9

df= (2-1)(2-1)=

P (a=.05)= 3.84


17(10.6)

29(35.9)


28(35.4)

126(120)


>50% Detritus <50% Detritus
Observed (expected) Observed (expected)


11(17.6)

33(26.8)


68(62.4)

88(95.2)


High Pine
Vegetation
Cover


Totals


45

155


200


Totals


200









Table 8. Chi-square tests of independence between shrub cover and herbaceous ground
cover within the 2 habitat categories on the Lower Suwannee NWR, Florida.


High Pine
Vegetation
Cover


>50% Shrub

<50% Shrub


Totals

X2= 37.9

df= (2-1)(2-1)= 1

P (a=.05)= 3.84


Pine Flatwoods
Vegetation
Cover

>50% Shrub

<50% Shrub


Totals

X2= 60.3

df= (2-1)(2-1)=


>50% Herb
Observed (expected)


2(23.4)

45(29.1)


>50% Herb
Observed (expected)


5(30.7)

54(29.4)


<50% Herb
Observed (expected)


87(66.6)

66(82.9)


200


<50% Herb
Observed (expected)


98(73.3)

43(69.1)


P (a=.05)= 3.84


Totals


Totals


103

97


200









the amount of shrub cover. With high chi-square values reached within both habitat

categories (high pine x= 60.3, pine flatwoods x= 50.5). Chi-square tests of

independence results for the association of shrub cover to bare ground cover, within both

habitat categories are shown in table 10. Within the high pine habitats bare ground cover

was not independent of shrub cover (x= 15.9), while the pine flatwoods category was

unable to show independence (x= 0), most probably due to a lack of bare ground within

this habitat.

Chi-square tests of independence results for the association of shrub cover to

detritus ground cover, within both habitat categories are shown in table 11. Within both

high pine and pine flatwoods categories, respectively the amount of detritus ground cover

was not independent from the amount of accompanying shrub cover (X2= 16.4 and 4.6).









Table 9. Chi-square tests of independence between shrub cover and woody ground
cover within the 2 habitat categories on the Lower Suwannee NWR, Florida.


High Pine
Vegetation
Cover


>50% Shrub

<50% Shrub


Totals

X2= 60.3

df= (2-1)(2-1)=

P (a=.05)= 3.84


Pine Flatwoods
Vegetation
Cover

>50% Shrub

<50% Shrub


Totals

X2= 50.5

df= (2-1)(2-1)=


>50% Woody
Observed (expected)


46(22.5)

4(28)


>50% Woody
Observed (expected)


56(33.3)

7(31.4)


<50% Woody
Observed (expected)


43(67.5)

107(84)


<50% Woody
Observed (expected)


47(71.8)

90(67.6)


200


P (a=.05)= 3.84


Totals

89

111


200


Totals









Table 10. Chi-square tests of independence between shrub cover and bare ground cover
within the 2 habitat categories on the Lower Suwannee NWR, Florida.


High Pine
Vegetation
Cover

>50% Shrub

<50% Shrub


Totals


>50% Bare <50% Bare
Observed (expected) Observed (expected)


0(8.1)

18(10)


89(81.9)

93(101.9)


182


X2= 15.9


df= (2-1)(2-1)=

P (a=.05)= 3.84


Pine Flatwoods
Vegetation
Cover

>50% Shrub

<50% Shrub


Totals

X2=0

df= (2-1)(2-1)=


>50% Bare
Observed (expected)


0(0)

0(0)


<50% Bare
Observed (expected) Totals


103(103)


97(97)


200


P (a=.05)= 3.84


Totals

89

111


200











Table 11. Chi-square tests of independence between shrub cover and detritus ground
cover within the 2 habitat categories on the Lower Suwannee NWR, Florida.


High Pine
Vegetation
Cover


>50% Shrub

<50% Shrub


Totals

X2= 16.4

df= (2-1)(2-1)= 1

P (a=.05)= 3.84


Pine Flatwoods
Vegetation
Cover

>50% Shrub

<50% Shrub


Totals

x2= 4.6

df= (2-1)(2-1)=


>50% Detritus
Observed (expected)


32(20.7)

13(25.8)


>50% Detritus
Observed (expected)


29(22.9)

15(21.6)


<50% Detritus
Observed (expected)


Totals


57(70.2)

98(87.4)


200


<50% Detritus
Observed (expected) Totals


74(81.1)

82(76.4)


P (a=.05)= 3.84















DISCUSSION

Vegetation cover survey results showed significant differences in all vegetation

categories except the bare soil ground cover form, since bare soil was uncommon within

the pine flatwoods compartments. These results supported the hypothesis that

compartments with high herbaceous ground cover and low shrub/canopy covers

supported higher densities of gopher tortoise burrows. While areas with low burrow

densities had higher shrub and canopy covers with lower herbaceous ground cover. Chi-

square results allowed the rejection of the null hypothesis that ground cover densities

were independent of canopy and shrub cover densities in almost all categories. Though

within the flatwoods habitats herbaceous and bare ground covers were not dependent

upon canopy cover. Also, within the flatwoods habitats bare ground cover was not

dependent upon the amount of shrub cover. The following discussion describes these

findings in the context of the respective habitat categories.

High Pine Habitats

In high pine habitats on the Lower Suwannee NWR, tortoise burrow densities were

higher within survey compartments with the most open shrub layers. A lack of burrows

within a survey compartment was associated with both shrub and canopy closure. The

shrub and canopy layers within low tortoise burrow density compartments were

characterized by a dense covering of young oaks. These successional changes within the

shrub and canopy layers appear to have led to lower herbaceous ground cover, less areas

of exposed bare soil with higher woody and detritus ground covers observed. Within









high pine habitat compartments paired comparisons indicate the strongest association

between tortoise burrow density and vegetation cover was at the shrub layer, this is likely

because of the rather young age of the planted pine component (11 years) where light

limitation to understory vegetation due to canopy closure has not yet occurred.

The vegetation cover differences, and therefore tortoise burrow density differences,

between survey compartments cannot be fully explained by past management practices.

The U. S. Fish and Wildlife Service assumed management of the lands containing the

high pine survey compartments in 1990, from the Georgia Pacific timber company. Soon

after refuge acquisition the area was completely clear-cut of planted slash pines by

Georgia Pacific in accordance with a deed agreement. After clear cutting, refuge

personnel began mechanical, V-blade planting of the sites to long leaf pine in 1993.

Intensive site prepping was not conducted prior to the 1993 plantings. A detailed

management history, including methods of site prep, vegetation control and planting

techniques on these sites prior to 1990 is unknown. Though some site prep techniques

could entomb tortoises, such as the piling of large windows directly on a burrow, gopher

tortoises are able to dig out of impacted burrows following certain types of site

preparation on sandy soils (Landers and Buckner 1981, Diemer and Moler 1982). Thus,

it is unlikely tortoises entombment can explain differences in tortoise density noted in the

2002 survey.

One explanation for the observed vegetation cover differences between survey

compartments is available soil moisture. The GIS soil analysis revealed that 29% of the

two low tortoise burrow density compartments occurred in a Leon soil series. This soil

series has a higher moisture content than other soils series present in this survey









compartment. However this particular Spodosol soil is very deep and has high

permeability in the A and E horizon with moderate permeability in the Bh horizon. Also,

plant community composition observed did not reflect an extreme difference in available

moisture compared to other high pine sites, as more mesic or flatwoods type vegetation

such as gallberry (Ilex glabra) or ericaceous shrubs were not observed to dominate any of

the high pine sites. A working hypothesis to be tested in the future is that higher soil

moisture contents may have increased the rate of shrub and canopy closure within these

sites, but that soil moisture differences were not dramatic enough to shift community

composition towards flatwoods type plants.

Pine Flatwoods Habitats

Within pine flatwoods compartments surveyed, differences in vegetation cover and

gopher tortoise burrow densities were most strongly associated with the amount of shrub

cover. Observed shrub cover was very low in compartments characterized as high

tortoise burrow density, while the opposite was observed in low tortoise burrow density

compartments. Since all compartments contained planted slash pines of very similar ages

and basal areas, canopy differences observed were related to canopy closure by plants

within the shrub layer. This successional closure of the shrub and canopy layers

presumably caused lower herbaceous ground cover along with higher woody and detritus

ground covers observed within the low tortoise density compartments.

The shrub cover observed in low tortoise burrow density compartments is

characterized by dense stands of gallberry and saw-palmetto with intermixed oaks.

Shifting to growing season burns may reduce the density of saw-palmetto and gallberry

stands. It has been theorized that top kill of many species early in the growing season can

halt carbohydrate production when carbohydrate reserves normally in the root system are









at their lowest level, thereby increasing kill (Waldrop et al. 1987). However, other

studies suggest that even after repeated, early growing season burns these very fire

resilient species will not be eradicated (Hough 1968, Hughes and Knox 1964). Though

growing season burns should decrease their densities due to the stress placed on

carbohydrate reserves.

Carbohydrate reserves in the rhizomes of gallberry and saw-palmetto have been

found at their lowest in August (Hough 1968, Hughes and Knox 1964). Therefore, if

the management goal is to decrease the densities of these two species late growing season

burns may be most effective.

As with the high pine survey compartments the observed differences in vegetation

cover can not be fully explained by past management practices; most probably, slight

differences in soil moisture may have accelerated succession within certain

compartments.















MANAGEMENT IMPLICATIONS

Gopher tortoise burrow densities were highest in high pine and pine flatwoods

communities on the Lower Suwannee NWR when shrub and canopy cover was relatively

low and herbaceous ground cover, as well as areas of bare soil, were relatively high.

Gopher tortoise densities are higher in open areas with herbaceous ground cover than in

brushy, shaded sites; the former have patches of bare ground needed for nest excavation

and also provide abundant herbaceous vegetation for feeding (Cox et al. 1987). This type

of habitat can be promoted by growing season fires (Robbins and Meyers 1992).

Furthermore, it has been suggested that growing season fires might increase the amount

of food available in late summer when food quality is declining and would provide food

conditions for new hatchlings, which emerge in late summer and early fall (Cox et al.

1987).

Under current conditions burning is problematic, if not impossible within high pine

areas on the refuge due to years of habitat degradation from a lack of fire. Fire

suppression has allowed these areas to begin succession to a mesic hardwood forest. This

process of succession has concentrated gopher tortoise populations to the higher, drier

sites, where inevitable succession is lagging behind more moist sites. Mesic hardwood

forest conditions are characterized by higher shading, greater detritus accumulation, and

less herbaceous ground cover than natural high pine forests. Since these areas have been

excluded from fire for at least 15 years, a lack of ground fuel greatly restricts a fire's









intensity and its ability to spread thereby rendering fire alone somewhat ineffective as a

management tool for restoration of these habitats.

Two alternate methods for restoring high pine habitats on the refuge include use of

a selective herbicide to remove hardwoods and restoration plantings. The utility of a

selective herbicide such as hexazinone, has been demonstrated for restoration of longleaf

pine communities (Hay-Smith and Tanner 1994). Hexazinone can facilitate the release of

wiregrass and aid in the reduction of scrub oak populations without damage to other

woody and herbaceous vegetation. Current research calls for forest managers to use

hexazinone rates ranging from 0.84 to 1.68Kg/ha, in liquid form applied in a grid pattern

to the soil surface. Applications should be made prior to rainfall, as rain is required to

distribute the herbicide through the root column. Broadcast applications of granular

hexazinone are discouraged as this may damage the herbaceous plant layer. It should

also be noted, the use of an herbicide is an important precursor to facilitate future

prescribed fire, not to replace fire.

After herbicide use restoration plantings of wiregrass should be pursued as

aggressively as possible. Restoration plantings of wiregrass will reintroduce this fire-

dependant species to these sites and further provide an ignition source to carry future

prescribed fires. These plantings will also provide forage for resident gopher tortoises

and other fauna. Depending upon the amount of site preparation needed on individual

sites and costs, refuge management can choose between sowing wiregrass seeds or

planting plugs. Wiregrass seeds do best when sown upon bare soil, while plugs, despite

their higher costs are better suited for a wider array of planting sites.









Following herbicide application and the establishment of wiregrass, prescribed fire

should be returned to these sites. Initially emphasis should be placed on growing season

burns to facilitate the further propagation of wiregrass. Subsequently, burning at various

times of the year under varying conditions will encourage plant diversity without

completely eliminating individual species. A fire frequency of 2 to 4 years would mimic

the natural burning frequency, maintain pine dominance and promote reproduction in

wiregrasses (Tanner et al. 1991).

Pine flatwoods habitats on the refuge exist as slash pine plantations established for

pulpwood production, therefore, these habitats differ markedly from natural flatwoods

habitats. To bolster pulpwood production, timber companies established dense

plantations of slash pine across landscapes. Currently slash pine plantations on the refuge

are planted too densely and should be thinned. Timber thinning opens the canopy, which

promotes herbaceous growth on the forest floor and aids in prescribed burning by

allowing the upward release of heat, the latter especially important for the safe

implementation of growing season bums. As with the high pine habitats, a return to

growing season burns and a varied bum cycle with bums occurring every 2 to 4 years

should be a management goal.

Compartments surveyed for this study contained slash pine basal areas of

approximately 21.75 sq. meters/hectare (94 sq. feet/acre) (pers. comm., D. Barrand,

Chiefland, FL). The refuge's habitat management plan appropriately calls for thinning of

these stands down to 11.5-15 sq. meters/hectare (50-65 sq. feet/acre) during second entry

thinnings (unpubl. Data, USFWS, LSNWR). This study highlights the importance of

timber thinning and the need to restore diversity to areas indiscriminately planted to slash









pine monocultures. Upon thinning, restoration plantings of longleaf pine and wiregrass

should take place.

Within the habitat management plan the refuge is divided into 9 compartments to

facilitate habitat management actions and aid record keeping. Management compartment

boundaries were established along major physiographical and legal features such as

roads, streams and landowner boundaries. Some of these compartments are further

compartmentalized into sub-compartments for more precise land management work.

The current system of compartmentalization should be further delineated since the

current compartments ignore or leave out habitats that were converted to slash pine

stands by the previous timber company management. For example, areas within the

compartments I sampled contained ridges of sandy, Entisol soils which would have

supported either high pine or scrub habitats prior to timber management. Also, within

these compartments, areas of wetter, Aquent soils were bedded and planted through.

Spodosols would be the primary soils dominated by pine flatwoods, as they form

under coniferous trees whose needles are low in base-forming cations and high in acid

resins. These acids bind with iron and aluminum and are carried downward until they

precipitate forming the characteristic spodic horizon (Brady and Weil 2002). A closer

look at each management unit, noting these areas of soil diversity could benefit future

habitat management activities on the refuge. For example, depending on area, higher

ridges could be clear cut of planted slash pine, then restored to high pine or scrub

habitats, while areas of spodosol soils could be maintained as pine flatwoods. Instead of

looking at one block of slash pine timber, this shift in management approach would focus






48


on restoring the natural interspersion of habitats that existed on the area prior to timber

company acquisition, thereby greatly increasing habitat diversity on the refuge.















APPENDIX
ANALYSIS OF VARIANCE TABLES


















Table 12. Analysis of variance results for high pine canopy cover comparisons.


Anova: Single Factor

SUMMARY
Groups Count
High Burrow Density 100
Low Burrow Density 100


ANOVA
Source of Variation
Between Groups
Within Groups


Total


SS
8.359
13.490


df MS F P-value
1 8.359 122.700 1.675E-22
198 0.068


21.850 199


Sum
29.53
70.42


Average
0.295
0.704


Variance
0.047
0.088


F crit
3 888













Table 13. Analysis of variance results for high pine shrub cover comparisons.

Anova: Single Factor

SUMMARY


Groups Count Sum
High Burrow Density 100 26.01
Low Burrow Density 100 106.15


Average
0.260
1.061


Variance
0.042
0.068


ANOVA
Source of Variation
Between Groups
Within Groups


SS
32.112
11.042


df MS
1 32.112
198 0.055


F P-value
575.799 1.629E-60


43.154 199


Total


F crit
3.888














Table 14. Analysis of variance results for high pine woody ground cover comparisons.

Anova: Single Factor

SUMMARY
Groups Count Sum Average Variance
High Burrow Density 100 29.01 0.290 0.057
Low Burrow Density 100 70.77 0.707 0.057


ANOVA
Source of Variation
Between Groups
Within Groups


Total


SS
8.719
11.319


df MS F
1 8.719 152.52
198 0.057


20.039 199


P-value
2.370E-26


F crit
3.888














Table 15. Analysis of variance results for high pine bare mineral soil cover comparisons.

Anova: Single Factor


SUMMARY
Groups Count
High Burrow Density 100
Low Burrow Density 100


Sum
49.32
18.2


Average
0.493
0.182


Variance
0.070
0.043


ANOVA
Source of Variation
Between Groups
Within Groups


Total


SS
4.842
11.328


df MS
1 4.842
198 0.057


F P-value
84.635 5.109E-17


16.170 199


F crit
3.888













Table 16. Analysis of variance results for high pine detritus ground cover comparisons.

Anova: Single Factor


SUMMARY
Groups Count
High Burrow Density 100
Low Burrow Density 100


Sum
33.52
53.3


Average
0.335
0.533


Variance
0.095
0.107


ANOVA
Source of Variation
Between Groups
Within Groups


Total


SS
1.956
20.111


df MS
1 1.956
198 0.101


F P-value
19.259 1.854E-05


22.067 199


F crit
3.888












Table 17. Analysis of variance results for high pine herbaceous ground cover
comparisons.

Anova: Single Factor


SUMMARY
Groups Count Sum
High Burrow Density 100 71.27
Low Burrow Density 100 31.31


Average
0.712
0.313


Variance
0.080
0.056


ANOVA
Source of Variation
Between Groups
Within Groups


Total


SS
7.984
13.541


df
1
198


MS
7.984
0.068


F P-value F crit
116.741 1.089E-21 3.888


21.525 199














Table 18. Analysis of variance results for pine flatwoods canopy cover comparisons.

Anova: Single Factor


SUMMARY
Groups Count
High Burrow Density 100
Low Burrow Density 100


Sum
49.39
79.48


Average
0.493
0.794


Variance
0.135
0.133


ANOVA
Source of Variation
Between Groups
Within Groups


Total


SS
4.527
26.640


df
1
198


MS F P-value
4.527 33.646 2.583E-08
0.134


31.167 199


F crit
3.888














Table 19. Analysis of variance results for pine flatwoods shrub cover comparisons.

Anova: Single Factor


SUMMARY
Groups Count
High Burrow Density 100
Low Burrow Density 100


Sum Average
26.36 0.263
113.48 1.134


ANOVA
Source of Variation
Between Groups
Within Groups


Total


SS df
37.949 1
9.375 198

47.325 199


MS F
37.949 801.44
0.047


Variance
0.061
0.033


P-value
1.564E-71


F crit
3.888














Table 20. Analysis of variance results for pine flatwoods woody ground cover
comparisons.

Anova: Single Factor

SUMMARY


Groups Count
High Burrow Density 100
Low Burrow Density 100


ANOVA
Source of Variation SS
Between Groups 5.996
Within Groups 12.933


Total


Sum
45.52
80.15


df
1
98


Average
0.455
0.801


MS
5.996
0.065


Variance
0.068
0.062


F
91.798


P-value
4.173E-18


F crit
3.888


18.929 199














Table 21. Analysis of variance results for pine flatwoods bare mineral soil cover
comparisons.

Anova: Single Factor

SUMMARY


Groups Count
High Burrow Density 100
Low Burrow Density 100


ANOVA
Source of Variation
Between Groups
Within Groups


Total


Sum
3.29
3


SS
0.00042
0.083


Average
0.032
0.03



df
1 0
198 0


Variance
0.000841
1.822E-18


MS
.000420
.0004205


P-value
0.318


F crit
3.888


0.083 199














Table 22. Analysis of variance results for pine flatwoods detritus ground cover
comparisons.

Anova: Single Factor

SUMMARY


Groups Count
High Burrow Density 100
Low Burrow Density 100


Sum
37.86
53.23


Average
0.378
0.532


Variance
0.095
0.084


ANOVA
Source of Variation
Between Groups
Within Groups


Total


SS
1.181
17.789


df
1
198


MS
1.181
0.089


F P-value
13.147 0.00036624


18.970 199


F crit
3.888














Table 23. Analysis of variance results for pine flatwoods herbaceous ground cover
comparisons.

Anova: Single Factor

SUMMARY


Groups Count
High Burrow Density 100
Low Burrow Density 100


Sum
81.06
40.45


Average
0.810
0.404


Variance
0.165
0.053


ANOVA
Source of Variation
Between Groups
Within Groups


Total


SS
8.245
21.714


df
1
198


MS F
8.245 75.187
0.109


29.960 199


P-value
1.541E-15


F crit
3.888
















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BIOGRAPHICAL SKETCH

Stephen E. Barlow was born on 25 July 1969 in Plant City, Florida. Soon after, his

family moved to Chiefland, Florida, where he was raised; he graduated from Chiefland

High School in May 1987. After serving five years in the U. S. Army and three years in

the U.S. Army Reserve he attended Pittsburg State University in Pittsburg, Kansas, from

which he received his Bachelor of Science degree in biology in December 1997. He

enrolled at the University of Florida in January 1998, majoring in environmental science

with a minor in wildlife ecology, and graduated with the Master of Science degree in

December 2004. During graduate school he began his professional career as a wildlife

technician with the Florida Fish and Wildlife Conservation Commission in January 1999.

He was soon promoted to biological scientist and after 3 years of service with the state of

Florida, he accepted his current position with the U.S. Fish and Wildlife Service as the

wildlife biologist on the Lower Suwannee National Wildlife Refuge. He and Elizabeth

Jane Works, of Humboldt, Kansas, were married on 21 December 1992. They have one

son, Seth Douglas Barlow, born in Gainesville, Florida, on 21 August 2001.