This file was created by scanning the printed publication. Errors identified by the software have been corrected; however, some errors may remain. Management of Amphibians, Reptiles, and Small Mammals in Xeric Pinelands of Peninsular Florida1 I. Jack S t ~ u tDonald ,~ R. Ri~hardson,~ and Richard E. Roberts4 Xeric pinelands seem incongruent with reference to Florida, a state with annual rainfall that ranges from 50-65 in (19.6-25.6 cm). None theless, the Florida peninsula contains thousands of acres of sandy soil derived from marine deposits dating to the Pleistocene (White 1970).Two distinct plant associations, longleaf pine (Pinus palustris)/ turkey oak (Quercus laevis) sandhill and sand pine scrub (Pinus clausa), have developed on these nutrient deficient and excessively welldrained soils. Significant areas of these plant associations occur at higher, albeit modest, elevations relative to the surrounding landscape. In fact, certain topographic features, e.g., the Lake Wales Ridge and the Marion Upland, were likely to have been true islands during interglacial periods while the remainder of Florida was covered by a shallow sea. Regardless of their exact origin, xeric pinelands support many relatively unusual species of amphibians, reptiles, and small mammals. 'Paper presented at symposium, Management of Amphibians, Reptiles, and Small Mammals in North America. (Flagstaff, AZ. July 79-21, 1988). 21. Jack Stout is Professor of Biological Sciences, Department of Biological Sciences, University of Central Florida, Box 250CO, Orlando, FL 328 16-0368. 3DonaldR. Richardson is Adjunct Professor, Department of Biology, University of South Florida, Tampa, FL 33620. 4RichardE. Roberfs is Biologist, Division of Recreation and Parks, Florida Department of Natural Resources, Hobe Sound, FL 33455. Abstract.-The primary xeric pinelands of peninsular Florida are lon~leafpinelturkey oak sandhills and sand pine scrub, Their management on public lands is largely confined to prescribed burning to maintain fire climax status of the vegetation. The regulation of large-scale developments on private land has stim ulated interest in preserve design and management. The suite of techniques used to solve conflict between natural system preservation and development includes: (I) conservation set asides (preserves) on site; (2) habitat restoration; (3) purchase and dedication of off-site preserves; (4) species relocation; and (5) wildlife resource mitigation fund. Human population growth (3.3% per year) and development in Florida continues to encroach on upland habitats and particularly on xeric pinelands. Most of the habitat loss is to agricultural uses, principally citrus. Oddly, the state's excellent wetlands protection acts have forced development into the uplands. Thus, xeric pinelands and their narrowly adapted fauna and flora are increasingly threatened by area reduction, fragmentation and isolation. It is our intent to discuss the management of these xeric pinelands in general and, more specifically, in the context of small preserves in an otherwise developed landscape. Management of xeric pinelands as ecosystems is yet in its infancy, and more detailed prescriptions for designated species are unproven. However, progress is being made (Cox et al. 1987) and improvement and revision of current thinking on management practices is anticipated. This paper summarizes selected literature on xeric pineland and the species associated with these communities to assess management practices. In addition, unpublished information has been used and identified in the text. Preserve design efforts by us have been on behalf of developers responding to development orders prepared by governmental agencies. These designs are site specific in detail, but nonetheless point to general problems and solutions. Our approach has been to focus on provid- ing the area required to support minimum viable populations of "keystone" or otherwise critical animal species of a given xeric pineland. Once this area is settled on, management should focus on those species whose minimum area requirements are met, whereas no special efforts are expended on species with larger area requirements. XERIC PINELAND HABITATS Longleaf Pinepurkey Oak Sandhills The longleaf pine/ turkey oak sandhill association (LLP/TO) was about 15% (2,110,256 ha) of the natural landscape of peninsular Florida in pre-Columbian times (fig. 1) (Auffenberg and Franz 1982).This xeric pineland occupies rolling topography of several ridge systems that run north-south, notably Trail Ridge, the Lake Wales Ridge, and the Brooksville Ridge; numerous lesser ridges and hills are identified by White (1970). These ridges consist of deep, well-drained soils of the Lakeland, Eustis, and Blanton associations (Beckenbach and Hammett 1962).Laessle (1942,1958a) describes the LLP/TO plant association as a fire climax system dominated by longleaf pine; slash pine (Pinus elliotti) replaces longleaf pine in the community in south Florida. Turkey oak is a minor tree, but can achieve co-dominance when fires are suppressed. The predominant understory plant is wiregrass, Aristida stricta; however, a rich assemblage of perennial herbs vary in prominence in concert with seasonal changes. Monk (1968) recognizes two additional phases of sandhill vegetation in north central Florida: (I) longleaf pine/sand post oak (Quercus nrargaretta) and (2) longleaf pine/southern red oak (Quercus falcata). A fourth phase, longleaf pine/scrub hickory (Cay a floridam), occurs in the southern portion of the Lake Wales Ridge (Abrahamson et al. 1984).Veno (1976), Givens et al. (1984), and Abrahamson et al. (1984) provide quantitative data on LLP/TO community structure and dynamics. Myers (1985)suggests that longleaf pine/ turkey oak and sand pine scrub associations are successionally linked in some portions of their geographic ranges. Differences in physical / chemical features of soils of LLP/TO and SPS communities in the Ocala National Forest are not considered to be sufficient to explain the local dis- Figure 1 .-Potential geographic distribution of longleaf pine/turkey oak sandhill and sand pine scrub xeric pinelands in Florida. Light shading indicates the sandhills and darker shading indicates the scrub. These distributions are based on Davis (1 980) and do not reflect minor sites of either community due to the scale of the itlustration. tribu tion of the communities (Kalisz and Stone 1984). Prior to settlement by European man, ground fires occurred in LLP/ TO sandhills at intervals of 1-5 years. These relatively "cool" fires favor regeneration of longleaf pine, flowering by grasses and herbs, and suppress growth of woody plants (Myers 1985). include those of Abrahamson et al. (1984) and Latham (1985). Outstanding examples of SPS include the "Big Scrub," part of the Ocala National Forest, scrubs of the Lake Wales Ridge, eg., the Archbold Biological Station, and stands along the Atlantic Coastal Ridge. SMALL VERTEBRATE SPECIES ASSEMBLAGES Sand Pine Scrub Compared with the LLP/TO sandhill association, sand pine scrub (SPS) has less area (250,000 ha) and a far more limited distribution (fig. 1). Scrub is associated with old shorelines, lake margins, and stream courses where extremely well washed, nutrient deficient sands were deposited during Pleistocene times (Kurz 1942; Laessle 1958a, b, 1967).The most widespread soils supporting SPS are the St. Lucie, Lakewood, and Pomello associations (Beckenbach and Hammett 1962). Sand pine scrub is a two-layered community. Sand pine (Pinus clausa) normally occurs as a relatively evenaged overstory species. The understory is comprised of 10-20 species of evergreen shrubs 1-5m in height. Four species of oaks comprise the bulk of the biomass, Quercus geminata, Q. myrtifolia, Q. chapmanii, and Q. inapina. Lesser numbers of other species including Ceratiola ericoides, Lyonia ferruginea, and Osmanthus americanus add to local diversity. Sand pine scrub is a fire climax community (Laessle 1958a, Abrahamson et al. 1984). In contrast with LLP/TO, SPS burns at intervals of 20-70 years; a combination of ground and crown fires destroys all the above-ground vegetation. Most of the woody plants, with the notable exception of the sand pine and Ceratiala, readily sprout from root crowns following fires. Laessle (1958a), Veno (1976), and Richardson (1977) provide data on plant community structure of scrubs. Recent quantitative studies Longleaf Pinepurkey Oak Sandhills Amphibians and Reptiles At least 47 species of herptiles, including 2 newts, 13 toads and frogs, 3 turtles, 10 lizards, 1 amphisbaenian, and 18 snakes, are reported to occur in LLP/TO habitats (table 1). Campbell and Christrnan (1982) list 5 categories of reptile and amphibian species that occur in LLP/TO and SPS: (1) characteristic (18 species); (2) associated with tortoise burrows (3 species); (3) frequent (8 species); (4) occasional (14 species); and (5) associated with aquatic habitats (21 species). Of the characteristic species, 7 are regarded as adapted to xeric conditions, 3 as sand swimmers, viz., Neoseps reynoldsi; Eumeces egregius, and Tantilla relicta, and the remainder (Sceloporus woodi, Masticophis flagellum, Stilosoma extenuaturn, Cnemidophorus sexlineatus) to other physical features of the habitats. The gopher tortoise (Gopherus polyphemus) is a terrestrial turtle that digs deep burrows in the welldrained sandhill soils (Auffenberg and Franz 1982).Stout (1981)and Eisenberg (1983) recognized the gopher tortoise was the keystone species in xeric pinelands. Some 80 species of animals may be classified as burrow commensals (Cox et al. 1987); however, the number of obligatory commensals is much smaller. Herptiles particularly associated with gopher tortoise burrows include Rana areolata, Pituophis melanoleucus, and D ymarchon corais. The snake fauna of LLP/TO sandhills is species rich (L 18 species). This diversity includes large forms, e.g., Dymarchon corais couperi and Crotalus adamanteus, and small, specialized species like Stilosoma extenuatum. This latter ophiophagous species feeds largely on Tantilla relicta; Tantilla, is in turn specialized on Tenebrionidae larvae (Mushinsky 1984). Small Mammals At least 19 species of small mammals with body masses less than 6.0 kg may be anticipated in LLP/TO sandhills (table 2). Two are fossorial, Scalopus aquatious and Geomys pinetia, 1 semi-fossorial, P. polionotus, and 2 occur in the surface litter, Blarina carolinensis and Cyptotis parua. Arboreal species include Sciurus carolinensis, S. niger, Glaucomys volans, P. gossypinus, and Ochrotomys nuttalli. Podomys floridanus nests in the burrows of the gopher tortoise and the pocket gopher (Layne 1969); it may enlarge other openings in the soil to establish burrows independently of the gopher tortoise (R. E. Roberts, personal observation). Dasypus novemcinctus is the only exotic species of mammal that is clearly established in the sandhill communi ty. Sand Pine Scrub Amphibians and Reptiles Campbell and Christman (1982) listed 64 species of reptiles and amphibians that may be found in LLP/ TO sandhills and SPS. Pitfall trapping in six different even-aged stands of SPS on the Ocala National Forest by Christman et al. (unpublished manuscript and personal communication) revealed 27 species (table 1). Of 1,624 individuals trapped, the common species were Bufo terrestris (n=332),Cnenridophorus sexlineatus (n=329), and Sceloporus woodi (n=216);five species were represented by single captures. Christman et al. concluded that the herpetofaunal diversity declined with increasing age of SPS stands. Copherus polyphemus is the keystone species in SPS but is less common there than in LLP/TO (Auffenberg and Franz 1982).Many, if not most, of the burrow commensals are common in SPS (Cox et al. 1987). Podomys floridanus is an example. Small Mammals Fourteen species of small mammals commonly inhabit SPS (table 2). Pod- omys floridanus is a predictable member of the assemblage throughout the range of scrubs in peninsular Florida (Layne 1978).Three subspecies of Peromyscus polionotus occur in scrubs of the interior and east coast portions of the peninsula. Common small mammals in central peninsular Florida scrubs include Podomys floridanus, Peromyscus goss ypinus, Ochrotomys nuttalli, and Glaucomys volans (Swindell 1987).Podomys floridanus is the predominate small mammal in scrubs of southeast Florida (Richardson et al. 1986). Limited data suggest Spilogale putorius is a major predator on small mammals in scrubs with lesser roles played by Mephitis mephitis and Mustela frenata (Stout and Roberts, personal observations). ENDANGERED AND THREATENED SPECIES Ten species of amphibians, reptiles, and small mammals associated with xeric pineland are currently listed as having some level of threatened, endangered, or sensitive status by either the state of Florida or the Department of Interior (table 3). The extensive overlap in species composition between the two pineland communities results from the high number of species common to both types. The Endangered Species Act charges federal agencies with the responsibility to manage federally listed species on federally owned lands. At the state level, preservation of these listed species is of major concern when they occur on parcels of land scheduled for large-scale development. Preserve design and management practices for these species have largely evolved on an ad hoc basis without adequate time for an evaluation of the management or the longterm implications for the species. MANAGEMENT OF XERIC PINELANDS ON PUBLIC LANDS Of three national forests in Florida, only the Ocala National Forest is located in the peninsula. It totals 153,846 ha of which 85,020 ha are SPS and 18,219 ha LLP/TO. The National Forest Management Act (1976) and pursuant regulations (36 CFR 219) require that each forest be managed to maintain well-distributed and viable populations of wildlife species, including species that are endangered or threatened (Norse et al. 1986). Silvicultural systems differ between the two pineland communities. On the Ocala National Forest sand pine scrub is routinely harvested in patchy clearcuts that range from 16-24 ha in area. Scrub understory vegetation is allowed to regenerate naturally; however, sand pine is seeded following site preparation 1 by a single roller chopping. The harvest rotation length is about 50 years. In contrast, LLP/TO is ostensibly managed on a 80-100 year rotation and shelterwood cutting favors natural regeneration of the longleaf pine (Don Bethancourt, personal communication). In practice, harvesting of longleaf pine may occur in 60 years. Effectiveness of ecosystem management in the SPS community will be judged by the response of designated indicator species, such as gopher tortoises and scrub jays Uphelocoma coerulescens) (table 3). The gopher tortoise is also a designated indicator species for the LEP/TO community. The significance of the gopher tortoise as a keystone species was emphasized in 1986 when harvesting of the species on national forests in Florida was made illegal through an agreement between the U.S. Forest Service and the Florida Game and Fresh Water Fish Csmmission. Other species-specific management practices involving amphibians, reptiles, or small mammals have not been deemed necessary to carry out on the Ocala National Forest (Don Bethancourt, personal communication). In fact, the impact of timber harvesting on small vertebrates of LLP/TO and SPS communities is simply not known. Public lands in Florida supporting xeric pinelands include, hut are not limited to, state forests and state parks. State forests with large acreages of LLP/TO, e.g., the Withlacoochee State Forest, are managed at the ecosystem level. Prescribed burning is done every 3-8 years and future timber sales will follow a rotation length of 80-120 years; currently rotation lengths are about 60 years and are not regarded as favorably for endemic wildlife. Wildlife management areas overlap the state forest holdings and are managed for sustained yields of wildlife by the Florida Game and Fresh h7aterFish Commission based on a memorandum of understanding between agencies (Cathy Ryan, personal communication). State parks are managed by the Division of Recreation and Parks of the Florida Department sf Natural Resources (FDNR). An ecosystem approach is taken in the restoration and management of xeric pinelands on state park lands (Jim Stevenson, personal communication). Prescribed burning has been used since 1969 to control hardwood invasion of LLP/ TO stands and to stimulate growth and flowering of grasses and herbs. Burning in spring and early summer appears to best duplicate the historic timing of lightning initiated fires in xeric pinelands. The impact of these management practices on the plant community has been documented (Davis 1984); the response of reptiles, amphibians, and small mammals is currently under study (Stout et al. unpublished). Generally, mature stands of SPS have not been burned until recently, due to the unpredictable behavior of fire in the community; however, a prescription for burning this fuel type has been written and tested on private land and state parks (Doran et al. 1987).Early recovery stages of SPS appear to support the greatest diversity of reptiles and amphibians. However, as canopy closure occurs in SPS, ground cover diminishes and habitat quality for gopher tortoises declines (Cox et al. 1987). In contrast, similar numbers of Podomys have been observed in early (R. E. Roberts, unpublished data, J. Dickinson State Park); intermediate (Stout 1982); and old growth SPS (James N. Layne, unpublished data, Archbold Biological Station). State parks, reserves, and preserves appear to be ideal lands to explore species-specific management measures for herptiles and small mammals. For example, sand swimming herptiles (Smith 1982)require openings that are relatively root free in LLP/TO and SPS habitats. The natural occurrence of such openings may have been due to "hot" spots associated with the combustion of high fuel loads, e.g., fallen trees (Ron Myers, personal communication). Concentration of natural fuels prior to prescription burns in SPS would offer a means to create microhabitat conditions favorable for the sand swimmers. MANAGEMENT OF XERIC PINELAND ON PRIVATE LAND Development of Regional Impact Concern with management of amphibians, reptiles, and small mammals on private lands in Florida derives from state and federal protection of endangered species and the development guidelines promulgated during the Development of Regional Impact (DRI) process. "The Florida Environmental Land and Water Management Act of 1972" (Chapter 380, Florida Statutes) defines developments of regional impact in Section 380.06(1),Florida Statutes, as "...any development which, because of its character, magnitude, or location, would have a substantial effect upon the health, safety, or welfare of citizens of more than one county (Anonymous 1976)." Large scale development projects in peninsular Florida commonly involve hundreds to several thousand acres of relatively natural landscape. The DRI process requires bona fide studies of wildlife populations and their associated habitats; emphasis is placed on listed species. Developers must prepare viable management strategies to accommodate wildlife resources dependent upon their lands (Cox et al. 1987; Richardson et al. 1986). Management strategies of developers with xeric pinelands generally follow one of two somewhat overlapping approaches to preserve habitat and/or species values: (1) conservation set asides or (2) mitigation. Conservation set asides are, in principle, the preferred solution. In practice some habitat is dedicated in perpetuity as a nature preserve; preserve design currently is a somewhat ad hoc process and will be discussed more completely in a subsequent section of this paper. Very high land values may dictate mitigation rather than on site preservation of habitat. Mitigation may take many forms to compensate for development of xeric pinelands. Restoration of degraded land (Humphrey et al. 19851, not necessarily xeric pinelands, is one method. Another tactic is to purchase comparable land or some other type of land of equivalent natural value elsewhere and dedicate it to preservation. A formal process for accomplishing this option is presently under study by the Florida Game and Fresh Water Fish Commission. Preservation of habitat is the basic purpose of conservation set asides and mitigations. The value of these efforts depends on the proximity to larger, undeveloped tracts of land, travel corridors, area of preserves, and future management options. Another form of mitigation is the relocation of sensitive species from tracts of land to be developed to land dedicated to purposes that are consistent with the long-term survival of the relocated species. In Florida, the gopher tortoise has been the focus of numerous relocation efforts. Diemer (1984) discussed the advantages and disadvantages of relocation of gopher tortoises as a species management strategy. Formal research on gopher tortoise relocation was recently reported (Proced. Gopher Tortoise Relocation Symp., 27 June 1987, Gainesville, FL, in press). The Florida Game and Fresh Water Fish Commission regulates relocations by a permit system based on a standardized relocation protocol. Preserve Design Preserve design is an evolving and controversial area of conservation biology (Diamond 1975,1978; Gilbert 1980; Higgs 1981; Margules 1982; Pickett and Thompson 1978; Pyle 1980; Soule and Simberloff 1986). Large preserves encompassing a mo- saic of xeric pinelands, mesic forests, and seasonal and permanent wetlands would perhaps offer the ideal landscape unit for long-term preservation of amphibians, reptiles, and small mammals in peninsular Florida. Because preserves on private lands must be justified and dedicated through the DRI process, economics dictates preserve units of minimal size. Rarely do we have the opportunity to cluster or juxtapose these small units to take advantage of the so called "rescue effect" (Brown and Kodric-Brown 1977). In practice, conservation set asides tend not only to be small in acreage but also only of one habitat type. The latter presents a dilemma for species whose requirements often include two or more contrasting habitats. For example, the gopher frog lives in tortoise burrows in LLP/TO sandhills during late spring, summer and early fall and migrates to temporary wet season depressions to breed in winter and early spring (Moler and Franz 1987).Thus a mosaic of upland-wetland habitats in close proximity are essential to maintain viable populations of this species. Other species such as the indigo snake have home range requirements that include 122-202 ha of several uplandwetland habitat types (Moler 1985; Moler unpublished data). It is obvious that large landscape units are necessary to preserve viable populations of these animals. We have prepared a detailed preserve design for a SPS community within the city of Boca Raton, Florida (Richardson et al. 1986; Stout et al. 1987; manuscript in preparation). The approach taken anticipated Soule and Simberloff (1986) and recommended the area of the preserve be sufficient to support a minimum viable population (Franklin 1980)of gopher tortoises because of their status as the keystone species. Although biologically reasonable, this basis-for determining preserve size is often economically unrealistic from the view point of the private land- owner. A consortium of public landowners would, however, permit the purchase and long-term management of the preserve as recommended. Cox et al. (1987) offer guidelines for the design of preserves on private lands to maintain gopher tortoise populations. They employed the computer simulation model POPDYN (Perez-Trejo and Samson manuscript) to determine population viability based on different initial sizes. Populations of 40-50 individuals were found to be likely (290%)to persist 200 years. Based on existing literature on home range requirements, Cox et al. (1987) recommended a minimum preserve of 10- 20 ha, depending on habitat quality, to support 40-50 tortoises. Another approach to determining the area of a preserve employs "incidence functions" (Diamond 1978). Incidence functions are species specific and derived from data sets which reveal the fraction of plots (discrete habitats) of different areas that actually support the species. It is a matter of judgement as to the probability of occurrence, e.g. 0.5 as opposed to 0.7, that would set a lower limit to area for an acceptable preserve. Data sets useful for evaluating this approach with respect to amphibians, reptiles, and small mammals in xeric pinelands are pres- ently lacking. Table 4 provides data we have gathered on area of discrete habitats and the presence or absence of gopher tortoises and Florida mice. It is apparent that tortoises are less area sensitive than Florida mice and that Florida mice are patchy in occurrence in LLP/TO, perhaps only secondarily related to area. Incidence functions do not necessarily reveal the minimum area required to support minimum viable populations (Franklin 1980).We believe preserve area should be based on providing this requirement, particularly when preserves are isolated relative to average dispersal distances of keystone species. However, clusters of preserves within dispersal distances of keystone species may be of less area per preserve due to a high likelihood of reinvasion from nearby populations following local population extirpations (Noss and Harris 1986). Management of Preserves in Xeric Pinelands Cedar Grove J. Dickinson 21.5 256.2 + + + + The future viability of preserves depends largely on their ownership after development of the surrounding landscape. It is unlikely that homeowners associations will assume the cost of management if preserves remain as a part of the overall development's "commons." Public ownership is an alternative and might rest with a city, county, or state. Local governments seem more appropriate; however, funds and expertise to manage may be lacking. One preserve in south Florida is designed to border a city park, thus allowing its maintenance and /or management costs to be assumed over time as part of the existing park system (Richardson, personal observation). Regardless of the ownership, a commi tment to long-term management must be achieved if a preserve is to retain natural values. Management options for nature preserves range from a decision I ) to d o nothing and let nature take its course; 2) to manage for maintenance of a viable ecosystem, which implies the natural biota, including amphibians, reptiles, and small mammals, will be present in proportion to their normal abundance; or 3) to focus management on the needs of one or more species. White and Bratton (1980) have exposed the folly of the first management option. The decision to emphasize ecosystem or species management depends on the entity responsible for management, type of preserve, management objectives, area of the preserve, nature of the surrounding lands, relative overall or regional rarity of particular species, and the resources available for management. Management objectives of any preserve should focus on: I) maintenance of normal ecosystem processes; 2) conservation of soil; 3) maintenance or restoration of normal hydrologic conditions; 4) prevention of establishment of exotic species.; 5) and prevention of human encroachment (e.g., dumping, ATVs, etc.) Beyond these generalities, management of preserves is an idiosyncratic process that may concern endemic species, genetics of inbred populations, or restoration of periodic wild fires. Xeric pinelands of peninsular Florida depend on periodic fires to maintain their structure and function (Laessle 1958a; Abrahamson 1984). Thus a burning program is essential in the management of LLP/TO or SPS preserves. Spring or early summer prescribed burns are routinely used to maintain LLP/TO communities on state parks. Doran et al. (1987) have documented prescribed burns of SPS preserves in an urban setting based on rather esoteric fire models developed by the U.S. Forest Service. ~ o ~ htortoises e r respond favorably to the bums (Stout et al. 1988).A mosaic of recovery stages in SPS may favor beta diversity of herptiles and small mammals. Mushinsky (1985) has carefully documented the response of the herpetofauna to a vari- ety of burning schedules in LLP/TO. Diversity and abundance of amphibians and reptiles was increased on experimental plots relative to unburned controls. Re-establishment of the pine overstory may be necessary to produce needle cast for carrying fire (Landers and Speake 1980). Management of conservation set a s i d s and/or easements may focus on particular species or combinations of species. The smaller the preserve the more likely that a reduced suite of species will be present (Richardson et al. 1986).Given that a fixed area is available for management, major efforts to enhance or maintain habitat should target those species that can maintain viable gopula tions within the preserve (Shaffer 1986). A species whose minimum area requirements for a minimum viable population exceeds the preserve area should not be of major concern (Shaffer and Samson 1985); nonetheless, such species can benefit from the preserves if travel corridors exist (Harris 1984). DISCUSSION Xeric pinelands of peninsular Florida support a species-rich assemblage of reptiles, amphibians, and small mammals. Growth and development continues to diminish LLP/TO and SPS habitats to the detriment of the associated biota. Land in public ownership, e.g. state parks and forests, national forests, and private holdings, e.g., the Archbold Biological Station, and institutional lands such as the Ordway and Swisher Preserves, jointly owned and managed by the University of Florida and The Nature Conservancy, will be increasingly valuable as other xeric pinelands are converted to land uses not favorable to the biota. Thus, management of these xeric pinelands will become more important in the future. At present management is largely limited to prescribed bums to maintain what were historically fire climax communities. Thus, fire management is tantamount to small vertebrate management. In the future as air quality standards are modified, prescribed burning, particularly in or near urbanized areas, will be restricted or eliminated as a management option. Alternative means of habitat manipulation need to be developed, particularly for SPS. Basic information on the life history of many amphibians, reptiles, and small mammals of xeric pinelands is lacking. The Nongarne Wildlife Program of the Florida Game and Fresh Water Fish Commission has initiated and f u ~ ~ d rather ed large scale studies of SPS and LLP/TO communities. These studies are at the community level and largely observational. Management needs of individual species may be derived only secondarily from this research. Studies that focus on particular species will ultimately lead to more refined habitat management guidelines. The report by Cox et al. (1987) will likely serve as a model for the preparation of habitat protection guidelines; management follows protection (White and Bratton 1980). Management alternatives at the ecosystem and species level are needed now for xeric pinelands on private lands undergoing development Regulation of development in these habitats as currently practiced will result in a patchwork of small, isolated nature preserves. Preservation of natural habitat in a developed landscape is, of course, desirable. However, several problems remain: (1) who will own the preserves, (2) how will a management plan be prepared, and (3) who will be responsible for management? Even another decade of rapid growth in peninsular Florida may result in a feG hundred nature preserves, which will not necessarily be restricted to xeric pineland habitat. Ignoring the question of ownership, no public land management agency is currently capable of assuming the charge of managing these preserves. Lack of manage- ment, e.g., failure to conduct prescribed burning, will allow successional changes to occur to the detriment of many small vertebrates narrowly adapted to xeric pinelands. Loss of habitat and species values originally used by jurisdictional agencies to secure preserve set asides provides a potential basis for private land owners to request development rights on the land. This action would defeat the entire purpose of having conservation set asides. An alternative to on site habitat protection is offered by Cox et al. (1987) in regard to preserving habitat for the gopher tortoise. The alternative, a Wildlife Resource Mitigation Fund (WRMF),allows a developer to contribute money to the fund to mitigate losses of valuable wildlife habitat on lands being developed. The collective monies of several development projects would allow an independent group such as the Trust For Public Lands to assist in the purchase of commensurate lands to expand an existing public park, preserve or forest. Management is more likely to be applied to these lands and ultimately the resources are better served by the public agencies. ACKNOWLEDGMENTS We thank the authors of the papers cited herein for their efforts and dedication to science. Biologists who contributed to our knowledge of xeric pineland include but are not limited to the following individuals: Dan Austin, Don Bethancourt, Russ Burke, Steve Christman, David Cook, David Corey, Jim Cox, Joan Diemer, Dick Franz, Larry Harris, Randy Kautz, Jim Layne, Wayne Marion, Paul Moler, Ron Myers, Reed Noss, Cathy Ryan, Jim Stevenson, and Don Wood. Support for research on the ecology of sandhill communities was provided by the Nongame Wildlife Program, RFP86-003, of the Florida Game and Fresh Water Fish Commission. Development groups that funded work by the authors on xeric pinelands include Hardy-Lieb Development Corporation, The Adler Group, and Deutsch-Ireland Properties. The Division of Recreation and Parks (FDNR), Department of Biology, University of South Florida, and the Department of Biological Sciences, University of Central Florida assisted in our studies in a variety of ways. We thank Beverly Bamekow, Rita Greenwell, Barbara Erwin, and Nancy Small for typing the manuscript. Lastly, we thank Paul E. Moler, James N. Layne and Robert C. Szaro for providing excellent suggestions to improve the paper. LITERATURE CITED Abrahamson, Warren G. 1984. Postfire recovery of Florida Lake Wales ridge vegetation. American Journal of Botany 7l:9-21. Abrahamson, Warren G., Ann F. Johnson, James N. Layne, and Patricia A. Peroni. 1984. Vegetation of the Archbold Biological Station, Florida: an example of the Southern Lake Wales Ridge. Florida Scientist 47:209-250. Anonymous. 1976. Developments of regional impact: guidebook for preparation of the application for development approval. 65 p. Division of State Planning, Department of Administration, Tallahassee, Florida. Arata, Andrew A. 1959. Effects of burning on vegetation and rodent populations in a longleaf pine turkey oak association in north Central Florida. Quarterly Journal Florida Academy Sciences 22:94104. Auffenberg, Walter, and Richard Franz. 1982. The status and distribution of the gopher tortoise (Gopherus polyphemus). p. 95-126. In North American tortoises: conservation and ecology. R. Bruce Bury, editor. Wildlife Research Report 12. 126 p. U.S. Fish and Wildlife Service, Washington, D.C. Beckenbach, J. R., and J. W. Hammett. 1962. General soil map of Florida. 1p. Florida Agricultural Experiment Stations and the Soil Conservation Service. Brown, James H., and Astrid KodricBrown. 1977. Turnover rates in insular biogeography: effects of immigration on extinction. Ecology 58:445-449. Campbell, Howard W., and Steven P. Christman. 1982. The herpetological components of Florida sandhill and sand pine scrub associations. p. 163-171. In Herpetological communities: a symposium of the Society for the Study of Amphibians and Reptiles and the Herpetologists' League, August 1977. Norman J. Scott, Jr., editor. 239 p. U.S. Fish and Wildlife Service, Wildlife Research Report 13. Christrnan, Steven P., Howard I. Kochman, Howard W. Campbell, Charles R. Smith, and Wallace S. Lippincott, Jr. Successional changes in community structures: amphibians and reptiles in Florida sand pine scrub. Unpublished manuscript. Cox, James, Douglas Inkley, and Randy Kautz, 1987. Ecology and habitat protection needs of gopher tortoise (Gopherus polyphemus) populations found on lands slated for large-scale development in Florida. Florida Game and Fresh Water Fish Commission Nongame Wildlife Program Technical Report no. 4.75 p. Tallahassee, FL. Davis, John H. 1980. General map of natural vegetation of Florida. Agric. Exp. Sta. Institute of Food and Agricultural Sciences Circular S-178.1 p. University of Florida, Gainesville. Davis, Mary M. 1984. Phenology of flowering plants on experimental fire plots, St. Marks National Wildlife Refuge. M. S. Thesis. 54 p. Florida State University. Tallahassee, FL. Diamond, Jared M. 1975. The island dilemma: lessons of modern biogeographic studies for the design of natural reserves. Biological Conservation 7:129-146. Diamond, J. M. 1978. Critical areas for maintaining viable populations of species. p. 2740. In The breakdown and restoration of ecosysHoldgate and M. J. tems. M. b7. Woodman, editors. Plenum Press, New York, N. Y. Diemer, Joan E. 1984. Tortoise relocation in Florida: solution or problem? Proc. 1984 Symposium Desert Tortoise Council: 131-135. Doran, R. F., Donald R. Richardson, and Richard E. Roberts. 1987. Prescribed burning of the sand pine scrub community: Yamato Scrub, a test case. Florida Scientist 50384192. Eisenberg, J. F. 1983. The gopher tortoise as a keystone species. p. 1-4. In The gopher tortoise: a keystone species. R. J. Bryant and R. Franz, editors. Proceedings of the 4th Annual Meeting of the Gopher Tortoise Council. Florida State Museum, Gainesville, FL. Franklin, Ian Robert. 1980. Evolutionary change in small populations. p. 135-149. In Conservation biology: an evolu tionary-ecological perspective. Michael E. Soule and Bruce A. Wilcox, editors. 395 p. Sinauer Assoc., Sunderland, MA. Gilbert, F. S. 1980. The equilibrium theory of island biogeography: fact or fiction? Journal of Biogeography 2209-235. Givens, Kerry T., James N. Layne, Warren G. Abrahamson, and Susan C. White-Schuler. 1984. Structural changes and successional relationships of five Florida Lake Wales Ridge plant communities. Bulletin Torrey Botanical Club 111%-18. Harris, Larry D. 1984. The fragmented forest. 211 p. The University of Chicago Press, Chicago,,IL. Higgs, A. J. 1981. Island biogeography theory and nature reserve design. Journal of Biogeography 8:117-124. Humphrey, Stephen R., John F. Eisenberg, and Richard Franz. 1985. Possibilities for restoring wildlife of a longleaf pine savanna in an abandoned citrus grove. Wildlife Society Bulletin 13:487496. Kalisz, Paul J., and Earl L. Stone. 1984. The longleaf pine islands of the Ocala National Forest, Florida: a soil study. Ecology 65:1743-1754. Kurz, Herman. 1942. Florida dunes and scrub vegetation and geology. Florida Geological Survey Bulletin 23:15-154. Laessle, Albert M. 1942. The plant communities of the Welaka Area. University of Florida Publication, Biological Science Series 4:l-143. Laessle, Albert M. 1958a. The origin and successional relationship of sandhill vegetation and sand pine scrub. Ecological Monographs 28:361-387. Laessle, Albert M. 1958b. A report on succession studies of selected plant communities on the Univ. of Ha. conservation reserve, Welaka, Florida. Quarterly Journal Florida Academy Science 21:101-112. Laessle, A. M. 1967. Relationship of sand pine scrub to former shore lines. Quarterly Journal Florida Academy Science 30:269-286. Landers, J. Larry, and Dan W. Speake. 1980. Management needs of sandhill reptiles in Southern Georgia. Annual conference Southeastern Association Fish and Wildlife Agencies. 34:515-529. Latham, Pamela J. 1985. Structural comparisons of sand pine scrubs of east-central Florida. M. S. Thesis. 106 p. University of Central Florida, Orlando. Layne, James N. 1969. Nest-building behavior in three species of deer mice, Perornyscus. Behaviour 35288-303. Layne, James N. 1978. Pevomyscus flon'danus (Chapman). p. 21-22. In Rare and endangered biota of FLorida. Vol. 1: mammals. James N. Layne, editor. 52 p. University Presses of Florida, Gainesville, Fl. Margules, C., A. J. Higgs, and R. W. Rafe. 1982. Modern biogeographic theory: are there any lessons for nature reserve design? Biological Conservation 24:ll5-128. Moler, Paul E. 1985. Distribution of the eastern indigo snake, D ymarchon corais couperi, in Florida. Herpetological Review 116:37-38. Moler, Paul E., and Richard Franz. 1987. Wildlife values of small, isolated wetlands in the southeastern coastal plain. p. 234-241. In Proc. 3rd S. E. Nongame and Endangered Wildlife Symposium. R. R. Odom, K. A. Riddleberger and J. C. Ozier, editors. Georgia Department Natural Resources, Game and Fish Division. Monk, Carl D. 1968. Successional and environmental relationships of the forest vegetation of North Central Florida. The American Midland Naturalist 79:441457. Mushinsky, Henry R. 1984. Fire and the Florida sand hill herpe tofaunal community: with special attenti on to responses of Cnemidophorus sexlinea tus. Herpetologica 41:333342. Myers, Ronald L. 3985. Fire and the dynamic relationship between Florida sandhill and sand pine scrub vegetation. Bulletin Torrey Botanical Club 122241-252. Norse, Elliott A., Kenneth L. Rosenbaum, David S. Wilcove, Bruce A. Wilcox, William H. Romme, David W. Johnston, and Martha L. Stout. 1986. Conserving biological diversity in our national forests. 116 p. The Wilderness Society. Noss, Reed F., and Larry D. Harris. 1986. Nodes, networks, and MUMS: preserving diversity at all scales. Environmental Management 10:299-309. Pickett, S. T. A., and John N. Thompson. 1978. Patch dynamics and the design of nature reserves. Biological Conservation 13:27-37. Pyle, Robert Michael. 1980. Management of nature reserves. p. 319327. In Conservation Biology: An evolu tionary-ecological perspective. Michael E. Soule and Bruce A. Wilcox, editors. 395 p. Sinauer Associates, Sunderland, Mass. Richardson, Donald Robert. 1977. Vegetation of the Atlantic coastal ridge of Palm Beach County, Florida. Florida Scientist 40:281-330. Richardson, Donald R., I. Jack Stout, Richard E. Roberts, Daniel F. Austin, and Taylor R. Alexander. 1986. Design and management recommendations for a sand pine scrub preserve: the Yamato scrub. 124 p. Ecological Consultants, Tampa, Florida. Shaffer, Mark L. 1986. The metapopulation and species conservation: the special case of the northern spotted owl. p. 86-99. In Ecology and management of the spotted owl in the Pacific Northwest, Ralph J. Gutierrez, and Andrew 8. Carey, tech editors. USDA Forest Service. General Technical Report PNW-185,119 p. Pacific Northwest Forest and Range Experiment Station, Portland, Oregon. Shaffer, Mark L., and F. B. Samson. 1985. Population size and extinction: a note on determining critical population sizes. American Naturalist 125:144-152. Smith, Charles R. 1982. Food resource partitioning of fossorial Florida reptiles. p. 173-178. In Herpetological communities: a symposium of the Society for the Study of Amphibians and Reptiles and the Herpetologistsf League, August 1977. Norman J. Scott, Jr., editor. 239 p. U.S. Fish and Wildlife Service, Wildlife Research Report 13. Soule, Michael E., and Daniel Simberloff. 1986. What do genetics and ecology tell us about the design of nature reserves? Biological Conservation 35:19-40. Stout, I. Jack. 1981. Conservation biology of Peromycus floridanus. Florida Scientist 44 (Supplement 1):37. Stout, I. Jack. 1982. Small mammal community structure in sand pine scrub. Bulletin Ecological Society America 63:68-69. Stout, I. Jack, Donald R. Richardson, and Richard E. Roberts. 1988. Response of resident and relocated gopher tortoises to a prescribed burn in a sand pine scrub community. Proced. Gopher Tortoise Relocation Symp., 27 June 1987, Gainesville, FL. Stout, I. Jack, Donald R. Richardson, Richard E. Roberts, and Daniel F. Austin. 1987. Design of a nature preserve in a subtropical urbanizing landscape: application of ecologic and genetic principles. Bulletin Ecological Society of America 68:423-424. Swindell, C. E. 1987. Multivariate analysis of competition and habitat use by small mammals in Florida Sand Pine Scrub. M. S. Thesis. 59 p. University of Central Florida, Orlando. Veno, Patricia A. 1976. Successional relationships of five Florida plant communities. Ecology 57:498-508. White, Peter S., and Susan P. Bratton. 1980. After preservation: philosophical and practical problems of change. Biological Conservation 18:241-255. White, W. A. 1970. The geomorphology of the Florida peninsula. Bureau of Geology, Tallahassee. 164 p. Geological Bulletin No. 51. Wood, Don A. 1987. Official lists of endangered and potentially endangered fauna and flora in Florida. 19 p. Florida Game and Fresh Water Fish Commission.