A FLORISTIC STUDY OF WEAVER CREEK WETLAND, SANTA ROSA COUNTY, FLORIDA Except where reference is made to the work of others, the work described in this thesis is my own or was done in collaboration with my advisory committee. This thesis does not include proprietary or classified information. __________________________________ Courtney R. Holt Certificate of Approval: ____________________ ____________________ Robert Boyd Debbie Folkerts, Chair Professor Assistant Professor Biological Sciences Biological Sciences ____________________ ____________________ Robert Lishak Joe F. Pittman Associate Professor Interim Dean Biological Sciences Graduate School A FLORISTIC STUDY OF WEAVER CREEK WETLAND, SANTA ROSA COUNTY, FLORIDA Courtney R. Holt A Thesis Submitted to the Graduate Faculty of Auburn University in Partial Fulfillment of the Requirements for the Degree of Master of Science Auburn, Alabama May 10, 2008 iii A FLORISTIC STUDY OF WEAVER CREEK WETLAND, SANTA ROSA COUNTY, FLORIDA Courtney R. Holt Permission is granted to Auburn University to make copies of this thesis at its discretion, upon request of individuals or institutions and at their expense. The author reserves all publication rights. _______________________________ Signature of Author _______________________________ Date of Graduation iv THESIS ABSTRACT A FLORISTIC STUDY OF WEAVER CREEK WETLAND, SANTA ROSA COUNTY, FLORIDA Courtney R. Holt Master of Science, May 10, 2008 (B.S., Centre College, 2005) 82 Typed Pages Directed by Debbie Folkerts Steepheads (streams that originate from the bases of valley walls) create a unique freshwater wetland habitat. Currently, steepheads are known to exist only in the southeastern United States along the Gulf Coastal Plain. The biology of these habitats is relatively unknown. The objectives of this study were to provide a comprehensive list of vascular plant species in the wetland of Weaver Creek (a steephead stream in Santa Rosa County, Florida), to compare the upstream and downstream reaches of Weaver Creek, and to compile lists of previously reported species from steephead habitats. Fourteen sampling visits over a period of one year (2006-2007) were made to the study site. Every effort was made to cover the entire length of the creek and to collect all species while in flower or fruit. In addition to the species list, species and taxonomic (generic) richness, v evenness (J?) and diversity (H?) were calculated for the study site. The Jaccard Index (JI) was used to compare the degree of floristic similarity between the upstream and downstream sections of the wetland. Species richness (102 species), evenness (J? = 0.86) and diversity (H? = 3.97) for the entire study site were all relatively high. Taxonomic/generic richness, evenness and diversity were high as well (67 genera, J? = 0.95, H? = 3.98). The degree of community similarity between the upstream and downstream sections of Weaver Creek was very low (JI = 13.33%). Because of this low degree of similarity between the two sections, species and taxonomic richness, evenness and diversity were calculated for each section of the creek to further illustrate the differences between them. The downstream section was both more rich and diverse than the upstream section. However, evenness was slightly greater upstream. This study added 68 species to the list of plants known to occur in steephead habitats. The richness and diversity of the study site illustrate the need for further scientific investigation into steephead habitats. The potential for studying the presence of genetically isolated populations and endemics, as well as rare species, in steepheads provides additional incentive to focus on these habitats. vi ACKNOWLEDGEMENTS This thesis is dedicated to Dr. George Folkerts whose passion and dedication to all aspects of nature has been an inspiration to myself and many others. Without his guidance, I may have never known about steepheads. His knowledge of field biology throughout my graduate career was absolutely indispensable. George?s unique approach to biology and life in general was an endless source of inspiration. I would also like to thank Dr. Debbie Folkerts for her invaluable help and guidance throughout the research and writing phases of this project. You were always available when I needed new ideas and inspiration. I am extremely grateful to you for taking me under your wing and helping me finish this project Thank you also to my committee members, Dr. Bob Lishak and Dr. Bob Boyd, for all of your assistance throughout the project. You were both very helpful in providing me with new perspectives and in helping me focus my study to a more manageable size. Thank you to Dr. Boyd for joining my committee on such short notice as well. Special thanks to Scott Pokswinski and Katie Glynn for their field assistance. Thanks to Eglin Air Force Base for granting me access to Weaver Creek, particularly Amanda Stevens and Bill Tate. Finally, thank you to my family and friends who listened to my frustrations as well as accomplishments during the project. vii Style manual or journal used: Castanea Computer software used: Microsoft Word, Microsoft Excel, MapTech Terrain Navigator, GW Basic viii TABLE OF CONTENTS LIST OF FIGURES............................................................................................................ix LIST OF TABLES..............................................................................................................x I. INTRODUCTION........................................................................................................1 II. METHODS.................................................................................................................11 DESCRIPTION OF THE STUDY SITE..............................................................11 COLLECTION METHODS..................................................................................15 ANALYSIS METHODS.......................................................................................15 III. RESULTS...................................................................................................................17 VASCULAR PLANTS OF WEAVER CREEK WETLAND..............................18 COMPILATION OF PREVIOUSLY REPORTED STEEPHEAD SPECIES...............................................................................................................41 IV. DISCUSSION............................................................................................................43 LITERATURE CITED......................................................................................................49 ix LIST OF FIGURES Figure 1. Map of western Florida panhandle with the location of Weaver Creek noted............................................................................................................56 Figure 2. Topographic map of Weaver Creek showing point of origin and point of entry into Weaver River...........................................................................57 Figure 3. Topographic map of Weaver Creek illustrating the difference in width between upstream and downstream sections...............................................58 Figure 4. Topographic map of Buck Pond collection site................................................59 x LIST OF TABLES Table 1. Species richness, species evenness, species diversity, and community similarity of Weaver Creek wetland..................................................60 Table 2. Taxonomic (generic) richness, evenness and diversity of Weaver Creek wetland........................................................................................................61 Table 3. Summary of vascular plant species reported in the literature to occur in steepheads................................................................................................62 Table 4. Plant species reported from steepheads with no specific location information............................................................................................................67 Table 5. Vascular plant species reportedly located in uplands (sandhills) surrounding steepheads.........................................................................................69 Table 6. Fauna reported from steepheads..........................................................................70 1 INTRODUCTION Steepheads (also listed as a type of seepage stream by the Florida Natural Areas Inventory [FNAI] [1990]) are little-known habitats found along the southeastern Coastal Plain. They were first described by Sellards and Gunter (1918). The Eglin AFB Steephead Monitoring Plan (2004) defined them as ?deep ravines at the headwaters of streams.? Means (1975) provided a more detailed definition: ?perennially wet, first order stream sources that issue from the base of valley headwalls.? The lack of a strict definition of ?steephead? speaks to the paucity of intense biological studies of this habitat type. In contrast to steepheads, gully-eroded ravines are formed by water running over the surface of the land, creating a valley by erosion (Wolfe et al. 1988, Means 1991, Means 2000). Water flow can be heavy at times but is not necessarily constant (Eglin AFB Steephead Monitoring Plan 2004). At the heads of these ravines, the slope is gentle and becomes steeper farther downstream. Due to the scouring action of the water, the ravine appears V-shaped in cross section (Means 2000). Over time, erosion decreases the grade of the slope and causes sediments to enter the stream. The formation of steepheads is strikingly different from that of gully-eroded ravines and somewhat more complex. Steepheads in the Florida Panhandle are commonly found in (but not limited to) the deep sands of the Citronelle formation as well as in younger sands south of the Cody Scarp, an ancient shoreline (Wolfe et al. 1988, 2 Enge 1998). Sediments are of Miocene or Pliocene age and are fine- to coarse-grained sand intermixed with gravel (Platt and Schwartz 1990, Means 2000, USGS 2006). These sands contain little clay or silt and were uncovered about two million years ago following the recession of the Gulf of Mexico (Means 1985). Some sources have indicated that these are the only areas in which steepheads occur (Wolfe et al. 1988, Platt and Schwartz 1990, Rasmussen 2004). However, Sheridan et al. (1999) reported on habitats they described as steepheads (although they called them seepage streams) from western Georgia Fall Line sandhills. The vegetational characteristics of steepheads in this area appear very similar to those in Florida. Steepheads are formed as a stream emerges from beneath the surface and causes the sand above the emergent point to slump. Schumm et al. (1995) referred to the actual water emergence as ?spring sapping? rather than seepage erosion because the water discharge is concentrated into specific point sources. This distinction is important in that it explains the difference between seepage slopes (bogs) and steepheads. It was previously thought that water rapidly percolates through the sand until it reaches a confining layer where it is forced out to form the head of the stream (Sellards and Gunter 1918). Resource Consultants and Engineers, Inc. (1994), however, contended that steepheads can form in homogeneous sediments, indicating that an impermeable layer is not necessary for spring formation. Drilling subsequently revealed that there is no hardpan below the bases of some steepheads, providing direct evidence that a hardpan is not required for seepage to occur (Schumm et al. 1995). Howard (1990) suggested that riverine valleys that form in homogeneous sediments are controlled by the slope of the 3 water table. However, more geologic/hydrologic investigation of steephead formation is needed to fully support this idea. The erosional processes occurring in steepheads, on the other hand, are fairly well understood. The origin of the steephead migrates farther into the sandhill as erosion occurs (Sharp 1938, Schumm et al. 1995). Over time, the outflow of water carries sand downstream and a U-shaped valley (in cross-section) develops. As the springwater removes sand from the base of the slope, the slope becomes undercut. Eventually, the sand slumps down into the emerging spring water and is carried away, beginning the cycle again and allowing the valley to migrate headward (Means 1991). The rate of headward erosion has been estimated to be about two to five centimeters each century; however, a maximum of 70 centimeters per century has also been suggested (Eglin AFB Steephead Monitoring Plan 2004). According to some sources this natural migration of the head is due in part to surface runoff occurring after fires or clear-cuts in the upland longleaf pine habitats (Schumm et al. 1995, Eglin AFB Steephead Monitoring Plan 2004). Means (2000) reported steephead progression as far as five kilometers into the sand body since the stream?s formation. The resultant head of the ravine takes on an amphitheater-like shape, usually slopes down at a 45-degree angle (approximately) and can be up to 35 meters deep. Abiotic factors in steephead ecosystems differ from those in gully-eroded ravines, allowing for a unique mixture of plant and animal species. Because steephead streams are formed by groundwater that has percolated through deep soils, some studies have suggested that the water remains the same temperature year round, usually between 20?C and 22.2?C. (Wolfe et al. 1988, Chafin et al. 1997). However, Enge (1998) found some 4 first-order steephead streams to vary by as much as 15?C during the year. More data are needed to determine actual temperature variation. Because steephead formation is caused by the steady trickle of groundwater through porous substrate, water flow is expected to be fairly constant. Although stream flow is constant from month to month, there is significant variation in water flow from year to year (Schumm et al. 1995). However, steephead streams typically do not run completely dry. Additionally, because the water is filtered through deep layers of sand, it emerges relatively neutral in pH and contains very few pollutants (organic or inorganic), unlike gully-eroded streams, which pick up nutrients and chemicals from a multitude of nonpoint sources such as fields, pastures, roads, and yards (Wolfe et al. 1988, Means 1991). Rainwater is quickly absorbed by the porous sands above steepheads, allowing little opportunity for surface runoff. For these reasons and because ambient temperatures are typically more stable year-round than in the surrounding uplands, steepheads have been said to act as refugia for cool-adapted and stenothermal species (Rasmussen 2004). Such cool-adapted species are present in this otherwise warm region due to the fact that average summer temperatures of the southeastern United States were once nearly 12?C colder than at present (Neill 1957, Watts 1980, Watts et al. 1992). Thus, relict populations of certain species that typically are not found along the Coastal Plain can persist in steepheads (James 1961). The biogeography of steepheads is also interesting in that there are certain genera with discontinuous distributions between eastern North America and eastern Asia. A number of these groups are found in steepheads and other ravines of the Gulf Coastal Plain (Gray 1875, Li 1952, James 1961). For instance, Torreya taxifolia Arn. is located 5 in ravines of the southeastern U.S. (specifically in northern Florida) and has related species (T. grandis Fortune. ex Lindl., T. fargesii Franch., T. jackii Chun., and T. nucifera (L.) Siebold. & Zucc.) in eastern China and Japan (Li 1952). There seems to be a pattern of fairly constant, undisturbed conditions in steepheads. Although fires are frequent in upland pine forests (every one to three years), they rarely reach into steepheads, creating a drastic distinction between uplands and ravines in this region (Delcourt and Delcourt 1977, Schwartz 1994). The environmental constancy and lack of disturbance may allow for some species that are sensitive to environmental change to thrive within steepheads. Due to unique abiotic factors, steephead organisms may evolve differences from related forms not occupying steepheads. Because most steepheads are separated from other steepheads by much drier habitats in which many steephead organisms could not survive, populations in isolation may develop genetic differences over time. Environmental characteristics (physical and vegetational) of steepheads are fairly constant within and among drainages into which they flow (Means 1975, Means 1977). Wolfe et al. (1988) proposed that organisms residing in steepheads might be genetically different from regional populations due to founder effects or intense local selection. Some steephead organisms may experience ecological release due to lack of immigration of more competitive species (Wolfe et al. 1988). Currently there is little evidence to support these statements. Means (2000) suggested that the aforementioned evolutionary processes have had a significant effect on species of plethodontid salamanders. Isolation due to sea level changes may have led to a number of speciation events in this region. For most groups of organisms in steepheads little evidence has yet been gathered to 6 indicate that this is the case. However, endemism has been documented in caddisflies that inhabit ravine systems of northern Florida (Rasmussen 2004). According to Rasmussen (2004), certain species are endemic to ravines (although not steephead ravines specifically) and appear to be associated with habitat type (stream size and ravine type). Intensive studies have not been performed on the vegetation along steephead streams, but some general information does exist (Means 1985, Chafin et al. 1997). More attention has been focused on the plant diversity of the steephead slopes than on the wetland flora. The general pattern from the sandhills above the steephead to the stream at the bottom begins with a xeric longleaf pine-scrub oak community surrounding the ravine grading to wetland species below. Historically, Pinus palustris Mill. dominated the upland areas. However, many of these areas are now converted into pine plantations and housing developments (Means 1991). Absence of fire has also negatively affected this habitat type by allowing for dense growth of understory species. On the upper ravine slopes the community becomes a mixture of xeric, deciduous trees. Halfway down the slope, a mesic forest develops, supporting trees typical of a beech-magnolia climax forest. The lower slopes of steepheads are dominated by an evergreen shrub zone, which contains a variety of species, including, in the Apalachicola region, many of Florida?s rarest plants, including Asarum arifolium Michx., Croomia pauciflora (Nutt.) Torr. and Rhododendron austrinum (Small) Rehder (Wolfe et al. 1988, Eglin AFB Steephead Monitoring Plan 2004). Kwit et al. (1998) supported this species zonation along the slope but mentioned that the abundances of species change gradually (rather than abruptly) from one zone to the next. 7 Means (1985) characterized the valley floor of steepheads as a wetland community dominated by Illicium floridanum Ellis and Magnolia virginiana L. Mosses, liverworts and ferns are also common along the stream. A more recent study by Chafin et al. (1997) reported the presence of three distinct ?baygall? plant associations along steephead streams on Eglin Air Force Base. At the head of the stream is the Illicium floridanum association. Downstream, the Cliftonia monophylla (Lam.) Britt. ex Sarg. association can be found in narrow channels (Chafin et al. 1997). The Chamaecyparis thyoides (L.) B.S.P. - Sphagnum association is found along wide channels with decreased water velocity (Chafin et al. 1997). Some wetland plants which are considered to be relatively uncommon are also found in and along steephead streams including Drosera intermedia Hayne, Kalmia latifolia L., Macranthera flammea (Bartr.) Pennnell, Nuphar advena (Ait.) Ait. f. subsp. ulvacea (Mill. and Standl.) Padgett, Peltandra sagittifolia (Michx.) Morong, Rhododendron austrinum, Rhynchospora crinipes Gale, Sarracenia leucophylla Raf., Sarracenia rubra Walt., and Xanthorhiza simplicissima Marshall (Chafin et al. 1997, U.S. Fish and Wildlife Service 2007). The presence of such species provides further incentive to protect steephead habitats. In addition to vegetation, animal diversity is also high in these ecosystems. Fairly constant water temperature and continuous flow of steephead streams allows for the survival of rare and sensitive species. The environment is buffered both thermally and chemically, providing a suitable year-round habitat for a number of species (Means 1985). 8 Constant water flow has obvious implications for the presence of aquatic species diversity. Darters, including the federally endangered Okaloosa darter (Etheostoma okaloosae Fowler), bluenose shiners (Pteronotropis welaka Evermann & Kendall) and creek chubs (Semotilus atromaculatus Mitchill) have been found year-round at the head of steephead streams (Means 1991, Chafin et al. 1997, Florida Fish and Wildlife Conservation Commission 2005, U.S. Fish and Wildlife Service 2007). Organisms along the banks also benefit from the continuous flow. A variety of salamander species are able to survive year-round in the continuously saturated leaf litter (Wolfe et al. 1988, Means 1991, Means 2000). Two notable amphibian species reported to occur in steephead ravines are the pine barrens tree frog (Hyla andersonii Baird) and the Florida bog frog (Rana okaloosae Moler), both of which are listed as species of special concern (SSC) in Florida (U.S. Fish and Wildlife Service 2007). Hubbell et al. (1956) also suggested that there are orthopterans endemic to such ravines in the Panhandle, although no specific species are mentioned. High water quality allows for survival of pollution-sensitive species. Entrekin et al. (1999) found high invertebrate diversity in all steephead streams studied in southwestern Georgia. In particular, EPT (Ephemeroptera, Plecoptera, Trichoptera) taxa were found, all of which are typically pollution-sensitive organisms. Because of the rugged terrain of steephead valleys, they are not directly under development or logging pressure. However, the uplands are experiencing degradation in a number of ways. Sites on Eglin Air Force Base are offered some degree of protection, but most steepheads on private land are under threat. Longleaf pine formerly dominated the upland habitat but is being replaced by pine plantations, housing and urban developments, and commercial/industrial developments (Florida Fish and Wildlife 9 Commission 2005). Fire exclusion has also led to a decline in quality of longleaf pine habitat in the area. Nearby roads contribute to slope disturbance, leading to erosion and siltation (Chafin et al. 1997). Large-scale disturbance of vegetation above a steephead can lead to increased run-off, and, in turn, a rise in stream turbidity (Schumm et al. 1995, Chafin et al. 1997). The Florida Fish and Wildlife Conservation Commission (2004) listed declining water quality as a threat to the habitat. On Eglin Air Force Base, Chafin et al. (1997) reported additional threats to the health of these ecosystems. They found evidence of direct trash dumping of items such as car parts and paint cans into the streams. Dumping of hazardous wastes in stream watersheds and runoff of fertilizers or biocides from surrounding land have the potential to pollute the shallow ground waters that flow into steephead streams. Invasive species are also a threat to steephead habitats and include feral hogs (Sus scrofa L.), Japanese climbing fern (Lygodium japonicum (Thunb.) Sw.) and Chinese tallow tree (Sapium sebiferum (L.) Roxb.) (Eglin AFB Steephead Monitoring Plan 2004). Research has only recently begun on steepheads and is still in its preliminary stages, making the need for conservation even greater. Not only are steepheads geologically unique, they also provide stable habitat for a variety of animal and plant species, some of which are found only in these ravines along the Gulf Coastal Plain. Means (1977, 1985) has also proposed that ravine ecosystems in the Panhandle act as paleorefugia (a place where ancient species that are extinct elsewhere persist). Much of this diversity is due to the fact that steepheads are actually a combination of four major ecosystems: an aquatic stream habitat, seepage-slope wetlands, moist beech-magnolia forests, and dry oak-cedar forests (Means 1991). Research is imperative in preserving 10 this system and protecting it from degradation. With thorough investigation of this habitat, it is probable that more endemic and perhaps undiscovered species will be found. The objectives of this study were (1) to describe the wetland flora of Weaver Creek steephead from the head of the stream to the foot (its point of entry into Weaver River), (2) to compare the upstream and downstream habitats of Weaver Creek and (3) to compile lists of plant and animal species reported from steephead habitats. 11 METHODS The steephead wetland of Weaver Creek was selected for the study based on its large size and accessibility for study. Additionally, Weaver Creek is relatively undisturbed by humans because of its location on Eglin Air Force Base. Permission was obtained from Eglin Air Force Base to conduct this study. Description of the Study Site Weaver Creek is located 18.8 km southeast of Milton, Florida on Eglin Air Force Base in Santa Rosa County, Florida (coordinates: 30?30?27.84? N, 86?54?46.95? W) (Figure 1). It is a steephead stream (flowing from south to north) with multiple heads (origin points), all of which are found at the bases of sandhills, and is part of the Yellow River drainage. Elevation on top of the ridge at the head is 33.2 m above sea level (asl). The elevation at the point of origin (base of ridge) is 11.3 m asl. At the point the creek enters Weaver River, the elevation drops to 0.91 m asl. Weaver Creek?s total length is 7.18 km (Figure 2), and its depth ranges from a few centimeters at the origin to over three meters deep near its foot. The approximate area of the wetland (area surveyed) is 0.55 km 2 . Valley width at the head is approximately 300 m (including the ravine slope) (Figure 3). The steep walls of the ravine at the head decrease the width of wetland habitat to include primarily only the creek itself, which is approximately 70 m wide at the narrowest point. Water depth at the head does not exceed a few centimeters, and the stream channel is relatively narrow. Due to the dense slope and wetland vegetation, the 12 upstream section of Weaver Creek is more heavily shaded than the downstream portion. At the foot of the stream, the valley is nearly 700 m wide (including the slope) (Figure 3). Downstream, the channel widens and deepens significantly. Thus, the majority of species found downstream are located on soil adjacent to the creek rather than in the stream channel itself. The area surrounding the creek downstream flattens out, creating a wide area (180 m wide) of wetland habitat characterized by constantly saturated soils and increased penetration by sunlight. This section is also heavily shaded by the overstory but contains patches in which the overstory is somewhat thinned, creating light gaps on the forest floor. In addition to the main stream channel, a side stream (Double Head Branch) was surveyed that originates from Buck Pond (Figure 4). A powerline intersects the stream at one point and creates a short stretch of intense sunlight. The remainder of Double Head Branch is typical of the upstream habitat of Weaver Creek. According to the Florida Geological Survey (1993), there are two geological formations on which Weaver Creek is situated. The majority of the creek (approximately three-fourths of its length from the head) is on the Citronelle Formation, which is characterized by fine to coarse-grained sands interspersed with gravel, silt and clay (Florida Geological Survey 1993). The remainder of the creek bed is in an area characterized by deposition of alluvium in the Weaver River floodplain. Sands in this section of the creek are very fine to coarse-grained (Florida Geological Survey 1993). The streambed is composed primarily of sand but is covered with deposits of organic matter in certain areas. Upstream areas of the creek are alluvial, whereas the majority of vegetation downstream grows in nonalluvial mucky areas on either side of the 13 stream channel. There are, however, pockets of alluvium deposited in the downstream section on which limited vegetation occurs as well. There are two main soil series in the streambed. Near the head of the creek are Lakeland sands (both in the stream and on the surrounding lower slopes) (Weeks et al. 1980). This soil type is on level to sloping (0- 30%) terrain and is well drained (Weeks et al. 1980). Downstream portions of the creek contain an association of Dorovan and Pamlico soils, creating a mucky substrate (saturated year round) that has formed via decomposition of plant material (Weeks et al. 1980). The water is remarkably clear for its entire length, indicating little impact from sediment runoff. Water temperature of the stream (23?C) remained fairly constant during the study period and along its length. This held true even when the air temperature had been near 37.8?C for almost a week. The water is acidic with a pH of 5.6. Above the ravine at the head of the stream is xeric sandhill habitat into which the ravine is carved. Plant composition of the sandhill above the steephead is drastically different from the slope and wetland vegetation. Dominant overstory trees of the sandhill habitat include Pinus elliottii Engelm., Pinus palustris, Quercus geminata Small, Quercus hemisphaerica Bartr. ex Willd., and some invading Pinus clausa (Chapm. ex Engel.) Sarg. Primary understory trees include Ilex opaca Ait., Ilex vomitoria Ait., Quercus incana Bartr., Quercus laevis Walt., Vaccinium corymbosum L., and Vaccinium elliottii Chapm. Crataegus uniflora Muenchh., Diospyros virginiana L. and Magnolia grandiflora L. are occasional in the understory. Other woody plants, vines and herbs common on the sandhill are Licania michauxii Prance, Pteridium aquilinum (L.) Kuhn, Satureja coccinea (Nutt. ex Hook.) Bertol., Serenoa repens (Bartr.) Small, Smilax 14 auriculata Walt., and Vitis rotundifolia Michx. Ground cover is sparse with scattered clumps of Aristida stricta Michx. var. beyrichiana (Trin. & Rupr.) D.B. Ward and lichens of the genus Cladina (Nyl.) Nyl., particularly C. evansii Abbayes. Near the head, the amount of relief is significant, creating a deep ravine. The ravine contains a variety of plant species, some of which are also found in the upland sandhill or the wetland at the base of the slope. Dominant large trees found on the steep slope are Magnolia grandiflora L., Nyssa sylvatica Marsh., Oxydendrum arboreum (L.) DC., Quercus nigra L., and Quercus phellos L. Understory slope trees include Carya glabra (P. Mill.) Sweet (very few), Cornus florida L., Ilex decidua Walt., Ilex opaca, Ilex vomitoria, Magnolia ashei Weatherby, Osmanthus americanus (L.) Benth. & Hook. f. ex Gray, Ostrya virginiana (P. Mill.) K. Koch, a few invading Pinus clausa, and Vaccinium corymbosum. Shrubs and small trees on the slope include Asimina parviflora (Michx.) Dunal, Hamamelis virginiana L., Illicium floridanum, Serenoa repens, and Vaccinium elliottii. Common vines and herbs on the slope are Mitchella repens L., Smilax pumila Walt. and Vitis rotundifolia. Downstream the terrain slopes more gradually. Many sandhill species extend nearly to the creek. The two main overstory trees present are Pinus palutris and Pinus elliottii. The understory consists primarily of Quercus incana Bartr. and Quercus laevis Walt. Acer rubrum L. begins to appear near the creek. Shrubs of the downstream slope include Chrysoma pauciflosculosa (Michx.) Greene, Ilex glabra (L.) Gray and Ilex vomitoria. The two common vines of the area are Smilax auriculata and Vitis rotundifolia (which is found near the creek). Ground cover consists primarily of Aristida stricta var. beyrichiana, Licania michauxii and Pteridium aquilinum. 15 Collection Methods Collections began on 20 August 2006 and were concluded on 18 September 2007. Fourteen visits were made during this period. Collection dates were scheduled in an effort to collect specimens when they were either in flower or fruit. Specific dates are as follows: 20 August 2006, 4 September 2006, 30 September 2006, 19 November 2006, 18 February 2007, 24 February 2007, 3 March 2007, 29 March 2007, 3 May 2007, 1 June 2007, 5 July 2007, 26 July 2007, 11 August 2007, and 18 September 2007. The entire wetland area along the stream was surveyed in an effort to obtain samples of every vascular species present. The wetland was defined as areas in and along the stream with saturated soil and emergent vegetation. At least one voucher specimen of most species found was deposited in the Freeman Herbarium (AUA) at Auburn University, Auburn, Alabama. Collection of federally protected species and those protected by the state of Florida was avoided. Photographs served as documentation for such species. Taxonomy follows Godfrey and Wooten (1981) unless otherwise noted. Location information of sensitive species was provided to Eglin Air Force Base. Analysis Methods Two distinct habitats within the wetland area (upstream and downstream) were described based on floristic composition. Visually, these two habitats are easily discernible based on vegetation present, stream width and relative vegetation density. The transition zone (ecotone) between these two separate habitats was sampled, but the specimens were not used to characterize either of the habitats. Species richness, species evenness, species diversity (the Shannon Index), and community similarity (the Jaccard 16 Index) were calculated for each section of the creek and for the creek as a whole. Species diversity and evenness were calculated using GW-Basic with the following formulas: H? = -? (p i * ln(p i )) and J? = H?/ln(S), where H? = diversity, S = number of species, p i = proportion of species i in total species, and J? = evenness (Ludwig and Reynolds 1988). Relative abundance values were used to calculate species evenness and diversity. Each species was assigned to one of the following categories: rare (0-5 individuals/colonies), infrequent (6-30 individuals/colonies), occasional (31-100 individuals/colonies), frequent (100-500 individuals/colonies), or abundant (500+ individuals/colonies). In order to measure diversity at the generic level, taxonomic evenness and diversity were also calculated using counts of species within genera instead of individuals within species. Community similarity between the upstream and downstream sections was determined using the Jaccard Index: JI = j/r x 100, where j = number of species found at both locations and r = number of species unique to either location. The Jaccard Index was chosen for this analysis because it requires only presence-absence data and is not biased even at small sample sizes (Ludwig and Reynolds 1988). 17 RESULTS Regarding the wetland floristic composition of Weaver Creek steephead, 102 species were recorded from 67 different genera and 46 families (see annotated list below). Of these, 10 species are listed in Florida as commercially exploited, threatened or endangered. The results of the species richness, evenness and diversity analyses reveal an interesting pattern (Table 1). Generic richness, evenness and diversity follow a similar pattern (Table 2). There is a distinct difference between the floristic composition of the upstream section (reaching approximately 3.40 km downstream from the head) and the downstream section (from 3.40 km from the head to the creek?s entry point into Weaver River) of Weaver Creek (Figure 3). This distinction is best illustrated by the low degree of similarity between the two habitats (Table 1). Composition differences between upstream and downstream sections are also well illustrated by the numbers of rare or infrequent species found at each site (Table 1). Below is a complete list of vascular plant species found in the Weaver Creek wetland along with general location, habitat information, collection numbers, and other significant notes. 18 Vascular Plants of Weaver Creek Wetland PHYLUM LYCOPODIOPHTYA FAMILY LYCOPODIACEAE Lycopodium alopecuroides L. ? foxtail clubmoss Holt 175 Foxtail clubmoss is rare along the lower portion of Weaver Creek. It is found exclusively in mucky soils alongside the stream beneath openings in the canopy, which allow increased light intensity. PHYLUM POLYPODIOPHYTA FAMILY BLECHNACEAE Lorinseria areolata (L.) T. Moore ? netted chain fern Holt 115 Netted chain fern is occasional along Weaver Creek. It is mostly found along the edge of the stream or in very shallow water in the stream itself. FAMILY THELYPTERIDACEAE Thelypteris palustris Schott var. pubescens (Lawson) Fern. ? marsh fern Holt 165 Marsh fern is rare along Weaver Creek. Its habitat is much like that of netted chain fern (moist soils alongside the stream or in shallow areas of the stream itself). North American populations belong to variety pubescens (Smith 1993). FAMILY OSMUNDACEAE Osmunda cinnamomea L. ? cinnamon fern Holt 126 Cinnamon fern is occasional in the upstream section of Weaver Creek. It is found growing in the stream (shallow areas) and along the edges in saturated soil. This species is listed as commercially exploited in Florida (USDA 2007). 19 PHYLUM PINOPHYTA FAMILY CUPRESSACEAE Chamaecyparis thyoides (L.) B.S.P. ? Atlantic white cedar Holt 141 Atlantic white cedar is abundant in the lower portion of Weaver Creek. It is found on the wide area of mucky soil bordering the creek. The largest individuals are approximately 22 cm DBH. A few specimens contain growths of vine wicky (Pieris phyllyreifolia). Weaver Creek populations fall within the range suggested by Li (1962) for Chamaecyparis henryae. Little (1966) considered C. henryae a variety. Michener (1993) considered C. henryae to simply be a variant of C. thyoides, a viewpoint which is adopted here. Taxodium ascendens Brogn. ? pond-cypress Holt 171 Pondcypress is abundant in the lower portion of Weaver Creek along the wide stream banks in mucky, inundated soils. The largest specimens are approximately 24 cm DBH. A number of individuals supported growths of vine wicky (Pieris phyllyreifolia). Watson (1983) considered both pond-cypress (Taxodium ascendens) and bald-cypress to be subspecies of Taxodium distichum. Godfrey (1988) supported the separation of these two into separate species, a view which is followed here. FAMILY PINACEAE Pinus elliottii Engelm. var. elliottii ? slash pine Holt 244 Slash pine is abundant along the entire length of Weaver Creek. It is found growing along the lower slopes and directly adjacent to the creek in upstream sections and in the broad expanse of saturated soil in the downstream portions. Identification of 20 this variety is based on its known range and the distinctive characteristics of its seedling as compared to P. elliottii var. densa Little and Dorman. PHYLUM MAGNOLIOPHYTA CLASS LILIOPSIDA FAMILY LILIACEAE Lilium iridollae Henry ? panhandle lily Holt 150 The panhandle lily is frequent along a section in the lower part of Weaver Creek. It grows in a shady expanse of mucky soil adjacent to the stream. The panhandle lily is of conservation concern and, therefore, is listed as G2, S2 and endangered in Florida by FNAI (2007). It is also under consideration for federal protection (Skinner 2002). The population at Weaver Creek appears quite healthy despite the fact it is not located in its typical habitat (open seepage slopes). FAMILY ARECACEAE Sabal minor (Jacq.) Pers. ? swamp palm Holt 136 Swamp palm is occasional along the creek and can be found growing in shallow water as well as along the banks. The largest specimen found was located in the stream and was approximately 1.5 m tall. FAMILY POACEAE Aristida patula Chapm. ex Nash ? tall threeawn Holt 217 Tall threeawn is infrequent along the upstream banks of Weaver Creek. It grows in clumps directly adjacent to the stream on very moist soils. 21 Arundinaria gigantea (Walt.) Walt. ex Muhl ? giant cane Holt 167 Giant cane is occasional along Weaver Creek. It can be found growing primarily on the stream bank in moist soil. Chasmanthium laxum (L.) Yates ? slender woodoats Holt 120 Slender woodoats is rare along Weaver Creek. It is found intermittently in small clumps along the bank. Dichanthelium ensifolium (Baldw. ex Ell.) Gould ? cypress witchgrass Holt 214 Cypress witchgrass is rare along Weaver Creek. It is found growing along a small path next to the creek. Godfrey and Wooten (1981) and LeLong (1984) place this species in the genus Panicum. Gould and Clark (1978) place it in Dichanthelium, which was previously considered a subgenus of Panicum. Aliscioni et al. (2003) support the elevation of Dichanthelium to genus level based on differences in the chloroplast ndhF gene. Additionally, some authors have considered this species a variety of D. dichotomum (Gould and Clark 1978). Godfrey and Wooten (1981) kept the species separate (although under the genus Panicum), and Aliscioni et al. (2003) support the distinction between the two at the species level. I adopt the viewpoints of elevating the former subgenus to genus and of keeping D. ensifolium a separate species from D. dichotomum. 22 FAMILY ARACEAE Orontium aquaticum L. ? golden-club Holt 176 Golden-club is frequent in the lower section of Weaver Creek and is found growing in the stream, both in shallow and deep water. Often, plants are found with their leaves completely submerged. It is more common, though, to find plants with primarily emergent leaves. Peltandra sagittifolia (Michx.) Morong ? spoonflower Holt 148 Spoonflower is occasional along the lower sections of the creek. It is found along the wide bank in mucky, slightly acidic soil, which is occasionally inundated by water, causing small pools to develop. Although not listed by the FNAI, spoonflower populations are monitored by conservation authorities where it occurs (Thompson 2000). FAMILY BURMANNIACEAE Apteria aphylla (Nutt.) Barnh. ex Small ? nodding-nixie Holt 108 The nodding-nixie is extremely rare (only one specimen found) at the head of the stream. It grows on the lower slope of the ravine in moist soil. FAMILY CYPERACEAE Carex atlantica Bailey ? Atlantic sedge Holt 211 Atlantic sedge is rare along the lower portion of Weaver Creek. It is found close to the edge of the stream in an area of mucky soil. Carex elliottii Schwein. & Torr. ? Elliott?s sedge Holt 209 Elliott?s sedge is infrequent along Weaver Creek. It is found close to the edge of the stream in areas of saturated soil. 23 Carex glaucescens Ell. ? clustered sedge Holt 224 Clustered sedge is occasional along Weaver Creek in the downstream portions. It grows in a wide area of saturated soil alongside the stream. Carex intumescens Rudge ? greater bladder sedge Holt 210 Greater bladder sedge is occasional along the downstream section of the creek. It is found growing in a wide area of saturated soil adjacent to the stream. Dulichium arundinaceum (L.) Britt. ? three-way sedge Holt 142 Three-way sedge is frequent in the lower section of Weaver Creek. This species can be found growing in the broad area of saturated soil alongside the stream as well as on islands of vegetation floating in the stream. Eleocharis elongata Chapm. ? slim spikerush Holt 221 Slim spikerush is occasional along the lower section of Weaver Creek. It was found growing in sandy soil in the stream. It is believed that the majority of this species grows with its leaves submerged. It cannot be determined definitely as to whether the submerged plants are also slim spikerush, but the two are found in close proximity, often growing directly next to each other. The submerged leaves are longer and narrower and contain no flowers or fruit. The emergent leaves are easily identifiable by their reproductive structures. Rhynchospora capitellata (Michx.) Vahl ? brownish beaksedge Holt 222 Brownish beaksedge is occasional in the lower section of Weaver Creek and can be found growing along the bank as well as on floating islands of vegetation in the stream itself. Godfrey and Wooten (1981) synonymized R. leptocarpa with R. capitellata. 24 Sorrie (2000) suggested that R. leptocarpa be distinguished as a separate species. I will adhere to Godfrey and Wooten (1981) on this subject. Rhynchospora cephalantha Gray ? bunched beaksedge Holt 193 Bunched beaksedge is occasional in the lower section of Weaver Creek. It grows most commonly in the area of saturated soil on either side of the stream. Rhynchospora chalarocephala Fern. & Gale ? loosehead beaksedge Holt 185 The loosehead beaksedge is infrequent along Weaver Creek. It is found on the wide expanse of saturated soil bordering the downstream section. Rhynchospora corniculata (Lam.) Gray ? shortbristle horned beaksedge Holt 181 Shortbristle horned beaksedge is rare along Weaver Creek. Its habitat consists of the wide expanse of saturated soil alongside the downstream portion of the creek. Rhynchospora curtissii Britt. ex Small ? Curtiss? beaksedge Holt 174 Curtiss? beaksedge is rare on Weaver Creek. The few specimens found are growing in clumps at the base of pondcypress trees, which are located in the saturated soil beside the creek. Rhynchospora filifolia Gray ? threadleaf beaksedge Holt 233 Threadleaf beaksedge is infrequent along Weaver Creek. This species grows along the edge of the stream in saturated soil. Rhynchospora glomerata (L.) Vahl ? clustered beaksedge Holt 117 Clustered beaksedge is occasional along Weaver Creek. It is found growing along the bank in both upstream and downstream sections of the creek. 25 Rhynchospora gracilenta Gray ? slender beaksedge Holt 155 Slender beaksedge is occasional along Weaver Creek and can be found along the banks of the stream as well as on the saturated soil further into the forested area alongside the stream. Rhynchospora macra (C.B. Clarke ex Britt.) Small ? large beaksedge Holt 172 Large beaksedge is rare along Weaver Creek. This species is found in the broad expanse of saturated soil alongside the stream, usually in areas of greater amounts of sunlight. Rhynchospora rariflora (Michx.) Ell. ? fewflower beaksedge Holt 234 The fewflower beaksedge is rare on Weaver Creek. The plant inhabits vegetation islands in the stream. Scirpus cyperinus (L.) Kunth ? woolgrass Holt 200 Woolgrass is infrequent along a tributary to Weaver Creek (originating from Buck Pond) (Figure 4). The habitat where the specimens are located is disturbed by a powerline road, thus creating an artificially sunny area along a small stretch of the tributary. Scirpus etuberculatus (Steud.) Kuntze ? Canby?s bulrush Holt 213 Canby?s bulrush is rare along the creek and grows in areas of saturated soil bordering the stream, on vegetation islands in the stream, and in shallow water in the stream. Smith (2002) placed this species in the genus Schoenoplectus (Reichenbach) Palla (formerly a subgenus). Godfrey and Wooten (1981) and Wunderlin and Hansen (2004) retained the species in the genus Scirpus, a viewpoint which is followed here. 26 Websteria confervoides (Poir.) S.S. Hooper ? algal bulrush Holt 248 Algal bulrush is rare along Weaver Creek. It grows in the lower section of the stream among other vegetation. This is a fairly atypical habitat for algal bulrush, which is usually found in lakes and ponds. FAMILY ERIOCAULACEAE Eriocaulon decangulare L. ? tenangle pipewort Holt 146 The tenangle pipewort is abundant in the lower section of Weaver Creek. It grows on the wide expanse of saturated soil bordering the creek. Lachnocaulon beyrichianum Sporl. ex Koern. ? southern bogbutton Holt 242 The southern bogbutton is occasional along Weaver Creek and can be found in the wide expanse of saturated soil along the downstream section of the creek. FAMILY JUNCACEAE Juncus canadensis J. Gay ex Laharpe ? Canadian rush Holt 198 Canadian rush is infrequent in a tributary of Weaver Creek (originating from Buck Pond) (Figure 4). The habitat is disturbed by a powerline road, creating an area of artificially greater sun intensity along the stretch in which this species is found. There are two varieties and two forms proposed to exist, but they do not seem to be biologically distinct (Brooks and Clemants 2000). Therefore, I will leave the identification of this specimen at the species level. Juncus marginatus Rostk. ? grassleaf rush Holt 220 The grassleaf rush is infrequent along the lower portions of Weaver Creek and is found growing only along small paths and clearings next to the creek. This species is 27 synonymized with a number of different species (Brooks and Clemants 2000). However, I will adhere to Godfrey and Wooten (1981) who use J. marginatus (with which three species are synonymized ? J. aristulatus Michx, J. biflorus Ell. and J. longii Fern.). Juncus polycephalus Michx. ? manyhead rush Holt 240 The manyhead rush is infrequent along Weaver Creek. It grows primarily on floating islands of vegetation in the stream but also inhabits the very edge of the stream bank where mucky soil is present. Juncus tenuis Willd. ? path rush Holt 207 Path rush is rare along Weaver Creek. The only specimens encountered were found growing on a small path next to the stream in compacted, moist soil. Juncus trigonocarpus Steud. ? redpod rush Holt 139 Redpod rush is infrequent along a tributary of Weaver Creek (originating from Buck Pond) (Figure 4). The habitat in which this species is found is disturbed by a powerline road, creating an artificially sunny stretch along the tributary. It grows along the raised bank of the tributary. FAMILY NARTHECIACEAE Lophiola aurea Ker Gawl. ? golden-crest Holt 218 Golden-crest is frequent in the lower section of Weaver Creek. It is present in a wide area of saturated soil that borders the creek. More specifically, it is found primarily in patches of greater sunlight within the broad areas of mucky soil. Godfrey and Wooten (1981) referred to this species as L. americana and synonymize L. aurea with it. Fernald (1921) recognizes these as two distinct species (L. aurea being the southeastern species), 28 a viewpoint which is adopted here. Family placement of golden-crest is not well resolved as this time. It has traditionally been placed in Haemodoraceae (Robertson 1976, Godfrey and Wooten 1981). Ambrose (1985) placed the species in Liliaceae; Wunderlin and Hansen (2004) suggested an affinity with Nartheciaceae; Simpson (1983) placed the species in Tecophiliaceae. I will adhere to the idea that golden-crest be placed in the Nartheciaceae based on morphological similarities to other species of that family. FAMILY ORCHIDACEAE Platanthera blephariglottis (Willd.) Lindl. var. conspicua (Nash) Luer - Holt 245 white fringed orchid The white fringed orchid is rare along Weaver Creek and grows in the wide expanse of saturated soil in the downstream sections of the creek. Godfrey and Wooten (1981) placed this species in the genus Habenaria. However, I will follow Sheviak (2002) in placing it in the genus Platanthera. Additionally, Godfrey and Wooten (1981) did not recognize the variety listed. Therefore, I will follow Sheviak (2002), Wunderlin and Hansen (2004) and USDA (2007) in listing the specimens at Weaver Creek as variety conspicua. Platanthera cristata (Michx.) Lindl. ? crested fringed orchid Holt 109 The crested fringed orchid is infrequent along the length of Weaver Creek. It is found growing in both the upper and lower reaches of the creek along the bank. Soil on which it grows is moist. Godfrey and Wooten (1981) placed this species in the genus Habenaria; however, I will follow Sheviak (2002) in placing it in the genus Platanthera. This species is listed as threatened in Florida (USDA 2007). 29 FAMILY SMILACACEAE Smilax laurifolia L. ? bamboo briar Holt 208 Bamboo briar is abundant throughout the length of Weaver Creek. Smilax smallii Morong ? lanceleaf greenbriar Holt 125 Lanceleaf greenbriar is abundant along the length of Weaver Creek. FAMILY SPARGANIACEAE Sparganium americanum Nutt. ? burr reed Holt 151 Burr reed is infrequent along the creek. It grows on the vegetation islands along the edges of the downstream portions of the creek. FAMILY XYRIDACEAE Xyris fimbriata Ell. ? fringed yelloweyed grass Holt 170 Fringed yelloweyed grass is occasional in the lower portion of Weaver Creek. It grows on the broad expanse of saturated soil that borders both sides of the creek as well as on the vegetation islands within the stream. Xyris smalliana Nash. ? Small?s yelloweyed grass Holt 147 Small?s yelloweyed grass is occasional along the lower section of the creek and grows in mucky soil along the banks of the creek as well as on the vegetation islands in the stream channel. 30 CLASS MAGNOLIOPSIDA FAMILY NYMPHAEACEAE Nuphar advena (Ait.) Ait. f. subsp. ulvacea Holt 152 (Mill. and Standl.) Padgett ? spatterdock Spatterdock is occasional in the lower portion of Weaver Creek. It grows both in the main stream and in side streams found along the broad flat areas bordering the main stream. Beal (1956) and Godfrey and Wooten (1981) considered all North American taxa in the genus Nuphar to be subspecies of Nuphar luteum. Wiersema and Hellquist (1997) suggested that Gulf coast populations of N. ulvacea (due to much intergradation) be listed as a subspecies of N. advena, a viewpoint which is adopted here. Nymphaea odorata Ait. subsp. odorata ? fragrant water-lily Holt 182 Fragrant water lily is infrequent along Weaver Creek and is found growing along the downstream portions in the stream itself as well as in side streams that contain sufficient amounts of water. The subspecies rank is assigned here based on Wiersema and Hellquist (1997). FAMILY MAGNOLIACEAE Magnolia grandiflora L. ? southern magnolia Southern magnolia is abundant in the upper section of Weaver Creek and is found growing on the stream banks. It also can be found growing abundantly on the steephead slopes. The largest individual is approximately 18 cm DBH. Magnolia virginiana L. ? sweet-bay Holt 106 Sweet-bay is abundant in the upper section of Weaver Creek and occasional in the lower section. Specimens are found growing in shallow water in the stream as well as on the lower slope and stream bank. The largest specimen is approximately 15 cm DBH. 31 FAMILY LAURACEAE Persea palustris (Raf.) Sarg. ? swamp red bay Holt 105 Swamp red bay is occasional in the upper reaches of the creek. Individuals grow mostly in shallow waters near the head. FAMILY ILLICIACEAE Illicium floridanum Ellis ? Florida anise Holt 194 Florida anise is abundant in the upper portion of Weaver Creek and grows in shallow areas of the stream, along the bank, and on the lower slope. CLASS ROSOPSIDA FAMILY HAMAMELIDACEAE Hamamelis virginiana L. ? witch hazel Holt 127 Witch hazel is infrequent along the upper section of Weaver Creek. Its typical habitat includes the moist, sandy soils on the banks of the stream. FAMILY MYRICACEAE Myrica cerifera L. ? wax-myrtle Holt 102 Wax-myrtle is abundant in the upper section of Weaver Creek. It can also infrequently be found growing in the lower portion of the creek. Typically, it grows in shallow areas of the stream itself (on sandy soils). In downstream areas, it is found in the broad expanse of mucky soil bordering the stream. 32 Myrica heterophylla Raf. ? evergreen bayberry Holt 128 Evergreen bayberry is occasional along the upper portions of Weaver Creek. It grows in shallows areas of the stream (on sandy soils). Wunderlin and Hansen (2004) referred to this species as M. caroliniensis Mill. However, I will adhere to Godfrey and Wooten (1981) in calling it M. heterophylla. Myrica inodora Bartr. ? odorless wax-myrtle Holt 130 Odorless wax-myrtle is infrequent along the upper portions of Weaver Creek. Much like the above two species, it grows in shallow areas of the creek. Additionally, it is rarely found along the lower section of the creek in the areas of mucky soil bordering the creek. FAMILY CLUSIACEAE Hypericum brachyphyllum (Spach) Steud. ? coastal plain St. John?s wort Holt 154 Coastal plain St. John?s wort is infrequent along Weaver Creek. It grows on areas of mucky soil bordering the creek. Hypericum fasciculatum Lam. ? peelbark St. Johnswort Holt 228 Peelbark St. John?s wort is infrequent along Weaver Creek. It is found inhabiting islands of vegetation growing in the stream. 33 FAMILY SARRACENIACEAE Sarracenia leucophylla Raf. ? whitetop pitcherplant Holt 145 The whitetop pitcherplant is occasional along Weaver Creek. It grows in the wide expanse of mucky substrate that borders the downstream portions of the creek. According to FNAI (2007), its global and state ranks are G3 and S3, respectively. It is also listed as endangered in Florida by FNAI (2007). Sarracenia psittacina Michx. ? parrot pitcherplant Holt 191 The parrot pitcherplant is occasional along lower Weaver Creek. It grows in the wide expanse of mucky substrate alongside the creek. The parrot pitcherplant is listed as threatened in Florida (USDA 2007). Sarracenia pupurea L. ? purple pitcherplant Holt 195 The purple pitcherplant is occasional along Weaver Creek. Its habitat consists of a broad area of saturated soil alongside the stream. Naczi et al. (1999) considered populations of purple pitcherplant in northwestern Florida, southern Alabama, southwestern Georgia, and southeastern Mississippi to be S. rosea. However, I will adhere to Godfrey and Wooten (1981) and Cheek et al. (1997) in retaining the species under its original name. This species is listed as threatened in Florida (USDA 2007). Sarracenia rubra Walt. ? red pitcherplant Holt 149 The red pitcherplant is occasional along Weaver Creek. It grows in the same general area as the other Sarracenia species found at the study site ? wet mucky soils bordering the stream. Godfrey and Wooten (1981) acknowledged the large amount of taxonomic studies completed in the decade prior to publishing their work. However they chose to remain with the original classification and not acknowledge any subspecies, a 34 view which is followed here. The red pitcherplant is listed as G4, S3, and threatened in Florida (FNAI 2007). FAMILY DROSERACEAE Drosera capillaris Poir. ? pink sundew Holt 189 Pink sundew is occasional along lower Weaver Creek and grows in areas of saturated soil. Drosera intermedia Hayne ? water sundew Holt 158 The water sundew is infrequent along lower Weaver Creek. Its habitat consists of a wide expanse of mucky soil on either side of the stream. FNAI (2007) lists the water sundew as G5, S3, and threatened in Florida. FAMILY CYRILLACEAE Cliftonia monophylla (Lam.) Britt. ex Sarg. ? black titi Holt 143 Black titi is frequent in the upper section of Weaver Creek. It grows both along the banks of the stream in moist soil as well as in shallow water in the stream. Cyrilla racemiflora L. ? white titi Holt 103 White titi is abundant in the upper section of Weaver Creek. Its habitat includes both the stream (shallow water) and the edges of the banks. FAMILY ERICACEAE Gaylussacia mosieri Small ? woolly huckleberry Holt 206 Woolly huckleberry is rare along the lower section of Weaver Creek. It grows in mucky soil that extends over a broad area away from the stream. 35 Lyonia lucida (Lam.) K. Koch ? fetterbush Holt 123 Fetterbush is present along the entire length of Weaver Creek. It is abundant in the upper portion of the creek and grows in the stream and along the edge. Downstream, it is frequent, though not abundant, and grows in a wide area of saturated soil bordering the stream. Oxydendrum arboreum (L.) DC. ? sourwood Holt 135 Sourwood is occasional in the upper section of Weaver Creek. It is found growing in the stream and along the edge in moist soil. Pieris phyllyreifolia (Hook.) DC. ? vine wicky Holt 179 Vine wicky is occasional on Weaver Creek. It grows under the bark of Taxodium ascendens and Chamaecyparis thyoides with its roots in saturated soil at the base of the trees. Up to 60 emergences on one individual of T. ascendens were counted. Many emergences may represent only one individual of the species. Rhododendron viscosum (L.) Torr. ? swamp azalea Holt 204 Swamp azalea is present throughout the length of Weaver Creek. Upstream, it is frequent within the stream in shallow water and along the banks in moist soil. Downstream, swamp azalea becomes more occasional and grows along the moist banks of the stream (usually in greater sunlight). It does not appear on the mucky substrate in the downstream areas dominated by pondcypress and Atlantic white cedar. Vaccinium corymbosum L. ? highbush blueberry Holt 118 Highbush blueberry is occasional on Weaver Creek. It grows in moist soil along the banks of the stream in addition to lower slopes of the ravine. 36 Vaccinium elliottii Chapm. ? Elliott?s blueberry Holt 124 Elliott?s blueberry is frequent in the upper portion of Weaver Creek. It grows primarily in shallow water in the stream or on moist soil along the banks. Wunderlin and Hansen (2004) have synonymized this species with V. corymbosum. However, Godfrey and Wooten (1981), Munoz and Lyrene (1985) and Uttal (1987) consider these separate species, a conclusion to which I will adhere. FAMILY SAXIFRAGACEAE Itea virginica L. ? Virginia willow Holt 104 Virginia willow is abundant in the upper section of Weaver Creek. It grows in shallow areas of the stream. Family placement of this species is uncertain. Wunderlin and Hansen (2004) place it in Iteaceae, USDA (2007) places it in Grossulariaceae and Godfrey and Wooten (1981) place it in Saxifragaceae. I will follow Godfrey and Wooten until further taxonomic research has been completed. FAMILY CELASTRACEAE Euonymus americanus L. ? strawberry bush Holt 156 Strawberry bush is rare along Weaver Creek. It was found growing on the bank of the stream in sandy moist soil. FAMILY AQUIFOLIACEAE Ilex coriacea (Pursh) Chapm. ? large gallberry Holt 162 Large gallberry is abundant in the upper portion of Weaver Creek. It grows in shallow water in the stream as well as along the banks in moist soil. 37 Ilex glabra (L.) Gray ? gallberry Holt 215 Gallberry is frequent in the upper portion of Weaver Creek. It is found growing in shallow water in the stream and along the bank in moist soil. Ilex opaca Ait. var. opaca ? American holly Holt 101 American holly is abundant in the upper portion of Weaver Creek. It grows from the stream up to mid-slope on the ravine. Ilex vomitoria Ait. ? yaupon Holt 116 Yaupon is occasional along Weaver Creek. It is found along the banks of the stream and lower slopes of the ravine in sandy soil. Downstream, it grows on the edges of the mucky areas of soil that border the creek in slightly less saturated soil. FAMILY POLYGALACEAE Polygala brevifolia Nutt. ? littleleaf milkwort Holt 169 Littleleaf milkwort is infrequent along Weaver Creek and is found growing in the broad expanse of saturated soil bordering the downstream sections of the creek. Polygala hookeri Torr. & Gray ? Hooker?s milkwort Holt 230 Hooker?s milkwort is infrequent along the downstream section of Weaver Creek. It grows in the broad expanse of saturated soil on either side of the creek. FAMILY FABACEAE Strophostyles helvola (L.) Ell. ? trailing fuzzybean Holt 227 Trailing fuzzybean is rare along Weaver Creek. This vine is not typically considered to be a wetland species. At the study site, it is found growing in the 38 downstream section in a depression in moist soil. The area in which trailing fuzzybean is found is adjacent to an area of disturbance (a dirt road that crosses Weaver Creek). FAMILY ROSACEAE Aronia arbutifolia (L.) Nieuwl. ? red chokeberry Holt 246 Red chokeberry is rare along Weaver Creek. The few specimens found were in the downstream section growing in the creek bed in shallow water. FAMILY VITACEAE Vitis rotundifolia Michx. ? muscadine Holt 133 Muscadine is frequent along the length of Weaver Creek. In upstream areas, it grows on a variety of trees and shrubs located in the stream as well on the banks and lower slopes of the ravine. Downstream it grows on a multitude of species as well along the banks and in the broad expanse of mucky soil present on either side of the stream. FAMILY MELASTOMATACEAE Rhexia petiolata Walt. ? fringed meadowbeauty Holt 197 Fringed meadowbeauty is infrequent along a tributary to Weaver Creek (originating from Buck Pond) (Figure 4). The area of the tributary in which this species is found is disturbed by a powerline, creating a short stretch of uncharacteristically intense sunlight. 39 FAMILY ONAGRACEAE Ludwigia maritima Harper ? seaside primrose-willow Holt 219 Seaside primrose-willow is rare along Weaver Creek. It is found in a disturbed area along the bank from which most of the overstory has been removed by human action, providing ample sunlight. The soil is moist and very sandy. FAMILY CORNACEAE Cornus florida L. ? flowering dogwood Holt 129 Flowering dogwood is rare in the wetland of Weaver Creek. However, one specimen was found growing along the bank in the upstream section of the creek in moist, sandy soil. This location is odd considering flowering dogwood is not typically a wetland species. Cornus foemina Mill. ? swamp dogwood Holt 160 Swamp dogwood is occasional on Weaver Creek. It is found growing along the bank of the stream in very moist, sandy soils. Nyssa biflora Walt. ? swamp tupelo Holt 144 Swamp tupelo is occasional along the stream. Its habitat consists of a broad area of saturated soil on either side of the stream. FAMILY CAPRIFOLIACEAE Viburnum nudum L. ? possumhaw Holt 114 Possumhaw is abundant in the upper section of Weaver Creek. It is found most commonly growing in shallow waters in the creek on sandy soil. 40 FAMILY RUBIACEAE Mitchella repens L.- partridgeberry Holt 122 Partridgeberry is infrequent near the head of Weaver Creek. It grows on moist, sandy substrate on the bank/lower slope of the ravine. FAMILY OLEACEAE Osmanthus americanus (L.) Benth. & Hook. f. ex Gray ? American olive Holt 205 American olive is frequent in the upper section of Weaver Creek. It grows along the bank as well as in the stream. Although Godfrey and Wooten (1981) call it O. americana, it appears O. americanus is more widely used. FAMILY BIGNONIACEAE Bignonia capreolata L. ? crossvine Holt 107 Crossvine is infrequent on the upper section of Weaver Creek along the bank and in the stream. FAMILY LENTIBULARIACEAE Pinguicula primuliflora Wood and Godfrey ? southern butterwort Holt 188 The southern butterwort is rare along the lower section of Weaver Creek. Its habitat along the stream consists of mucky soil in areas with a significant amount of sun due to openings in the canopy. FNAI (2007) lists the species as G3G4, S3 and FNAI endangered in Florida. 41 Utricularia cornuta Michx. ? horned bladderwort Holt 226 Horned bladderwort is rare along the lower section of Weaver Creek. It grows along the broad area of saturated soil on either side of the creek. Typically, it is found growing near the edge of the stream. Utricularia gibba L. ? humped bladderwort Holt 186 Humped bladderwort is rare along Weaver Creek and grows in the broad area of saturated soil along the creek. The few specimens are located in areas of greater sunlight caused by openings in the canopy. FAMILY ASTERACEAE Balduina uniflora Nutt. ? oneflower honeycombhead Holt 243 Oneflower honeycombhead is occasional along the lower portions of Weaver Creek. It is found in saturated soil alongside the creek. Bidens laevis (L.) B.S.P. ? smooth beggarticks Holt 168 Smooth beggarticks is rare along the lower portion of Weaver Creek. It is found growing in the broad expanse of saturated soil alongside the creek near the edge of the forested area. Compilation of Previously Reported Steephead Species In addition to my personal observations, I have compiled comprehensive lists of species reported in the literature to occur in steepheads. Plant species found within steepheads are listed in Tables 3 and 4. The reported floristic composition of the uplands surrounding steepheads can be found in Table 5. In comparison to previous studies, 68 species found during this study had not been previously reported from steepheads. The 42 majority of these unreported species (52 species) are found exclusively downstream at Weaver Creek. A list of steephead fauna was also compiled (Table 6). There are extensive lists of herpetofauna and invertebrate families provided by Enge (2005) and Entrekin et al. (1999), respectively. Please refer to these papers for further detail regarding herpetofauna and invertebrates of steepheads. 43 DISCUSSION As a whole, Weaver Creek appears to have a high degree of species and generic diversity compared to other wetland habitats. Much of this diversity is due to the distinction between the floristic communities found along the stream ? one upstream and the other downstream. The change from upstream to downstream is somewhat gradual; there is no distinct line at which the two sections can be delineated. Some species were located throughout the study site, but the majority of species were unique to one section or the other. The two sections consist of very different habitat, a fact which helps to explain the low degree of floristic similarity between the communities. Upstream, the wetland is more heavily shaded due in large part to the slope vegetation. Downstream, the slope is very gradual, so no additional overstory is created by the slope vegetation. There, only one canopy layer provides shade, creating a habitat with greater light intensity which allows for the growth of many of the small herbaceous species found exclusively downstream. These herbaceous species, nearly half of which are rare or infrequent at the study site, increase the richness and diversity (both species and generic) of the downstream section. However, because the majority of individuals downstream are one of two species, either Chamaecyparis thyoides or Taxodium ascendens, the evenness of the downstream section is slightly less than that of the upstream section. Even though evenness is one aspect of diversity, the difference in evenness between the two stream sections does not seem to be great enough to have 44 much of an effect on the diversity of each section. On the steep slope upstream, numerous canopy layers exist and shade the wetland to a greater extent, limiting the growth of a wide variety of small herbaceous species (and, thus, decreasing richness). Species and generic evenness, however, is slightly greater upstream because there are a greater number of dominant species/genera, only about one-fifth of which are rare or infrequent. Furthermore, the greater water depth downstream allows for the growth of floating vegetation islands, which occur along the edges of the stream. These islands are dominated primarily by Dulichium arundinaceum, Eleocharis elongata, Eriocaulon decangulare, Orontium aquaticum, and Xyris spp. They are relatively large in size, reaching up to approximately 8-10 m in length and 2-3 m wide. Upstream, the minimal water depth and extensive shading seem to have prohibited the formation of vegetation islands. Finally, the soils upstream and downstream are drastically different, as described previously. Loose, sandy, well-drained soil composes the majority of the substrate upstream. Downstream soils contain a large amount of decomposing material and are very mucky with a noticeably finer texture. These soils remain saturated year-round. Upstream soils are moist year-round but do not hold as great a volume of water as downstream soils. As a result of these variations in habitat, each section of the creek is characterized by different dominant species. Near the head of Weaver Creek, the wetland is dominated by Cyrilla racemiflora, Ilex coriacea, Ilex opaca, Illicium floridanum, Itea virginica, Lyonia lucida, Magnolia grandiflora, Magnolia virginiana, Myrica cerifera, and 45 Viburnum nudum. The foot of the stream is dominated primarily by Chamaecyparis thyoides and Taxodium ascendens. Both sections of the creek contain an abundance of Pinus elliottii. The drastic difference in dominant vegetation and the low degree of community similarity strongly support the classification of these two habitats of Weaver Creek as highly distinct. There are a number of notable species found in Weaver Creek steephead that have not been previously mentioned in the literature. In total, 67% of the vascular plant species found in this study were not noted by previous authors. The majority of these newly reported steephead species (76%) reside exclusively in the downstream portion. Most of the previous authors have focused heavily on slope vegetation and, to a lesser extent, wetland area near the head of a steephead creek. Focusing on both the wetland habitat of the steephead and the downstream section of Weaver Creek led to this increase in diversity recorded. Because the downstream habitat is drastically different, it is not a surprise that many species have not been accounted for in previous studies. Although it may not have been considered typical steephead habitat in previous studies, it is useful to include the downstream section in steephead studies in order to gain a broader understanding of the entire stream system. Including the vegetation near the foot of the stream adds a new dimension to the floristics of steephead wetlands and adds 72% of the overall species richness. Weaver Creek appears to have a relatively high degree of species richness, evenness and diversity when compared to other wetlands of similar size. Other wetland systems that have a similar hydrologic regime (constant water levels) appear to have species richness values comparable to those of Weaver Creek. For instance, herbaceous 46 plant species richness of pine savannas in southern Louisiana was similar to that of Weaver Creek wetland (140 species in 1 km 2 ) (Keddy et al. 2006). In this study I found 102 species in 0.55 km 2 . However, there have been studies in other wetland types that report greater species richness than was found at Weaver Creek. For instance, Kirkman and Sharitz (1994) reported the presence of 105 species in only 0.025 km 2 in Carolina bay wetlands in South Carolina. However, other research on Carolina bays has shown a lower species richness and/or diversity compared to Weaver Creek (Tyndall et al. 1990, Mulhouse et al. 2005). In the case of Carolina bays in particular, greater variation in hydroperiod (among bays) could be the underlying cause of this divergence in species richness/diversity between studies. Carolina bays have a wide range of sizes, which can affect their hydroperiod since water levels are heavily dependent upon rainfall and rates of evapotranspiration (Kirkman and Sharitz 1994). Differences in hydroperiod would create significant differences in floristic composition. Regarding the differences in richness/diversity between Carolina bays and Weaver Creek steephead, it is important to note the drastic difference in hydrologic regime between these two wetland types. As mentioned above, most Carolina bays do not maintain a constant water level. Steephead water levels remain fairly constant year round due to seepage from the sandhill at the head of the stream. This factor most likely plays a significant role in determining the richness and diversity of the systems. It may be that only Carolina bays with nearly constant water levels approach the diversity of steephead wetlands. Wetlands that are alluvial throughout and in which water levels are not constant appear to have richness levels below those of Weaver creek. Brandt et al. (2003) found a 47 species richness value of 19 over an area of 0.2 km 2 and an evenness of 0.68 (using the Jaccard Index) in tree islands in Everglades National Park, FL. Although their study area was slightly smaller than the wetland of Weaver Creek, the authors determined (prior to the study) that the area sampled was sufficiently large to provide an accurate estimate of species richness. In a study of bottomland hardwood-Pinus taeda forest only 71 vascular plant species were recorded over an area of 9 km 2 (Grell et al. 2005). Diversity of this bottomland forest (mean H? = 1.68) was also much lower than that of Weaver Creek (Grell et al. 2005). Tidal wetlands also appear to be much less species rich than Weaver Creek. Morzaria-Luna et al. (2004) found only 13 species in a salt marsh within a study area of 0.54 km 2 (nearly the size of the wetland of Weaver Creek). The intertidal zone of Boston Harbor Islands National Park contained only 15 species in an area approximately 3.66 km 2 (Bell et al. 2005). It is not surprising that tidal wetlands have a lower species richness considering the water level fluctuation and salinity of these habitats. Comparisons in this study were based only on a survey of one steephead wetland. Investigation of a large number of steepheads would be extremely beneficial. There would likely be a great number of species found in other steephead wetlands that were not found in this study. Other steepheads contain habitat types (within the steephead wetland) that are not present at Weaver Creek. For instance, close to the head of a few steephead streams there are open marshy areas which contain islands of herbaceous species, many of which are not present at Weaver Creek (personal observation). The upstream area of Weaver Creek does not contain the large openings necessary for such islands to develop. Although Weaver Creek is a fairly typical, undisturbed steephead, 48 there is inherent variation in floristic composition among steephead habitats (despite relatively consistent abiotic factors) regarding floristic composition. More intensive field surveys among a number of steepheads would provide a much more complete picture of the true diversity of steephead wetlands. Future research should be aimed at faunal diversity in steepheads, as well. Herpetological fauna has been the most extensively studied thus far, but the use of steepheads as refugia by rare herpetofauna should be investigated further because of obvious conservation implications. Invertebrate diversity research would also be beneficial since very little work has been completed on this topic. The array of invertebrate taxa found in steepheads is almost entirely unknown. Emphasis on wetland research has grown over the past few decades, but steepheads remain relatively unknown even within the scientific community. The variety of plant species found at Weaver Creek and the fact that two very distinct communities are found over a relatively small area support the idea that steepheads are indeed unique wetland systems and worthy of scientific focus. Many steephead wetlands are unaffected by human development because of their topography and/or location. However, human development will eventually put pressure on steepheads as well. Thus, concentrating research on steepheads while they are in a relatively pristine state is of great importance. Cataloguing the unique aspects of these habitats may ultimately lead to their preservation. Because of high diversity, community distinction, and their importance as paleorefugia, preservation of these habitats is critical in helping to protect a large number of species. 49 LITERATURE CITED Aliscioni, S.S., L.M. Giussani, F.O. Zuloaga, and E.A. Kellogg. 2003. A molecular phylogeny of Panicum (Poaceae: Paniceae): tests of monophyly and phylogenetic placement within the Panicoideae. American Journal of Botany 90: 796-821. Ambrose, J.D. 1985. Lophiola, familial affinity with the Liliaceae. Taxon 34: 149-150. Beal, E.O. 1956. Taxonomic revision of the genus Nuphar Sm. of North America and Europe. Journal of the Elisha Mitchell Scientific Society 72: 317-346. Bell, R., R. Buchsbaum, C. Roman, and M. Chandler. 2005. 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In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 12+ vols. New York and Oxford. Vol. 3. Wolfe, S.H., J.A. Reidenauer, and D.B. Means. 1988. An ecological characterization of the Florida Panhandle. U.S. Fish and Wildlife Service Biological Report 88(12): 1-277. Wunderlin, R. P., and B. F. Hansen. 2004. Atlas of Florida Vascular Plants http://www.plantatlas.usf.edu/ [S. M. Landry and K. N. Campbell (application development), Florida Center for Community Design and Research.] Institute for Systematic Botany, University of South Florida, Tampa. 56 Figure 1. Map of the western Florida panhandle with the location of Weaver Creek noted. Coordinates: 30?30?27.84? N, 86?54?46.95? W. Weaver Creek 57 Figure 2. Topographic map of Weaver Creek showing point of origin and point of entry into Weaver River. Head of creek Foot of creek N 58 Figure 3. Topographic map of Weaver Creek illustrating the difference in width between upstream and downstream sections. The point of delineation is only approximate due to the fact that the stream becomes gradually wider downstream. Approximate point of delineation between upstream and downstream sections. N 59 Figure 4. Topographic map of Buck Pond collection site. The powerline along which the stream was sampled is shown as a dashed line with dots interspersed. The point at which the powerline crosses Double Head Branch was the center of the sampling area. N 60 Table 1. Species richness (with number of species unique to each section in parentheses), evenness and diversity (the Shannon Index) of both the upstream and downstream sections of Weaver Creek as well as the entire length of the creek. Community similarity (the Jaccard Index) between upstream and downstream sections is also given. The last column is included to illustrate the numbers of species that occurred in rare (0-5) or infrequent (6-30) abundance categories with percentages of total species per section in parentheses. Richness (S) Evenness (J?) Diversity (H?) Similarity (JI) Rare/ Infrequent Species Upstream 41 (29) 0.88 3.28 9 (22%) Downstream 73 (61) 0.84 3.61 13.33% 35 (48%) Entire Creek 102 0.86 3.98 -- 44 (43%) 61 Table 2. Taxonomic (generic) richness (with number of genera unique to each site in parentheses), evenness and diversity (the Shannon Index) of upstream and downstream portions along with the entire length of the creek. Richness Evenness (J?) Diversity (H?) Upstream 34 (25) 0.97 3.43 Downstream 43 (34) 0.93 3.49 Entire Creek 67 0.95 3.98 62 Table 3: Summary of vascular plant species reported to occur in steepheads. The table is organized by position within the steephead (upper slope, middle to lower slope, and wetland). Source information is provided for each species as well. FAMILY UPER SLOPE MID/LOWER SLOPE WETLAND Aceraceae Acer rubrum (Clewel 1981) Acer rubrum (Enge 198) Apiaceae Hydrocotyle umbelata (Enge 198) Aquifoliaceae Ilex opaca (Enge 198, Kwit et al. 198) Ilex vomitoria (Enge 198) Ilex coriacea (Enge 198, Kwit et al. 198) Ilexopaca (Enge 198, Kwit et al. 198, Eglin AFB Stephead Monitoring Program 204) Ilex vomitoria (Enge 198) Ilex coriacea (Enge 198) Ilex opaca (Enge 198) Araceae Orontium aquaticum (SAIC 206) Peltandra sagitifolia (Chafin et al. 197, SAIC 206) Aristolochiaceae Hexastylis arifolia (Wolfe et al. 198) Hexastylis arifolia (U.S. Fish and Wildlife Service 207) Betulaceae Ostrya virginiana (Enge 198) Carpinus caroliniana (Enge 198) Ostrya virginiana (Wolfe et al. 198, Enge 198, Kwit et al. 198) Blechnaceae Wodwardia virginica (Enge 198) Caprifoliaceae Viburnum dentatum (Enge 198) Clethraceae Clethra alnifolia (Enge 198) Cornaceae Cornus florida (Clewel 1981, Enge 198) Cornus florida (Enge 198) Nysa sylvatica (Enge 198) Nysa biflora (Means 1975, Clewel 1981) Nysa sylvatica (Enge 198) Cupresaceae Juniperus silicicola (Enge 198) Chamaecyparis thyoides (Chafin et al. 197) Cyperaceae Eleocharis sp. (SAIC 206) Rhynchospora crinipes (Chafin et al. 197) 63 FAMILY UPER SLOPE MID/LOWER SLOPE WETLAND Cyrilaceae Cliftonia monophyla (Clewel 1981) Cyrila racemiflora (Clewel 1981, Enge 198) Droseraceae Drosera intermedia (SAIC 206, U.S. Fish and Wildlife Service 207) Dryopteridaceae Onoclea sensibilis (Enge 198) Ebenaceae Diospyros virginiana (Clewel 1981) Ericaceae Kalmia latifolia (Kwit et al. 198) Oxydendrum arboreum (Kwit et al. 198) Vacinium arboreum (Clewel 1981, Enge 198, Kwit et al. 198) Kalmia latifolia (Wolfe et al. 198, Enge 198, Kwit et al. 198, Eglin AFB Stephead Monitoring Plan 204) Lyonia lucida (Wolfe et al. 198, Eglin AFB Stephead Monitoring Plan 204) Oxydendrum arboreum (Clewel 1981, Kwit et al. 198) Rhododendron austrinum (Wolfe et al. 198) Vacinium arboreum (Clewel 1981, Wolfe et al. 198, Enge 198, Eglin AFB Stephead Monitoring Plan 204) Kalmia latifolia (Enge 198, U.S. Fish and Wildlife Service 207) Lyonia lucida (Enge 198) Oxydendrum arboreum (Clewel 1981) Rhododendron austrinum (U.S. Fish and Wildlife Service 207) Euphorbiaceae Sebastiana fruticosa (Enge 198) Fabaceae Cercis canadensis (Enge 198) Cercis canadensis (Enge 198) 64 FAMILY UPER SLOPE MID/LOWER SLOPE WETLAND Fagaceae Fagus grandifolia (Kwit et al. 198) Quercus alba (Clewel 1981, Kwit et al. 198) Quercus hemisphaerica (Clewel 1981, Wolfe et al. 198, Enge 198, Kwit et al. 198, Eglin AFB Stephead Monitoring Plan 204) Quercus incana (Enge 198) Quercus laevis (Clewel 1981) Quercus laurifolia (Enge 198) Quercus nigra (Wolfe et al. 198, Enge 198, Eglin AFB Stephead Monitoring Plan 204) Quercus stelata (Clewel 1981) Quercus virginiana (Clewel 1981) Fagus grandifolia (Clewel 1981, Wolfe et al. 198, Enge 198, Kwit et al. 198, Eglin AFB Stephead Monitoring Plan 204) Quercus alba (Clewel 1981, Wolfe et al. 198, Enge 198, Kwit et al. 198) Quercushemisphaerica (Clewel 1981, Enge 198, Kwit et al. 198) Quercus laurifolia (Enge 198) Quercus michauxi (Wolfe et al. 198, Enge 198) Quercus nigra (Wolfe et al. 198) Hamamelidaceae Hamamelis virginiana (Enge 198) Liquidambar styraciflua (Enge 198) Liquidambar styraciflua (Enge 198) Iliciaceae Ilicium floridanum (Enge 198, Kwit et al. 198) Ilicium floridanum (Means 1975, Clewel 1981, Wolfe et al. 198, Enge 198, Eglin AFB Stephead Monitoring Plan 204, U.S. Fish and Wildlife Service 207) Juglandaceae Carya glabra (Clewel 1981, Wolfe et al. 198, Enge 198) Carya tomentosa (Clewel 1981, Wolfe et al. 198, Enge 198, Eglin AFB Stephead Monitoring Plan 204) Carya palida (Clewel 1981) Caryatomentosa (Clewel 1981) Lauraceae Persea borbonia (Clewel 1981, Enge 198) Persea borbonia (Clewel 1981, Enge 198) Liliaceae Lilium iridolae (SAIC 206) 65 FAMILY UPER SLOPE MID/LOWER SLOPE WETLAND Magnoliaceae Magnolia grandiflora (Kwit et al. 198) Magnolia ashei (Wolfe et al. 198) Magnolia grandiflora (Clewel 1981, Wolfe et al. 198, Enge 198, Kwit et al. 198, Eglin AFB Stephead Monitoring Plan 204) Magnolia pyramidata (Wolfe et al. 198) Magnolia virginiana (Enge 198) Liriodendron tulipifera (Means 1975) Magnolia virginiana (eans 1975, Clewel 1981, Wolfe et al. 198, Enge 198, Eglin AFB Stephead MonitoringPlan204) Myricaceae Myrica cerifera (Enge 198) Nymphaeaceae Nuphar lutea subsp. ulvacea (Chafin et al. 197, SAIC 206) Oleaceae Osmanthus americanus (Clewel 1981, Enge 198, Kwit et al. 198) Osmanthus americanus (Clewel 1981, Enge 198, Kwit et al. 198) Osmanthus americanus (Clewel 1981) Pinaceae Pinus glabra (Clewel 1981, Kwit et al. 198) Pinus glabra (Clewel 1981, Wolfe et al. 198, Enge 198, Kwit et al. 198) Pinus glabra (Enge 198) Potamogetonaceae Potamogeton sp. (SAIC 206) Ranunculaceae Xanthorhiza simplicisima (U.S. Fish and Wildlife Service 207) Rosaceae Amelanchier arborea (Clewel 1981) Crataegus marshali (Enge 198) Prunus americana (Enge 198) Prunus caroliniana (Enge 198) Saraceniaceae Saracenia leucophyla (Chafin et al. 197, SAIC 206) Saracenia rubra (Chafin et al. 197, U.S. Fish and Wildlife Service 207, SAIC 206) Saururaceae Saururus cernus (Enge 198) Saururus cernus (Enge 198) Scrophulariaceae Bacopa monieri (Enge 198) Macranthera flamea (U.S. Fish and Wildlife Service 207) 66 FAMILY UPER SLOPE MID/LOWER SLOPE WETLAND Stemonaceae Cromia pauciflora (Wolfe et al. 198) Symplocaceae Symplocos tinctoria (Enge 198) Symplocos tinctoria (Enge 198) Taxaceae Taxus floridana (Wolfe et al. 198, Enge 198) Toreya taxifolia (Wolfe et al. 198) Theaceae Stewartia malacodendron (Wolfe et al. 198) Tiliaceae Tilia heterophyla (Clewel 1981, Enge 198) Tilia heterophyla (Clewel 1981) Verbenaceae Calicarpa americana (Enge 198) Vitaceae Vitis rotundifolia (Enge 198) Vitis rotundifolia (Enge 198) 67 Table 4: Plant species reported from steepheads with no specific location information. Some are repeated from Table 1 if listed by a different source than in Table 3. Reference information is provided for each species. FAMILY SPECIES Anacardiaceae Toxicodendron radicans (Clewel 1981, Enge 198) Anonaceae Asimina parviflora (Clewel 1981) Ilex coriacea (Clewel 1981) Aquifoliaceae Ilex opaca (Clewel 1981) Araceae Colocasia esculenta (Enge 198) Araliaceae Aralia spinosa (Clewel 1981) Arecaceae Sabal minor (Enge 198) Aristolochia serpentaria (Clewel 1981) Aristolochiaceae Hexastylis arifolia (Clewel 1981, Means 1985) Ascelpiadaceae Matalea alabamensis (Eglin AFB Stephead Monitoring Program 204) Betulaceae Ostrya virginiana (Clewel 1981) Bignoniaceae Bignonia capreolata (Clewel 1981) Blechnaceae Wodwardia areolata (Enge 198) Bromeliaceae Tilandsia usneoides (Clewel 1981) Sambucus canadensis (Enge 198) Caprifoliaceae Viburnum nudum (Clewel 1981, Enge 198) Celastraceae Euonymus americanus (Clewel 1981) Cornaceae Cornus foemina (Enge 198) Cupresaceae Juniperus silicicola (Clewel 1981) Cyperaceae Carex baltzeli (Eglin AFB Stephead Monitoring Program 204) Dioscoreaceae Dioscorea quaternata (Clewel 1981) Athyrium filix-femina (Enge 198) Dryopteris ludoviciana (Enge 198) Dryopteridaceae Polystichum acrostichoides Epigaea repens (Means 1985) Leucothoe racemosa (Clewel 1981) Oxydendrum arboreum (Enge 198) Rhododendron austrinum (Eglin AFB Stephead Monitoring Program 204) Ericaceae Vacinium fuscatum (Clewel 1981) Sapium sebiferum (Eglin AFB Stephead Monitoring Plan 204) Euphorbiaceae Sebastiana fruticosa (Clewel 1981) Fabaceae Cercis canadensis (Clewel 1981) Fagaceae Quercus arkansana (Eglin AFB Stephead Monitoring Program 204) Grosulariaceae Itea virginica (Clewel 1981, Enge 198) Hydrangeaceae Hydrangea arborescens (Means 1985) Juglandaceae Carya ovata (Entrekin et al. 199) 68 FAMILY SPECIES Calamintha dentata (Clewel 1981) Lamiaceae Conradina glabra (Clewel 1981) Lauraceae Sasafras albidum (Clewel 1981) Lilium iridolae (Means 191) Lilium superbum (Clewel 1981) Liliaceae Trilium underwodi (Enge 198) Loganiaceae Gelsemium sempervirens (Clewel 1981) Lygodiaceae Lygodium japonicum (Eglin AFB Stephead Monitoring Plan 204) Liriodendron tulipifera (Enge 198) Magnolia ashei (Eglin AFB Stephead Monitoring Program 204) Magnoliaceae Magnolia pyramidata (Enge 198, (Eglin AFB Stephead Monitoring Program 204) Oleaceae Chionanthus virginicus (Clewel 1981) Orchidaceae Godyera pubescens (Means 1985) Osmundaceae Osmunda cinamomea (Enge 198) Pinaceae Pinus taeda Poaceae Arundinaria gigantea (Enge 198) Actaea pachypoda (Means 1985) Ranunculaceae Clematis reticulata (Clewel 1981) Rosaceae Crataegus uniflora (Clewel 1981) Rubiaceae Mitchela repens (Clewel 1981, Enge 198) Sapotaceae Bumelia lanuginosa (Clewel 1981) Smilax sp. (Enge 198) Smilax bona-nox (Clewel 1981) Smilacaceae Smilax pumila (Clewel 1981) Staphyleaceae Staphylea trifolia (Means 1985) Stemonaceae Cromia pauciflora (Means 1985) Halesia diptera (Clewel 1981) Styracaceae Styrax grandifolia (Clewel 1981) Symplocaceae Symplocos tinctoria (Clewel 1981) Taxus floridana (Clewel1981, Means 1985) Taxaceae Toreya taxifolia (Means 1985) Theaceae Stewartia malacodendron (Eglin AFB Stephead Monitoring Program 2004) Celtis tenuifolia (Clewel 1981) Ulmaceae Ulmus floridana (Enge 198) Verbenaceae Calicarpa americana (Clewel 1981) Violaceae Viola sp. (Enge 198) Parthenocisus quinquefolia (Enge 198) Vitaceae Vitis rotundifolia (Enge 198) 69 Table 5: Vascular plant species reportedly located in uplands (sandhills) surrounding steepheads. Reference information is provided as well for each species. FAMILY SPECIES Agavaceae Yuca filamentosa (Enge 198) Chrysobalanaceae Licania michauxi (Enge 198) Vacinium arboreum (Enge 198) Ericaceae Vacinium yrsinites (Enge 198) Loganiaceae Gelsemium sempervirens (Enge 198) Quercus geminata (Enge 198) Quercus incana (Wolfe et al. 198, Enge 198, Eglin AFB Stephead Monitoring Plan 204) Quercus laevis (Wolfe et al. 198, Enge 198, Eglin AFB Stephead Monitoring Plan 204) Quercus laurifolia (Enge 198) Quercus margareta (Enge 198) Quercus marilandica (Wolfe et al. 198) Fagaceae Quercus virginiana (olfe et al. 198, Enge 198) Pinus clausa (Enge 198) Pinus elioti (Enge 198) Pinaceae Pinus palustris (Wolfe et al. 198, Enge 198, Eglin AFB Stephead Monitoring Plan 204) Smilacaceae Smilax sp. (Enge 198) 70 Table 6: Fauna reported from steepheads. Reference information is provided for each species. FAMILY SPECIES Invertebrates Acrididae Melanoplus apalachicolae (Chafin et al. 197) Baetisca becki (Florida Fish and Wildlife Conservation Comision 204) Baetisca laurentina (Florida Fish and Wildlife Conservation Comision 204) Baetiscidae Baetisca rogersi (Florida Fish and ildlife Conservation Comision 204) Behningidae Dolania americana (Chafin et al. 197) Cambarus pyronotus (Florida Fish and Wildlife Conservation Comision 204, Florida Fish and Wildlife Conservation Comision 205) Procambarus rathbunae (Florida Fish and Wildlife Conservation Comision 205) Cambaridae Procambarus rogersi expletus (Florida Fish and Wildlife Conservation Comision 205) Cordulegastridae Cordulegaster sayi (Florida Fish and Wildlife Conservation Comision 205) Ctenizidae Cyclocosmia toreya (Chafin et al. 197) Gyrinidae Spanglerogyrus albiventris (Epler 205) Leptoceridae Oecetis parva (Florida Fish and Wildlife Conservation Comision 205) Lestidae Lestes inaequalis (Florida Fish and Wildlife Conservation Comision 205) Sericostomatidae Agarodes ziczac (Florida Fish and Wildlife Conservation Comision 205) Fish Pteronotropis hypselopterus (FNAI 190) Pteronotropis welaka (Chafin et al. 197) Semotilus atromaculatus (Wolfe et al. 198, FNAI 190) Cyprinidae Semotilus thoreauianus (Florida Fish and Wildlife Conservation Comision 204) Ictaluridae Noturus leptacanthus (FNAI 190) Etheostoma edwini (FNAI 190) Etheostoma okalosae (Means 191, Chafin et al. 197, Eglin AFB Stephead Monitoring Program 204, Florida Fish and Wildlife Conservation Comision 204, Florida Fish and Wildlife Conservation Comision 205, SAIC 206) Etheostoma parvipine (Florida Fish and Wildlife Conservation Comision 204, Florida Fish and Wildlife Conservation Comision 205) Percidae Percina nigrofasciata (FNAI 190) Amphibians Amphiumidae Amphiuma pholeter (Means 197, Florida Fish and Wildlife Conservation Comision 204, Florida Fish and Wildlife Conservation Comision 205) Hylidae Hyla andersoni (Means and Longden 1976, Chafin et al. 197, Florida Fish and Wildlife Conservation Comision 204, Florida Fish and Wildlife Conservation Comision 205, U.S. Fish and Wildlife Service 207) 71 FAMILY SPECIES Desmognathus apalachicolae (Wolfe et al. 198, Means 191, Means 200, Florida Fish and Wildlife Conservation Comision 204, Florida Fish and Wildlife Conservation Comision 205) Desmognathus auriculatus (Means 1974, Means 1975, Wolfe et al. 198, FNAI 190, Florida Fish and Wildlife Conservation Comision 204, Florida Fish and Wildlife Conservation Comision 205) Desmognathus cf. conanti (Florida Fish and Wildlife Conservation Comision 204, Florida Fish and Wildlife Conservation Comision 205) Desmognathus fuscus (Means 1974) Desmognathus fuscus conanti (Means 1975, Wolfe et al. 198, Means 200) Desmognathus monticola (Means 1974, Florida Fish and Wildlife Conservation Comision 204, Florida Fish and Wildlife Conservation Comision 205) Eurycea bislineata (Means 1975, Wolfe et al. 198, FNAI 190) Eurycea chamberlaini (Florida Fish and Wildlife Conservation Comision 204, Florida Fish and Wildlife Conservation Comision 205) Eurycea cirigera (Means 200) Eurycea cf. quadridigitata (Florida Fish and Wildlife Conservation Comision 204, Florida Fish and Wildlife Conservation Comision 205) Hemidactylium scutatum (Florida Fish and Wildlife Conservation Comision 204, Florida Fish and Wildlife Conservation Comision 205) Pseudotriton montanus (FNAI 190) Pseudotriton ruber (Means 1975, Wolfe et al. 198, Means 200) Plethodontidae Pseudotriton ruber subsp. vioscai (FNAI 190) Proteidae Necturus alabamensis (FNAI 190) Rana clamitans subsp. clamitans (FNAI 190) Ranidae Rana okalosae (Wolfe et al. 198, Chafin et al. 197, Means 191, Eglin AFB Stephead Monitoring Program 204, Florida Fish and Wildlife Conservation Comision 204, Florida Fish and Wildlife Conservation Comision 205, U.S. Fish and Wildlife Service 206, U.S. Fish and Wildlife Service 207) Reptiles Chelydridae Macroclemys temincki (Chafin et al. 197) Farancia erytrograma (FNAI 190) Nerodia taxispilota (FNAI 190) Colubridae Storeria ocipitomaculata (FNAI 190) Kinosternidae Sternotherus minor (FNAI 190) Scincidae Eumeces anthracinus (Chafin et al. 197) Viperidae Agkistrodon contortrix contortrix (Chafin et al. 197) Birds Acipitridae Elanoides forficatus (Florida Fish and Wildlife Conservation Comision 204, Florida Fish and Wildlife Conservation Comision 205) 72 FAMILY SPECIES Butorides virescens (Florida Fish and Wildlife Conservation Comision 204) Ardeidae Egreta caerulea (Florida Fish and ildlife Conservation Comision 204, Florida Fish and Wildlife Conservation Comision 205) Parulidae Seiurus motacila (Florida Fish and Wildlife Conservation Comision 204, Florida Fish and Wildlife Conservation Comision 205) Mamals Eumops glaucinus (Florida Fish and Wildlife 204) Molosidae Nyctinomops macrotis (Florida Fish and Wildlife Conservation Comision 204) Mustelidae Lutra canadensis lataxina (Florida Fish and Wildlife Conservation Comision 204, Florida Fish and Wildlife Conservation Comision 205) Corynorhinus rafinesqui (Florida Fish and Wildlife Conservation Comision 204, Florida Fish and Wildlife Conservation Comision 205) Eptesicus fuscus (Florida Fish and Wildlife Conservation Comision 204, Florida Fish and Wildlife Conservation Comision 205) Lasiurus borealis (Florida Fish and Wildlife Conservation Comision 204, Florida Fish and Wildlife Conservation Comision 205) Lasiurus cinereus (Florida Fish and Wildlife Conservation Comision 204, Florida Fish and Wildlife Conservation Comision 205) Lasiurus intermedius (Florida Fish and Wildlife Conservation Comision 204, Florida Fish and Wildlife Conservation Comision 205) Lasiurus seminolus (Florida Fish and Wildlife Conservation Comision 204, Florida Fish and Wildlife Conservation Comision 205) Myotis austroriparius (Florida Fish and Wildlife Conservation Comision 204, Florida Fish and Wildlife Conservation Comision 205) Myotis grisescens (Florida Fish and Wildlife Conservation Comision 204, Florida Fish and Wildlife Conservation Comision 205) Vespertilionidae Pipistrelus subflavus (Florida Fish and Wildlife Conservation Comision 204, Florida Fish and Wildlife Conservation Comision 205)