Floristic Inventory of Tiger Creek Preserve and Saddle Blanket Scrub Preserve, Polk County, Florida

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Floristic Inventory of Tiger Creek Preserve and Saddle Blanket Scrub Preserve, Polk County, Florida
Corogin, Paul T.
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[Gainesville, Fla.]
University of Florida
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1 online resource (107 p.)

Thesis/Dissertation Information

Master's ( M.S.)
Degree Grantor:
University of Florida
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Committee Chair:
Judd, Walter S.
Committee Members:
Gordon, Doria R.
Putz, Francis E.
Hall, David W.
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Subjects / Keywords:
Blankets ( jstor )
Creeks ( jstor )
Floodplains ( jstor )
Highlands ( jstor )
Scrub vegetation ( jstor )
Species ( jstor )
Taxa ( jstor )
Tigers ( jstor )
Vegetation ( jstor )
Wetlands ( jstor )
Botany -- Dissertations, Academic -- UF
blanket, conservancy, conservation, creek, endemic, endemism, florida, floristic, lake, plant, polk, rare, ridge, saddle, scrub, tiger, wales
City of Lake Wales ( local )
Electronic Thesis or Dissertation
bibliography ( marcgt )
theses ( marcgt )
Botany thesis, M.S.


A floristic inventory of vascular plants for The Nature Conservancy?s Tiger Creek Preserve and Saddle Blanket Scrub Preserve, Polk County, Florida was conducted from August 2004 through February 2008. Twelve plant communities were characterized: scrub, scrubby flatwoods, sandhill, xeric hammock, mesic flatwoods, wet flatwoods, bayhead, seepage slope, depression marsh, floodplain wetlands, blackwater stream, and ruderal. The 2308 combined hectares of the Preserves yielded vouchers for 554 taxa, of which 54 are non-native to Florida, 37 are at or near the southern or northern limits of their ranges in Florida, and 24 are new Polk County records. The Preserves are located on the Lake Wales Ridge. This area's unique geological history has contributed to the evolution of a high number of endemic taxa, many of which are rare and of conservation concern. This study found 55 taxa endemic to Florida, of which 27 are restricted to Florida peninsular scrubs or scrub/sandhill ecotonal habitats. Of these 27 taxa, 15 are restricted to Florida's central ridges, especially to the ancient scrubs on the Lake Wales, Lake Henry, and Winter Haven Ridges. Nine of these taxa are restricted to four or fewer counties. Twenty-seven are listed in Florida, and 18 are listed federally, as endangered, threatened, or of management concern. Endemism on Florida's sand ridges is discussed. ( en )
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Thesis (M.S.)--University of Florida, 2008.
Adviser: Judd, Walter S.
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by Paul T. Corogin

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2008 Paul T. Corogin 2


ACKNOWLEDGMENTS My sincere thanks go to Dr. Walter Judd, my supervisory committee chairman. His academic professionalism, his belief in my abilities, and his enthusiastic willingness to share his vast botanical expertise, have gui ded me invaluably in my academic aspirations, as well as in the production of this thesis. I also thank the othe r members of my committee. Dr. Doria Gordons knowledge of Florida habitats and her affiliation with The Nature Conservancy have given me insights concerning my study sites. Dr. David Halls expertise in the Florida flora has been of great assistance as I have puzzled over stacks of unidentified specimens. And the work and teaching of Dr. Francis Jack Putz have re fined my understanding of ecology and natural history, and instilled in me a drive for academic perfection in my work. For their help in using the Universi ty of Florida Herbarium, I thank staff members Kent Perkins and Trudy Lindler. Thanks are also due to Richard Abbo tt, whose insightful pointers on many occasions helped me solve plant identification puzzles. I am grateful as well to the staff of The Nature Conservancys Lake Wales Ridge Program, especially Beatriz Pace-Aldana, Adam Peterson, and Steve Morrison, for their support and assistance throughout this project. I also thank Dr. Terry Lucansky, who guided me through the bureaucratic maze when I came back to academia in mid-life as a post-baccalaureate student. His encouragement did much to propel me toward eventual entrance into graduate school. Finally, I thank my mother for instilling in me the importance of believing in oneself. I thank my father for his generosity, and for be ing a continual inspira tion, showing by example that many of our perceived limitation s are illusory. Most of all, I thank Ananda Devi, my wife her unfailing love and support have smoothed many a bump in the road to this journeys completion. 3


TABLE OF CONTENTS page ACKNOWLEDGMENTS...............................................................................................................3 LIST OF TABLES................................................................................................................. ..........6 LIST OF FIGURES.........................................................................................................................7 ABSTRACT.....................................................................................................................................8 CHAPTER 1 INTRODUCTION................................................................................................................. ...9 The Preserves............................................................................................................................9 Geology and Physiography.....................................................................................................11 Climate........................................................................................................................ ............15 Hydrology and Soils...............................................................................................................17 History....................................................................................................................................20 Pre-Human Florida..........................................................................................................21 Human Florida.................................................................................................................2 3 History of the Preserves...................................................................................................25 2 PLANT COMMUNITIES......................................................................................................33 Upland Communities..............................................................................................................34 Scrub................................................................................................................................36 Scrubby Flatwoods..........................................................................................................42 Sandhill....................................................................................................................... .....43 Xeric Hammock...............................................................................................................46 Mesic Flatwoods..............................................................................................................47 Lowland Communities............................................................................................................ 48 Wet Flatwoods.................................................................................................................48 Bayhead...........................................................................................................................49 Seepage Slope..................................................................................................................50 Depression Marsh............................................................................................................50 Floodplain Wetlands........................................................................................................51 Blackwater Stream...........................................................................................................53 Ruderal............................................................................................................................54 3 FLORISTIC METHODS AND RESULTS............................................................................58 Methods..................................................................................................................................58 Results.....................................................................................................................................59 Taxa of Special Inte rest or Concern................................................................................59 Endemism on Floridas Ancient Sand Ridges.................................................................63 4


4 ANNOTATED LIST OF VASCULAR PLANTS.................................................................73 LITERATURE CITED..................................................................................................................94 BIOGRAPHICAL SKETCH.......................................................................................................107 5


LIST OF TABLES Table page 3-1 Endemic vascular plant species found on Tiger Creek and Saddle Blanket preserves that are restricted to Florida penins ular scrubs or scrub/sandhill ecotones.......................71 3-2 Endemic vascular plant species found on Tiger Creek and Saddle Blanket preserves that are not restricted to peninsular scrubs.........................................................................72 6


LIST OF FIGURES Figure page 1-1 Peninsular Florida, showing the central ridge system and Polk County............................30 1-2 Polk County area, showing the Polk Upla nd, ridges, and locations of Tiger Creek and Saddle Blanket Preserves...................................................................................................31 1-3 Tiger Creek Preser ve acquisition map...............................................................................32 2-1 Vegetation map of Tiger Creek Preserve...........................................................................56 2-2 Vegetation map of Saddle Blanket Scrub Preserve...........................................................57 7


8 Abstract of Thesis Presen ted to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science FLORISTIC INVENTORY OF TIGER CREEK PRESERVE AND SADDLE BLANKET SCRUB PRESERVE POLK COUNTY, FLORIDA By Paul T. Corogin May 2008 Chair: Walter S. Judd Major: Botany A floristic inventory of vascular plants for The Nature Conservancys Tiger Creek Preserve and Saddle Blanket Scrub Preserve, Polk C ounty, Florida was conducted from August 2004 through February 2008. Twelve plant communities were characterized: scrub, scrubby flatwoods, sandhill, xeric hammo ck, mesic flatwoods, wet flat woods, bayhead, seepage slope, depression marsh, floodplain wetl ands, blackwater stream, and ruderal. The 2308 combined hectares of the Preserves yielded vouchers for 541 taxa, of which 51 are non-native to Florida, 37 are at or near the southern or northern limits of their ranges in Florida, and 24 are new Polk County records. The Preserves are located on the Lake Wales Ridge. This areas unique geological history has contribute d to the evolution of a high nu mber of endemic taxa, many of which are rare and of conservation concern. This study found 55 taxa endemic to Florida, of which 27 are restricted to Florida peninsular scrubs or scrub/sandhill ecotonal habitats. Of these 27 taxa, 15 are restricted to Florid as central ridges, especially to the ancient scrubs on the Lake Wales, Lake Henry, and Winter Have n Ridges. Nine of these taxa ar e restricted to four or fewer counties. Twenty-seven are listed in Florid a, and 18 are listed federally, as endangered, threatened, or of management concern. Endemi sm on Floridas sand ridg es is discussed.


CHAPTER 1 INTRODUCTION The Preserves Tiger Creek Preserve and Saddl e Blanket Scrub Preserve are two conservation properties in Polk County, Florida, owned and operated by Th e Nature Conservancy (TNC). Tiger Creek Preserve is located about eight km southeast of the town of Lake Wales (population 15,000), on the eastern edge of the Lake Wales Ridge (Figures 1-1, 1-2), a sandy upland region of central Florida noted for its unique ecosystems and large number of rare endemic species, both animal and plant (Ward 1979). The area co ntaining the Preserve is bounde d on the north by State Road 60, on the east by Walk-in-Water Road and La ke Weohyakapka, on the south by County Road 630, and on the west by State Road 17. To the west of the Preserve are citrus groves, to the north and east are low-density residential developments interspersed with native vegetation, and to the south is a mosaic of pasture, citrus groves, and residential developmen t. The Preserve was established in 1971 to protect the watershed of Tiger Creek. Since then the Preserve has grown through the piecemeal acquisition of parcels of surrounding land (Figure 1-3), to 1974 hectares (ha), including upland habitats. At its current ex tent, encompassing a blackwater stream and its adjacent uplands, the Preserve represents an ou tstanding remnant of Lake Wales Ridge habitats, harboring 15 species of plants and seven species of animals with Natural Heritage Inventory element rankings of G3 or higher (Chafin 2000; TNC 2006a). Saddle Blanket Scrub Preserve is located in extreme southern Polk County near the Highlands County line, at the western edge of the Lake Wales Ridge, approximately 24 km southwest of Tiger Creek Preserve (Figure 1-2) The northern boundary of the Preserve fronts on Avon Park Cutoff Road approximately 1.6 km west of its intersection with U.S. Highway 27. To the north and east are citrus groves, and to the south and west are moist grassy pasture lands. 9


This Preserve was acquired by TNC in 1988. A 36 ha parcel was added in 1992, bringing the total area to 334 ha. It is prim arily a mosaic of upland habitats including scrub, punctuated by a few depression wetlands. The Preserve features one of the largest, most intact, and biologically diverse tracts of ancient scrub ha bitat remaining on the Lake Wale s Ridge. It harbors 18 of the rare plant species and eight of the rare animal species that are restricted to th ese habitats. This is one of the largest concentra tions of these species now re maining (TNC 2006b, 2006c). Because of their locations on the Lake Wales Ridge, these Preserves are at the center of ongoing conservation efforts. The habitats of the Ri dge and their unique biological denizens have long been under severe threat from human land-use changes; the already naturally fragmented Lake Wales Ridge scrub community has been redu ced to less than 10% of its original extent (Christman and Judd 1990; Huck et al. 1989). The accelerated pace of human encroachment injects an urgency into efforts to understand and preserve these areas. Over the last several decades, conservationists at the federal, state, county, and private levels have risen to the challenge, acquiring land for pr eservation and devising strategi es for studying and protecting what remains of these species and their vanishing habitats (FWCC 2002). One result of these efforts is the Lake Wales Ridge Protected Area s Network, a patchwork of protected lands both public and private, totaling some 34,400 ha th at includes Tiger Creek and Saddle Blanket Preserves (Martin 1998). These two Preserves co mprise the centerpiece of TNCs Lake Wales Ridge Program, whose goal is to pr eserve biodiversity in this region, using the preserves as a platform to promote ecological management prac tices on conservation lands of the Lake Wales Ridge, and as a laboratory for study of the Ridges ecosystems (TNC 2006a). Despite efforts to promote research, prior to this investigatio n, no vouchered floristic inventories had been published for these two Pres erves. This is surprising, in view of the 10


biological importance of this area and the urgency of its preservation. With this in mind, a vouchered floristic inventory of Tiger Creek and Saddle Blanke t Scrub Preserves was conducted over a three-year period, from August 2004 through February 2008. The objective of this study was to catalog the flora and plant communities of these preserves as they now occur, providing a baseline of information for use by current and future researchers a nd managers in their efforts to understand and conserve the biota of these unique ecosystems. In addition, phylogenetic relationships and biogeographical patterns of th e areas endemic species are briefly assessed. Geology and Physiography Florida today presents a seemingly simple tableau a sandy peninsula of low relief, recently emerged from the sea, suggesting a simp le biogeography yet since its emergence from the sea, it has undergone a variet y of geologic processes includi ng numerous changes in size and shape (Webb 1990). To understand its biogeogra phy, therefore, one must understand Floridas geological history. Modern peninsular Florida is the exposed portion of a larger block known as the Florida Platform, measuring some 560 by 720 km in size, extending southeast from continental North America (Scott 2001). The Florida Platform had its beginnings in the Paleozoic era, in the Cambrian period [ca. 600 million years before present (MYBP)], as undersea volcanic rocks attached to the Gondwana land mass near what is now the West African continental margin. Over these igneous basement rocks were depos ited sedimentary layers of Ordovician through Devonian age (ca. 500 350 MYBP). In the Tr iassic period (ca. 240 MYBP), processes of rifting began to separate Africa from North and South America, creating the Atlantic Ocean, and the Florida Platform separated from Africa and became attached to the North American plate (Smith and Lord 1997). 11


By the mid-Jurassic (ca. 180 MYBP), the Fl orida platform had ceased to undergo violent tectonic movements and to this day it remains one of the most stable areas of the earths crust (Cooke 1939). From the Jurassic until at least th e mid-Oligocene (ca. 28 MYBP), the platform underwent a long period of marine deposition of ca rbonate sediments in shallow seas, building up the thick limestone bedrocks characterist ic of Florida (Randazzo 1997; Scott 1997). Rainwater is slightly acidic (Jordan 1984), and these carbonate limestone bedrocks, when above sea level, are subject to dissoluti on when exposed to these acidic su rface waters. This leads either to denudation (general lowering of land surface) or to karst formation. The latter occurs when surface water flows down through cr acks, joints, and softer places in the rock, dissolving the rock, forming cavities, depressions, and sinkholes. Karst topography is widespread in Florida due to this process. Terrestrial vertebrate fo ssils deposited in sinkholes indicate that karst formation had taken place by the late Oligocen e in some areas of peninsular limestone, suggesting this as the earliest emergence of Florida from the sea, during low sea-level stands associated with climatic cooling. The limestone layers formed in the Eocene, Oligocene, and early Miocene (ca. 56-24 MYBP) form the Florid an aquifer. These limestone layers, made porous by karst-forming solution processes, carry huge amounts of groundwater throughout much of the peninsula. (Brown et al. 1990; Upchurch and Randazzo 1997; Webb 1990). By the end of the Oligocene, renewed uplift and erosion in the Appalachian region brought increasing loads of siliclastic sediments washi ng out onto the Florida platform. This deposition predominated from mid-Oligocene to Holocene (recent) times, covering much of the limestone bedrock with sandand clay-cont aining sediments, which form re latively impermeable confining layers over the limestones of the Floridan aquifer (Brown et al. 1990; Scott 1997, 2001; Scott et al. 2001; Webb 1990). These deposits, often lying over older karstic features, have been worked 12


and re-worked by wave action over ages of sea-level fluctuations, and subjected to erosion while above ground, to shape the surficial formations recognized in modern Fl orida (Scott 2001). Polk County lies within the Central Highlands physiographic provin ce of Florida (Cooke 1945). The central and southern part s of Polk County, including th e Lake Wales Ridge, lie within the Central Lake District, an ar ea of karst terrain with sandhill s and numerous lakes and solution basins (Brooks 1982; Brown et al. 1990). Underlying Polk County, and forming the Floridan aquifer, are the Eocene Avon Park and Ocala Group limestones, and the Oligocene Suwannee limestone. Over these are the less permeable si liclastic-containing Miocene sediments of the Hawthorne Group, which include the Arcadia formation, in the southwestern part of Polk County, forming an intermediate aquifer, overlai n by the Peace River formation, which is present over much of the county, forming the upper confini ng layer for the intermediate aquifer. Over this formation are undifferentiated sandy to clay ey sediments forming the unconfined surficial aquifer of Pliocene to Holocene age (Campbell 1986; Tighe 1987). Overlying the entire county are quartz sands from which the surfa ce soils have developed (Tighe 1987). The Lake Wales Ridge, along with the similar Winter Haven and Lake Henry Ridges to its west, are prominent features rising from th e surrounding Polk Upland (Figure 2). The topography of all the ridges is quite irregular due to karstic processes, erosion, and dune building; elevations on the Lake Wales Ridge range from 46 to 93 m above mean sea level (Campbell 1986; White 1958). These ridges are ca pped with sediments once thought to be of Miocene origin (White 1958), but now recognized as sandy sediments of the Cypresshead Formation, deposited when sea level rose dram atically in the Pliocene, inundating much of peninsular Florida (Scott 1997, 2001). White (1970) considered these ridges remnants of a once more extensive upland, preserved as ridges because they sit in pl aces where siliclastic sediments 13


were thicker over underlying limestone, limiting dissolution in those places while surrounding limestone, with thinner overlying layers, diss olved, and subsided. Surrounding the ridges are undifferentiated reworked Cypresshead sediments of Pliocene to Pleist ocene age (Scott 2001). The ridges are oriented parallel to the current Atla ntic coast; in aerial view they exhibit a striped pattern typical of beach ridges, with dunes on the eastern flank of the Lake Wales Ridge, indicating that they we re shaped by wave and wind action along an ancient shoreline (White 1970). The maturity of karst on the ridges (deeper lakes, roughe r and more dissected terrain) compared with that of the surrounding areas suggests that the ridges have been above sea level for a considerable period of time (White 1970). Of several ancient Florida shorelines recognized by geologists (e.g., MacNeil 1950), the most promin ent is the Wicomico, which encircles the Central Highlands and represents a high sea-level stand probably in the early Pleistocene (Webb 1990). A higher and older shoreline, the Okefe nokee, occupies some higher elevations on the ridges, and may date from the late Pliocene (Webb 1990). These high points on the ridges may have once formed a string of islands, or a narrow peninsula, in a shallow Pliocene sea, their flora and fauna evolving in isolation, a possible explanation for the la rge number of endemic species restricted to these ridges (Christman 1988a; Myers 1990). Tiger Creek Preserve lies on th e eastern flank of the Lake Wales Ridge, occupying an area covered by Pleistocene dunes extending to the ea st shore of Lake Weohyakapka. The creek system has cut into these dunes forming a broa d floodplain in the norther n and western portions of the Preserve, while downstream to the east the cr eek is more steeply incised in the dunes as it flows into Lake Weohyakapka. The upland areas of the Preserve are undulating sandhills with numerous depressions, remnants of the Pleistoc ene dune and swale system. The swales are now ephemeral depression wetlands. Elevations with in the Preserve range from 21 to 41 m, the 14


highest point being on the northwe st side of the Preserve wher e it abuts the steep eastern escarpment of the Lake Wales Ridge proper. (Tighe 1987; TNC 2006a). Saddle Blanket Scrub Preserve lies on the western edge of the Lake Wales Ridge. The eastern portion of the Preserve is on the ridge proper, on se diments of the Pliocene-aged Cypresshead formation, while the western pa rt lies in the Polk Upland, on an area of undifferentiated reworked Cypresshead sediments of Pliocene to Pleistoc ene age (Scott et al. 2001). At the surface are quartz sands from which the soils have developed. With the exception of a few depression wetlands, the Preserve is a relatively flat sandy upland, with elevations ranging from 41 m over a broad cen tral area, to 32 m at its s outhwestern edges (Namm 1997). Two small lakes, the Saddle Blanket Lakes, are lo cated at the northern edge of the Preserve. Climate Although peninsular Florida lies at a latitude dominated by dry high-pressure air, its proximity to maritime tropical air masses over the Caribbean Sea and the Gulf of Mexico makes its climate humid and subtropical (Chen and Gerb er 1990). Based on a 29-y ear averaging period (1971-2000) at Mountain Lake near Tiger Creek, the mean annual temperature in Polk County is 22.1 degrees C, and mean annual rainfall is 1212 mm; mean monthly temperatures vary from 16.1 degrees C in January, to 27.3 degrees C in August (NOAA 2005). Frost or freezing temperatures generally occur at least once a year in the Lake Wales Ridge area (Waldron et al. 1984). Central peninsular Florida experiences a di stinct summer rainy season from June through September, and a dry season from November th rough April. The region receives more than 60% of its annual rainfall in the summer (Jordan 1984). Most of this rain comes from thunderstorms that develop along seabreezes through convergence, or inland through convection; both processes cause sun-heated, moisture-laden air to rise, and the moisture condenses to form 15


rainstorms (Winsberg 1990). These storm-genera ting processes result from solar heating of the land surface. Air over land heats and rises faster than air over water, creating low pressure over land, whereupon moist air from over water rush es toward the low pressure, creating the seabreezes (Chen and Gerber 1990). Florida s narrow peninsular shape assures overall good coverage of storm rainfall throughout, as no part of the peninsula is far from water. The distribution of rainfall from thunderstorms, however, is naturally irregular, and unusual rainfall events can cause localized flooding that can affe ct low-lying plant communities (Jordan 1984). Floridas thunderstorm activ ity is the greatest of anywhere in the United States, and it ranks with the worlds other high thundersto rm areas (Winsberg 1990). Th is abundance of thunderstorms creates a vast amount of lightning, which can and does start fires in flammable vegetation. Over evolutionary time, natural fires have been a dom inant ecological disturbance in central Floridas sandy upland ecosystems, greatly influencing plan t species characterist ics and the makeup and distribution of plant communities (Menge s and Kohfeldt 1995; Myers 1990). Tropical weather systems are another significan t source of rainfall in summer and early fall in this region. Between 1852 and 2007, 53 hurricanes have passed within 40 km of the town of Lake Wales (NOAA 2007). Three of these, Char ley, Frances, and Jeanne, occurred in 2004, near the start of this study period. Flooding and wind damage from such events can have pronounced disturbance effects on plant communities. By November, the rainy season has ended in central Florida, and winter rainfall is dependent upon occasional low-pressure systems, and the passage of continental cold fronts, which drop rain in north Florid a but often dissipate over the ce ntral peninsula, making winter drier there than in north Fl orida (Chen and Gerber 1990). In spring, the Bermuda high, a semipermanent subtropical high-pressure ridge, extends westward from the Atlantic Ocean, 16


creating dry sunny conditions that can persist through May. The stre ngth of this high can delay the onset of the rainy season. Drought conditions can occur when a dry spring follows a dry winter (Jordan 1984). Hydrology and Soils The ultimate source of all fresh water in central peninsular Florida is rainfall (Clemens et al. 1984). Some of this water flows as runoff, entering streams and lakes. The rest filters through the soil into the surficial aquifers wher e it can percolate latera lly, entering wetlands and water bodies, or downward, to re charge the deep aquifers (M iller 1997). The Central Lake District, where the limestone bedrocks are ove rlain by deep, permeable sands, discontinuous and leaky confining beds, and numerous deep lakes, is an important recharge area for the Floridan aquifer (Barcelo et al. 1990; Brown et al. 1990). At Tiger Creek and Sa ddle Blanket Preserves, the deeper aquifers do not contribute directly to water in wetlands, streams or ponds; these are fed by runoff and seepage from the surficial (water table) aquifer (Tighe 1987; TNC 2006c). Politically, Tiger Creek and Sa ddle blanket Preserves fall wi thin the Southwest Florida Water Management District (Barcelo et al. 1990). Hydrologically, Tiger Creek is part of the Kissimmee River basin. It drains a 13,786 ha drainage basin situ ated to the west of, and draining into, Lake Weohyakapka, which in turn drains into the Kissimmee River through a series of streams and lakes (Tighe 1987; TNC 2006a). Tighe, who conducted a detailed two-year hydrological study of Tiger Creek, considered it a relatively undisturbed seepage blackwater stream system (1987, p. 85). Its southern branch Patrick Creek, flows nor theastward from its source, Lake Patrick (Lake Leonore on some maps), through a culvert under Murray Road, to its confluence with Tiger Creek. Its floodplain is quite broad except where it flows through some upland areas. The source of the northern br anch is groundwater seepage from surrounding uplands into a bayhead north of the Preserve. For the first 1.6 km of its southward flow, its 17


floodplain is relatively narrow, surrounded by xeri c uplands, after which the floodplain broadens to about 0.8 km in width, remaining broad up to the confluence. From the confluence it flows eastward into Lake Weohyakapka, through sandy uplands where its banks are high and its floodplain narrow. The channels of the creek system are flat and sandy-bottomed, choked in many places with fallen trees. The terrain of th e Preserve outside the creek floodplain is an undulating mosaic of uplands and ephemeral de pression wetlands. Upland soils are deep, permeable sands, while those of depressions a nd flatwoods may be underlain by a hardpan that holds the water table perched, where water can stand for some time following rainfall (Tighe 1987). Some of these wetlands, especially se veral bayheads at the base of sandy slopes, contribute flow to the creeks via small tributary streams. These tributary wetlands have a large water storage capacity, and can discharge water to the creeks at steady ra tes during much of the year (Tighe 1987). Central Florid a receives more than 50% of its total yearly rainfall during the summer (June-September), leaving the water tabl e highest in September, lowest in May. The hydrology of Saddle Blanket Scrub Preserve is similar to that of Tiger Creek, except for the absence of a creek system at Saddle Blanke t. Rainwater runs off or percolates into the permeable upland sands, where it flows laterally to feed several depression wetlands and the small lakes, or downward to recharge the Floridan aquifer. The average level of the water table of this surficial aquifer, 34-37m, corresponds to the ground level of the bayheads and the water level of the small lakes (TNC 2006c). At highe r elevations than the bayheads are some wetprairie depressions whose water tables are pr obably perched above hardpan layers. Surface water in the area of this Preserve, on the wester n side of the Lake Wale s Ridge, drains westward toward the Peace River (Barcelo et al. 1990). 18


Of the various land uses surrounding Tiger Cr eek and Saddle Blanket Preserves, citrus groves have the greatest potential to affect the hydrology due to their enormous consumption of water for irrigation, most of which is drawn fr om deeper aquifers (Tighe 1987; TNC 2006a, 2006c). General widespread declines in regional lake levels in recent decades have been linked to reduced rainfall and groundwater pumpage (Barce lo et al. 1990). Because the confining beds of the surficial aquifer are lea ky in this region, withdrawals from the deeper aquifers can cause local declines in the surficial water table, pot entially threatening the plant communities of the wetland areas of these Preserves. Another land use that occurs n ear Tiger Creek Preserve is sand mining. Hydrological monitoring is ongoing at Tiger Creek in response to concerns about the possible effects on the hydrology of the creek system of prop osed new sand mines near the Preserve (TNC 2006a). Variations in rainfall patterns over the year s have had dramatic effects on the hydrology of Tiger Creek Preserve, and this probably affected the composition of wetland plant communities. The first half of the 20th century was generally wetter than the second half (Tighe 1987). Of interest in this regard ar e some aerial photos from 1922 commissioned by Edward Bok, founder of Historic Bok Sanctuary near Lake Wales (on file at TNC o ffice at Tiger Creek Preserve), showing that a large area of the floodplain at the confluence of Tiger and Patrick Creeks was open grassland at that time, perhaps a sawgrass (Cladium jamaicense) marsh. In 2004 this same area was forested wetland with sparse occurren ces of sawgrass in the understory. The drier period from the 1950s through 1985 had seen wetla nd areas going dry sometimes for decades, with mesic and xeric species invading (Tighe 1987; TNC 2006a). During 1998-2007, rainfall was on the increase, and in 2004-2005, during the period of this floris tic study, three closepassing hurricanes followed by an unusually we t summer left all floodplains and depression 19


wetlands inundated, some for up to 18 months (NOAA 2004, 2005; TNC 2006a). This inundation exceeded the limits of flooding tolerance for many mesic and hydric tree species in the forested creek floodplains, causing a remark able die-off of canopy trees. This was in addition to the considerable number of trees dow ned by hurricane winds. Patterns of rainfall and fire in the decades following this event will dete rmine whether these areas become reforested, or remain more open and marshy. Evidently, hydrol ogic processes have contributed to wholesale changes in wetland community makeup, and time will tell if we are witnessing another such change in the making. Soils of Lake Wales Ridge uplands are deep, we ll-drained sands. At Tiger Creek Preserve, upland soils are mainly of the yellow sand Astatu la series, which typically support sandhill and yellow sand scrub vegetation. Poor ly drained sandy soils of the Smyrna and Myakka series are found in pine flatwoods areas. Basinger muc ky fine sands occur in seasonal depression wetlands, while bayheads and the creek floodplain are dominated by organic soils of Samsula and Hontoon muck series. (TNC 2006a; USDA 1990) At Saddle Blanket, the predominant upland soils are of the Archbold sand series, coarse white well-dr ained sands that support scrub vegetation. In wet prairies and flatwoods areas are poorly-drained Placid and Myakka depressional sands, and in the bayheads is Samsula muck (TNC 2006c; USDA 1990). History The biota of Florida evolved over millions of years through many environmental changes before human beings came on the scene. The arri val of humans ushered in an era of changes that continue at an ever-increasing pa ce to the present day. To set the stage for the snapshot in time that this study represents, I offer the following synopsis of the biotic history of terrestrial Florida, divided into three parts. My account of pre-human Florida is based primarily on Webb (1990), with other references as noted. In summarizing human Florida, I have consulted Milanich (1994, 20


1995) for the pre-European period; for the rest I have drawn upon various authors as cited. For the local history of Tiger Creek and Saddle Bl anket Preserves, I have drawn heavily from anecdotal information collected by Preserve managers. Pre-Human Florida The oldest fossil evidence for a la nd biota in peninsular Florid a dates to the late Oligocene (ca. 25MYBP), consisting of two ve rtebrate fossil sites, one near Gainesville in north Florida, and the other in Hillsborough County east of Tampa. The kinds of animals represented suggest that mesic forest may have been the predominant ha bitat at that time. Vertebrate fossils from the early to middle Miocene (23-11 MYBP) are found in the panhandle and in Polk and Hillsborough Counties; they suggest a tropical climate and a mosaic of mesic forest and open savanna habitat. Middle Mi ocene plant fossils from the Alum Bluff area of the panhandle include a mixture of temperate and tropical speci es. According to Graham (1964), the flora of eastern North America was undergoing a change from tropical to temperate throughout the Tertiary period, and the Alum Bluff flora was about 42 percent tropical by the middle Miocene. In the late Miocene (11-5 MYBP ), two dramatic drops in sea-level occurred, during which Florida reached perhaps its great est land area, and the climate probably was dry. Vertebrate fossils from this period show a great diversity of savanna-adapt ed species. A high sea-level stand in the early Pliocene (5-4 MYBP) redu ced Floridas land area, and corresponds to the deposition of siliclastic sediments like the Cypres shead formation in the central peninsula, which underlies the Lake Wales Ridge. Vertebrate fossils from this time suggest Florida still supported tropical savanna and forest habitats. Florida vertebrate fossils dati ng from the late Pliocene (ca. 2.5 MYBP) record extinctions of Miocene fauna, and give evidence of the wave of immigrant species that had begun to arrive from South America over the newl y-formed isthmus of Panama. Tropical savanna probably predominate d, along with some forested areas. 21


Whether parts of the central peninsula were on ce islands, or were alwa ys a peninsula, has been debated. Based on the existence of endemism in the biota of the central peninsula, many researchers have postulated the existence of a Suwannee Strait cutting across north Florida even as late as the Pliocene. But geologists ( e.g., Hine 1997) believe this seaway had filled in with sediments by the Oligocene. Webb (1990) ar gues that Florida has probably always been a peninsula, and that central Floridas endemism can be explained as easily by the existence of habitat islands as by geographical ones. Fossils from the early Pleistocene (ca. 2 MYBP ) indicate that, especi ally during periods of low sea level, there was likely a continuous corr idor of semi-arid subt ropical savanna habitat connecting Florida with wester n North America. This connection was broken in the midPleistocene, as evidenced by the disappearance of many western taxa from the Florida fossil record, although a number of reptiles, birds, a nd plants have remained as disjunct relics, separated from their western rela tives by wetter habitats of the Mississippi delta region. This period also saw the development of longleaf pine habitats, inferred from certain vertebrate fossils. Numerous oscillations of sea level throughout the Pleistocene co rresponded to cycles of glaciation, and these created various relict shorelines visible today in Florida. Floridas Central Highlands region was above water for much of the Pleistocene, and the Lake Wales Ridge probably for all of it. During low sea-level sta nds the water table droppe d, wetlands disappeared, and the uplands became more xeric than today, supporting a scrublike vege tation that was likely quite extensive. Late Pleistocene pollen reco rds from Lake Annie on the Lake Wales Ridge indicate the presence during the last glacial period of grasslan ds, and a xeric scrub vegetation analogous to todays rosemary scr ub, which is found on the driest dune tops in that area (Watts 1980). At the end of the last glacial period, this pollen record shows changes in the composition 22


of the flora, with pines and mesic and wetland species increas ing, and grasses decreasing, as water tables rose and climate became wetter (Watts 1980). Todays lakes, bayheads, and swamps were formed relatively recently, begi nning around 5000 years before present (YBP) as water tables reached near present levels. The pr esumed increase in thunderstorms as the climate warmed and moistened, as well as probably the ac tions of newly-arrived humans, increased the importance of fire. Upland ecosystems began to assu me their modern aspects, with longleaf pine savannas becoming more widespread acr oss upland Florida (Myers 1990). Human Florida Humans entered Florida at leas t 12,000 YBP, during the late Pl eistocene, when the climate was much drier than today, and the land area mu ch greater. These early Paleoindian nomadic hunter-gatherers hunted Pl eistocene megafauna, perhaps leading to the extinction of these large vertebrates by the end of the last glaciation. Before ca. 9500 YBP, water sources were scarce in the interior peninsula due to lower water tables; ar tifacts from this period are largely restricted to sites where there were watering holes, mostly in the karstic Tertiary lim estone regions of the northwest peninsula, and along ol d coastal areas that are now submerged. The earliest evidence of human habitation near the Lake Wales Ridge is at Nalcrest in Polk County, on Lake Weohyakapka, dating from around 9500 YBP, a time when the environment was beginning to change, becoming warmer and less arid, although still drier than toda y. Between 9500 and 5000 YBP, gradually moistening climate, rising sea level, and shrinking land area characterized the Archaic period, when human populations increased and people found a greater number of wellwatered places to settle. By around 5000 YBP, climate and habitat conditions had become similar to those of today. Humans had become less nomadic, settling throughout the peninsula. By the end of the Archaic period (2500 YBP), pottery was widespread and regional cultures had proliferated. With the spread of agricu lture after A.D. 750, many cultures became more 23


complex. Certain areas of Polk and Highla nds Counties are associated with scattered archaeological remains from the Belle Glade culture, which flourished from the Archaic period up until colonial times, in the Okeechobee basin and Kissimmee River drainage regions. At the time of first European contact, a nd throughout the Spanish mission period, a huntergatherer tribe known as the Jororo lived in the area of Polk and Highlands Counties. This inland area was relatively isolated from Spanish colonial activity until the late 17th century, when the Spanish established several missions among the Jo roro, which were abandoned after only a few years (Hann 1996; Milanich 1995). By the late 18th century, the Jororo, along with the rest of Floridas indigenous peoples, thei r populations decimated by diseases and slaving raids, had all but vanished from Florida (Hann 1996). As th ese populations were dwindling, Creek Indians from Georgia began settling in north Florida, ev entually becoming the Seminoles. After Florida became a U.S. territory in 1819, Am erican settlers moved into nor th Florida, and the resulting friction between settlers and Seminoles led to the Seminole Wars, which forced a few hundred remnant Indians to flee south toward the Ev erglades, passing through a virtually unsettled central Florida (Milanich 1995). Schafer (1996) states that the Armed Occupation Act of 1842 encouraged white people to settle in the Hills borough County area, the e dge of wilderness (p. 218), where they established cat tle ranches and slowly pushed inland toward what was to become Polk County. By the time of the Civi l War, there was a fle dgling open-range cattle industry in the area, which became Polk C ounty in 1861 (Brown 1996). By 1880, Polk County had a population of just over 3000 people, but the arrival of railroads in the 1880s spurred an explosion of growth and development; population expanded, and the citrus industry especially thrived (Frisbie 1976). After rec ouping from disastrous freezes in the 1890s, the citrus industry has prospered throughout the 20th century (Proctor 1996). Af ter the 1970s, new farming 24


techniques accelerated the conversion of scrub land s to citrus groves, increasingly imperiling this already fragmented ecosystem (TNC 2006c). Du e mostly to immigration, Floridas population has increased more than sixfold since 1950, and the building of Disney World and other themepark tourist attractions near Orlando since 1971 has brought accelerated urban development to central Florida, which is spilling into Polk County (Mohl and Mormino 1996). Since 1950, more than 70 percent of Lake Wales Ridge xeric upland habitats have been lost to citrus groves and, increasingly, residential development (Mye rs 1990; Peroni and Abrahamson 1985). For conservationists, this human impact injects so me urgency into the task of preserving the remnants of these habitats and the constellation of rare species they support. History of the Preserves In the areas of Tiger Creek and Saddle Bla nket Preserves, human impacts have been relatively light, given the surrounding changes. The following account is based primarily on the Tiger Creek and Saddle Blanket management plan documents (TNC 2006a, 2006b), which in turn draw upon largely anecdotal information fr om local sources. One such source, for Tiger Creek, is a 1992 interview by preserve managers of Mr. Bob Byrd, a land manager since 1949 for Alico, Inc., a citrus company that ow ns groves and other land in the area. At both Preserves there is evidence of preh istoric Native American habitation. Preserve staff have found projectile points, pottery sherds, and mounds that may be shell middens within Tiger Creek Preserve boundaries; these have never been formally studied. At Saddle Blanket, artifacts from a mound were unearthed in 1971 by Talbot Lewis, owner of the property at the time, which were donated to the University of Florida. Settlement of the Lake Weohyakapka area did not begin until the turn of the 20th century; by the 1940s there were just a few scattered homesteads around Tiger Creek. There is also some evidence of a few early-20th century homesteads in the vicinity of Saddle Blanket Preserve. From colonial times through the 19th 25


century, human-related impacts on Preserve lands we re primarily by way of introduced animals. The Spanish did not create settlements in thes e areas, but they had in troduced feral hogs to Florida, whose impacts on vegetation can be signifi cant. Hog-rooted soil is still a fairly common sight in the Preserves; managers regularly hire local residents to trap and remove these animals. By the 1850s, open-range cattle grazing was common in both Preserves, and this continued until 1947, when fence laws were enacted. Grazing continued at Tiger Creek until 1957, when the area west of the Preserve was converted to citrus groves. At Saddle Blan ket, grazing continued in the scrublands until the 1970s; westernmost ar eas of this Preserve had been converted to pasture and are still being grazed. One of the earliest recorded owners of the la nd that now includes Tiger Creek Preserve was Consolidated Land Co., or Consolidated Tomoka, which owned large tracts of Lake Wales Ridge land in the early 1900s. The company supplie d naval stores, and although it carried out extensive turpentine extraction from longleaf pi nes, reportedly no turpentining was ever done at Tiger Creek. Railroad companies at that time were among the larg est users of timber, and in 1906 the Atlantic Coastline Railroad Co. bought the Tiger Creek area from Consolidated as an investment in the timber resources of the land. Timber was not harvested from the Tiger Creek area until 1940, when the L.D. Mullins Lumb er Co. cut most of the longleaf pine (Pinus palustris) in the uplands of the Pres erve, leaving perhaps only two pi ne trees per hectare, with no replanting. Additional harvests were done by other companies in 1949 and 1950, taking remaining longleaf and some slash pines (Pinus elliottii) Other than the timber harvests, Mr. Byrd asserted that the land was otherwise undisturbed. No ditche s had been dug, no pesticides or fertilizers had been used, nor had any roller-c hopping or root-raking been done. In the early 1960s, the Hercules Company harvested the old pine stumps for their resin, which was used as a 26


binder in dynamite. This activity probably damage d some of the vegetation; it left some scars and depressions in the soft sand that are still vi sible today. In 1960, Alico, Inc. was formed as a company separate from the railroad co mpany, to oversee the land holdings. Mr. Byrd did not recall any large wildfires at Tiger Creek in his time, but from 1970 to 1985, Don McLean, who held a cattle and hunting lease on 324 ha of what would become the Preserve, conducted periodic contro lled burns in what is now the Preserves central highlands. Besides these burns, fires were generally suppre ssed in the Tiger Creek area. Mr. Byrd and Steve Morrison, the current manager of Tiger Creek Preserve, recollect two occasions of people harvesting Osmunda from the creek floodplain for its fiber, which was sold as a potting medium for orchids. Until the 1960s, the lands around Tiger Creek were also used by the public for hunting. Lands now bordering Tiger Creek Preserve on the north, east, and south were largely undeveloped until the 1980s, when low-density ru ral housing developments were established. Citrus groves on the west side were well established by 1960. Interest in preserving Tiger Creek began in the 1920s when Edward Bok, founder of Bok Tower Gardens (now Historic Bok Sanctuary), bega n efforts to acquire the property, but he gave up in 1929 when Polk County would not promise ne ver to build a road through it. In the 1960s, interest was rekindled when Ken Morrison, direct or of Historic Bok Sanctuary, and Cary Bok, Edward Boks son, convinced The Nature Conservancy of the importance of the property, and in 1971 TNC purchased the first 235 ha from Otto and Dora Pfundstein. By 1986, five additional purchases from Alico, Inc. brought the total ar ea to 1821 ha. Additional purchases have been added since then (Figure 1-3). From 1985 to 1988 the Preserve was managed by Historic Bok Sanctuary, then TNC established a staffed office at the Preserve. Managers hope to acquire 27


additional lands as buffers for the island-like Pr eserve, as development steadily encroaches upon it. The land that became Saddle Blanket Scrub Preserve was used primarily as pasture until 1971, when Talbot Lewis purchased the land and began clearing ground for a water-park tourist attraction. The project was neve r completed, but significant clearing was done around the larger of the two lakes, and in the large bayhead fr onting Avon Park Cutoff Road. The wetland has largely recovered, but ditches a nd earthworks are still apparent amid the forested growth. Logging probably never occurred in the scrub uplands because of the low commercial value of its sand pine trees (Pinus clausa) but according to a local resident, a temporary logging railroad had been built through the property in 1922 for ha uling logs from nearby stands. This old railroad bed still exists as a berm and ditch cutting east-west through the Preserves uplands. In 1972, the property was bought by a south Florida company, subdivided into small lots, and named Frostproof Estates. This probably spar ed the property from bei ng converted to citrus groves, and the fact that roads were never built prevented access to the lots, so no houses were ever built. The present Preserve lands, totaling 334 ha, were acquired over more than a decade by TNC starting in 1988. The original intention wa s to transfer all thes e acquisitions to the Florida Division of State Lands to be manage d by the Florida Department of Environmental Protection. However, the TNC Florida Chapter d ecided to retain ownership of the Saddle Blanket property, and keep it as one of its La ke Wales Ridge Preserve s. In 1996 the Florida Cabinet approved canceling its option to purchase Saddle Blanket, allowing the Conservancy to retain it as a Preserve. Ownership of a 36 ha segm ent that was previously transferred to the State of Florida in 1992 was transferred back to the Co nservancy after it decided to keep the property. All surrounding lands today are ag ricultural. Uplands to the nor th and east, probably formerly 28


scrub and sandhill habitats, have been in citrus production for decades, and lower lying lands to the west and south are cattle pastures. A 40 ha ar ea bordering the south of the Preserve features scrub degraded by grazing, which still harbors so me listed endemic plant species that are found in the Preserve. This area is be ing considered for acquisition. 29


Figure 1-1. Peninsular Fl orida, showing the central ridge system and Polk County. (Data source: Florida Geographic Data Library: ). 30


Figure 1-2. Polk County area, showing the Polk Upland, ridges, and locations of Tiger Creek and Saddle Blanket Preserve s. (Data source: Florida Geographic Data Library ). 31


32 Figure 1-3. Tiger Creek Preser ve acquisition map (TNC 2006a).


CHAPTER 2 PLANT COMMUNITIES What exactly is a plant community? Early in the 20th century, F. E. Clements propounded the idea of the plant community as a tightly integrated, well-defined entity which, like an organism, is born, lives, and dies through the process of succession, along a predictable trajectory, to a predictable endpoint or climax, having an ontogeny and phylogeny that can beexperimentally studied (Clements 1936, p. 17). The cooperative interrelationships among the species making up a Clementsian community work together to give the community a distinct set of emergent properties, and the boundaries of the species ranges equal the boundary of the community (Gurevitch et al. 2002). In counter point was the view of H.A. Gleason, who believed that plant communities are simply aggregations of individual plants; each plant species has its own environmental tolerances and responses, and its presence in the community is governed by chance events (Gleason 1917). The properties of a Gleasonian community are simply the aggregate of the properties of the individuals th at make it up, and its boun daries are indistinct, the ranges of its individual species being independent of each other (Gleason 1926). Work since the mid 20th century has tended to vindicate Gl eason in certain re spects, but not in others. Curtis and McIntosh (1951) mapped vegetation in Wisconsin, showing that species do appear to be distributed indi vidualistically according to thei r tolerances, making community boundaries often indistinct, as Gleason thought. Other work, how ever, supports the idea that communities are more than just the Gleasonian sums of their constituent species. Interactions among these species may lend certain properties to the community. For example, fire-dependent associations like Floridas grassy longleaf pine sandhills appear to have emergent properties influenced by the flammability of some of their member species (see Kerr et al. 1999; Mutch 1970). Yet, studies of pollen records from lake bottoms ( e.g., Davis 1981, 1983; Watts 1980) 33


show that plant species that occu r together today did not necessarily occur together in the past, supporting Gleasons individualistic view of communities. While definable associations of species do sometimes exist and can persist over ti me, and can even display emergent properties, the communities we see today are clearly snapsh ots in time, and frequently intergrade. Our various designations of them are likely ar bitrary, but useful, human constructs. These human constructs are never more useful than when they are put to the service of conservation. Recognizing that conservation is often more effective when focused on communities rather than individual species, The Nature Conservancy makes extensive use of the U. S. National Vegetation Classification (USNVC ), a vegetation-based system the Conservancy helped to develop (Maybury 1999). This system draws together information on ecosystems and rare species compiled by groups at state and local levels, one of which is the Florida Natural Areas Inventory (FNAI). Manage rs of Tiger Creek and Saddle Bl anket Preserves, more or less following FNAIs Guide to the Natural Communities of Florida (1990), recognize the following major plant communities at Ti ger Creek: sandhill, xeric hammock, scrubby flatwoods, yellow sand scrub, mesic flatwoods, wet flatwoods, ba yhead, floodplain forest, depression marsh, and seepage slope (TNC 2006a) (Figure 2-1). At Saddle Blanket, the following communities are recognized: scrub, sandhill, xeric hammock, sc rubby flatwoods, mesic flatwoods, bayhead, depression marsh (wet prairie), seepage slope, sandhill upland la ke, and disturbed areas (TNC 2006b; Figure 2-2). In this st udy, I have generally adhered to these designations, with some additions and modifications as will be seen be low. My descriptions of the communities are based on FNAI (1990), various author s, and my own field observations. Upland Communities The upland communities at Tiger Creek and Sadd le Blanket Preserves are where many rare and/or endemic taxa are found, thus most mana gement is focused on their maintenance or 34


restoration. These communities are scrub, scru bby flatwoods, flatwoods, sandhill, and xeric hammock. All of these are affected in one way or another by fire. Fire caused by lightning is th e most common natural distur bance in Floridas upland habitats, shaping and maintaining their composition and structure (Menges et al. 1993; Reinhart and Menges 2004). Disturbance has often been seen as upsetting to a stable ecosystem, creating chaos and initiating the process of succession (Clements 1936). Certain communities, however, rapidly recover their pre-fire co mposition after a burn; few if any invasive or disturbance-loving species arrive. Thus fire does not initiate succes sion in the Clementsian sense; rather, it is the absence of fire that leads to successional cha nges in this kind of plan t community (Abrahamson 1984). These communities are pyrogenic associations, historically maintained by natural fires. Each kind of pyrogenic community depends for its continued persistence on an optimal range of fire intensities and fire-return intervals to whic h it has become uniquely adapted. Of the upland habitats at both Preserves, scrub, scrubby flat woods, flatwoods, and sandhill are pyrogenic and managed using fire. (Xeric hammock on the Pr eserves is considered by managers to be successional to the other three habitats due to th e longtime absence of fire; these areas are being aggressively managed for restoration to one of these other upland habitats .) Natural fires are patchy and variable, creating a diversity of fire regimes across the landscape to which the various communities, along with their unique species, ar e adapted (Menges 2007). Based on this understanding, managers of both Preserves have drafted elaborate fire management plans (Peterson 2006; TNC 2006c), with the aim of ensuring the persis tence of the unique species on the Preserves whose fire requirements are diverse and often narrow. Plant communities on both Preserves were overgrown at the time of acquis ition due to decades of fire suppression. By 2007, most upland areas and adjacent wetlands on both Preserves had received prescribed 35


burning with the aim of preserving, or in many cases restoring, these habitats. It should be noted that the current appearance of the plant communities on these Preserves (with the exception of areas of creek floodplain habitat) has been shaped to a great extent by more than two decades of prescribed burning. Complex interactions of fire patterns, soils, elevation, hydr ology, and species dispersal characteristics cause these upland communities to occur in an intricate mosaic. Where one community grades into another, ecotones and seem ingly hybrid habitats can occur, which defy attempts to characterize them. This is particul arly true at Tiger Creek, where a patchwork of upland and wetland, created by the undulating terrain of an ancient dune and swale system, has probably caused historical irregulari ties in the frequency and intens ity of natural fires (Christman 1988b; Myers and Boettcher 1987). These peculia rities will be discussed in the community descriptions below. Scrub Scrubs generally are xeromorphic communities dominated by sclerophyllous shrubs that lack a groundcover of grasses or herbs (Chr istman 1988a). Florid a scrubs and scrubby flatwoods, in terms of their vegetation structur e, relationships with fire, and life history characteristics of their plant spec ies, are similar to dry shrubla nds in other parts of the world, such as chaparral in Californ ia, and the fynbos of South Africa (Menges and Kohfeldt 1995). Scrubs, once more extensive in Florida than th ey are today, occupy drough tier areas of the inland sand ridge system, as well as some coastal dune environments. Ocala National Forest features the largest single block of inla nd scrub, while the second largest, now reduced to fragments by human development, occupies the Lake Wale s Ridge (Myers 1990). Although Florida scrubs vary in species composition, most of those on the Lake Wales Ridge include the following: a dense shrub layer of xero phytic evergreen oaks ( Quercus chapmanii, Q. geminata, Q. inopina, 36


and Q. myrtifolia ), along with Lyonia ferruginea Lyonia fruticosa Serenoa repens and often Ceratiola ericoides with or without an overstory of Pinus clausa (Myers 1990). Other species often found in scrub include Carya floridana Galactia elliottii Ilex opaca var. arenicola, Persea humilis Rhynchospora megalocarpa Sabal etonia Selaginella arenicola Sideroxylon tenax, and ground lichens (especially species of Cladonia and Cladina). Open patches of bare sand are common, and herbs and grasses are sp arse if present at all. Florida scrubs are remnants of a xeric habitat that was more widespread in the geological past, when climate was drier and water tables lower (Christman 1988a). Over the last 5000 years, as climate moistened, water tables rose and lightning-caused fires became more common, the range of scrub has contracted, becoming frag mented, existing today as patches in a complex mosaic of vegetation types (Myers 1990). The large number of endemic plant and animal species in Florida scrub suggest that it is an ancient associ ation. There are 56 endemic plant species found in Florida scrub; some 45 of these are listed, either federally or by the state, as endangered or threatened (Stout 2001). Scrubs on the Lake Wales, Lake Henry, and Winter Haven Ridges in Polk and Highlands Counties contain the highest number of endemic plant species of any Florida scrub; many of these ende mics occur only on these three ridges, thus these scrubs, about 200 in number, are referred to as ancient scrubs (Christman 1988a; Christman and Judd 1990). Other scrubs can be called pione er scrubs, presumably being younger and thus lacking the slow-dispersing anci ent endemic species (Christman 1988a). Since no single scrub contains all of the rare species and no one rare species occurs in all the scrubs, many patches must be preserved in order to prevent the exti nction of these taxa (Christman and Judd 1990). Saddle Blanket Preserve is part of a small numb er of protected areas where ancient scrubs are found. Tiger Creek Preserve has only pioneer scrubs (Christman 1988b). 37


Scrub soils are deep, well-drained, nu trient-poor sands, either white sand ( e.g., Archbold series, as at Saddle Blanket ,), or yellow sand ( e.g., Astatula series, as at Tiger Creek, which also supports sandhill vegetation). Most ancient scrubs are white sand scrubs, rarely sharing species with sandhills, while yellow sand scrubs are more likely to be pioneer scrubs, often with some species in common with sandhills (Christman 1988a, 1988b). It was once thought that these vegetation differences were due to differences in fertility between the two soils, but research by Kalisz and Stone (1984) on sandy upl and soils in Ocala National Fo rest failed to find any such differences. Myers (1990) suggested that white sands are just yellow sands that have been bleached by leaching of rainwater made more acidi c by leaf litter in scrubs; in sandhills such acidic leaf litter is burned off in frequent ground fires and cannot build up as it can in scrub, whose fires are infrequent. Thus, a deeper white sand substrate may indicate that the site has supported scrub for a longer period of time than one with yellow sand. Scrubs depend on fire for their persistence. The relatively non-flam mable scrub vegetation burns when the fuel load is sufficient, at an interval now thought to be from 15 30 years for white sand scrubs (rosemary scrubs), 5 12 year s for yellow-sand scrubs (oak-hickory scrubs), and 8 16 years for scrubby flatwoods (Menges 2007) The fires can be cata strophic, top-killing all vegetation. Scrub plants have various life-history strategies in re sponse to fires. Most of the shrub layer resprouts from extensive root system s, while other species regenerate from seeds; Ceratiola ericoides and some of the woody and herbaceous rare endemics have seed banks in the soil, while Pinus clausa regenerates from fire-induced s eed release from serotinous cones (Menges and Kohfeldt 1995). Without fire, Ceratiola ericoides and the short-lived Pinus clausa eventually fail to regenerate, as do certain rare endemic plants, which require fire-cleared bare sand patches, and the habitat becomes a xeric hammock dominated by mixed evergreen oaks 38


(Menges et al. 1993; Myers 1990; Peroni and Abra hamson 1986). Scrubs ofte n occur in the fire shadows of wetlands; natural fire s usually spread to scrubs fr om more flammable adjacent vegetation, such as sandhills, flatwoods, or grassy depressions (Christman 1988a). Fires burn somewhat unevenly in scrubs with many open sa ndy patches, helping maintain a mosaic of patchiness in the vegetation. Scrub is the predominant vegetative comm unity at Saddle Blanket Scrub Preserve, occupying most sites between 38-41 m above mean sea level, covering about 198 ha, a large percentage of the entire preser ve area (TNC 2006b). Of this total area, Preserve managers designate 32 ha as sand pine or rosemary-oak scrub, and 158 ha as scrubby flatwoods (TNC 2006c). Pinus clausa is widely scattered over much of the area; fires and hurricanes have thinned what was once a denser overstory. S ubtle changes in elevation, and thus hydrology, affect the shrub composition of scrubs (Christman 1988a). At higher elevations, where water table is deep, are areas referred to as rosemary-o ak scrub (denoted as sc rub in Figure 2-2). Here, shrubs are somewhat widely spaced and of ten dwarfed, and open bare sandy patches are abundant. Frequent species include Carya floridana, Ceratiola ericoides, Ilex opaca var. arenicola, Lyonia fruticosa, Persea humilis, Quercus chapmanii, Q. geminata, Q. inopina, Q. myrtifolia, Sabal etonia, and Sideroxylon tenax. Less frequent are Asimina obovata Chionanthus pygmaeus Conradina brevifolia Licania michauxii Lyonia ferruginea, Mimosa quadrivalvis var. floridana, Opuntia humifusa, and Serenoa repens. Prunus geniculata (an endemic) is encountered only rarely. Ground c over is sparse, and includes Cnidoscolus stimulosus, Dichanthelium chamaelonche subsp. breve, Euphorbia polyphylla, Galactia elliottii Lechea deckertii, Paronychia americana, Polygonella polygama, Rhynchospora megalocarpa, Selaginella arenicola, Si syrinchium xerophyllum, Stipulicida setacea var. setacea, and ground 39


lichens. Non-native taxa are ra rely encountered. Scattered ab out in open sandy patches, and earning these high elevation areas the designati on of ancient scrub, are the scrub endemics Asclepias curtissii, Bonamia grandiflora Crocanthemum nashii, Crotalaria avonensis Hypericum cumulicola, Lechea cernua, Liatris ohlingerae Paronychia chartacea Polygonella basiramia, and P. myriophylla. Many of these endemics, as well as Ceratiola ericoides, are restricted to bare-sand gaps, which are created and maintained mainly by fires (Hawkes and Menges 1996). Along a gradient toward lower elevations closer to the water table, shrub vegetation becomes more dense, with a greater abundance of Lyonia fruticosa and Serenoa repens, and often Bejaria racemosa Ilex glabra, Lyonia lucida Smilax auriculata, Vaccinium darrowii, V. myrsinites and Vitis rotundifolia Many of the dominant shrubs in these areas, such as sclerophyllous oaks, blueberries, and Lyonia spp ., are resprouters and clonal spreaders, which aggressively close any open sandy patches (M enges and Kohfeldt 1995), hence these areas generally lack the rare scrub endemics and Ceratiola ericoides. Christman (1988a) calls these areas low scrubs, but Preserve managers designate much of this as scrubby flatwoods (Figure 2-2). At Tiger Creek Preserve, the only type of scrub recognized by managers is yellow sand scrub, in patches totaling about 5.7 ha. This is considered a pioneer scr ub as it lacks the rare endemics found at Saddle Blanke t. Christman (1988a) thought that some of Tiger Creeks scrublike vegetation was old pioneer scrubs succeeding to xeric ha mmock due to fire exclusion. Christman and others have noted that yellow sand scrub could also be seen as intermediate between scrub and sandhill, possessing some spec ies found in sandhill but rarely in white sand scrub, including the endemic taxa Clitoria fragrans, Eriogonum longifolium var. gnaphalifolium, and Polygala lewtonii species that are actually more typica l of ecotonal habitats than of either 40


scrub or sandhill (Christman 1988a, 1988b; USFWS 1996). Also characteristic of ecotones are the endemics Chionanthus pygmaeus, Nolina brittoniana and Prunus geniculata (Christman 1988b). Yellow sand scrubs at Tiger Creek (s ee Figure 2-1) feature a dense shrub layer dominated by Quercus geminata, Q. myrtifolia, and Serenoa repens, with abundant Carya floridana and widely scattered Pinus clausa Other frequent species include Callicarpa americana Diospyros virginiana Garberia heterophylla, Osmanthus megacarpus, Sideroxylon tenax, Vaccinium stamineum, and Vitis rotundifolia Less frequent are Clitoria fragrans, Opuntia humifusa, Persea humilis, Rhus copallinum and Rhynchospora megalocarpa. Christman (1988a) observed that scrubs often form ecotones with xeric hammock, in which Chionanthus pygmaeus, Ilex ambigua, and Osmanthus megacarpus are abundant. There are areas on the Preserve that fit this description quite well. Besides yellow sand scrub, othe r scrublike vegetation on Tiger Creek Preserve is placed by managers in the scrubby flatwoods category, afte r consultations with scientists at Archbold Biological Station (TNC 2006a). My own observations suggest that scrubby flatwoods tends to be a rather flexible category. For example, certain scrublike areas in the far eastern part of the Preserve near the creek appear more like ove rgrown pioneer scrubs than scrubby flatwoods, resembling the extensive sand pine dominated scrub forests found in Ocala National Forest. These areas feature the frequent occurrence of species such as Carya floridana, Ilex opaca var. arenicola, Persea humilis, and Sabal etonia, along with sclerophyllous oaks, and a fairly dense canopy of Pinus clausa. Chasteen (1982) surveyed areas of sand pine scrub forest just southeast of this part of the Pres erve, and reported these species plus Bonamia grandiflora, an endemic species found in both white sand and yellow sand scrubs, and Nolina brittoniana and Polygala lewtonii, endemic species found in scrubs and san dhills and their ecotones. Currently, 41


Preserve managers consider the areas descri bed above as scrubby flatwoods, and they are designated as such on the vegetation map (Figure 2-1). Community designation of such areas is problematic and probably somewhat arbitrary, given the intergrading na ture of scrub, scrubby flatwoods, and sandhill. Although it is beyond the sc ope of this study to re-draw the Preserve vegetation maps, I believe further study of scrublike vegetation coul d bring some finer resolution to a currently somewhat catch-all concept of scrubby flatwoods. Scrubby Flatwoods Preserve managers consider scrubby flatwoods to be a type of scrub, although Myers (1990) speaks of it as a form of flatwoods, illustrating the fact that it has characteristics of both, often occurring as an ecotone between the two. This category, as used at Tiger Creek, can be seemingly arbitrary and somewhat of a catch-a ll, including habitats that are ecotones or hybrids between scrubs, sandhill s, and flatwoods. Managers c onsider it to be the most common shrub-dominated habitat at both Tiger Cr eek (166 ha) and Saddle Blanket (158 ha). Scrubby flatwoods occur on deep well-drained sandy soils but with the water table higher than that of scrub or sandhill, and c onsist of a shrubby unders tory of moderate to high density that includes species typical of both scrub and mesic flatwoods, with or without a scattered pine overstory (FNAI 1990; Myers 1990). At Tiger Cree k, much of the scrubby flatwoods is ecotonal between depression marshes or flatwoods, and hi gher elevation sandhills, and is dominated by sclerophyllous oaks ( Quercus chapmanii, Q. geminata, Q. inopina, Q. myrtifolia ) Lyonia fruticosa, Lyonia lucida, Serenoa re pens, Vaccinium myrsinites, and V. darrowii Less frequent are Bejaria racemosa, Garberia heterophylla, Gaylu ssacia nana, Licania michauxii, and Quercus minima. A scattered overstory of pines (either Pinus elliottii, P. palustris, or P. clausa ) may or may not be present. Herbaceous ground cover is sparse and patchy, and may include Aristida stricta A. purpurascens, Andropogon spp., Lechea deckertii, Palafoxia feayi, Pityopsis 42


graminifolia, and Smilax auriculata. The rare endemic annual herb Warea carteri is found mainly in scrubby flatwoods at Tiger Creek. At Saddle Blanket, many areas designated as sc rubby flatwoods could also be thought of as low scrubs as defined by Christman (1988a). I have described these above, under the scrub category. Management for scrubby flatwoods is si milar to that of scrub, including prescribed burning to simulate a natural fire interval of 8 16 years. Scrubby flatwoods burns more frequently than high scrub due to the hi gher shrub density, with fewer open sandy gaps, allowing fires to carry somewhat more evenly th rough the habitat. Some of what is currently xeric hammock at Tiger Creek may have been scrubby flatwoods prior to decades of fire exclusion. Sandhill Sandhill, also known as high pine, or longleaf pine savanna, occurs on deep, well-drained yellow sand soils, often on the tops and slopes of ridges. It is a pyrogenic savanna-like community whose most classic form is dominated by a scattered overstory of Pinus palustris, and deciduous oaks such as Quercus laevis and Q. incana, with Q. geminata occurring singly or in clonal oak domes, a grassy ground cover of Aristida stricta (wiregrass), and a great diversity of herbs (FNAI 1990). In contrast to scrubs, whose high-intensity fires are infrequent due to the relative nonflammability of their shrub-dominated vegetatio n, sandhills are adapted to low-intensity ground fires occurring roughly every one to five years (R einhart and Menges 2004). Historically these fires were caused by lightning, and most likely also by human beings. The dense flammable carpet of wiregrass, herbs, and fallen pine needles provide s the perfect fuel for this kind of fire, which keeps the shrub layer sparse, giving the habitat an open, parklike aspect, and providing bare mineral soil for the germinati on of pine seeds (Myers 1990). 43


Not all sandhill communities conform exactly to the classic pattern of scattered pines with grassy ground cover. One common variant is known as turkey oak barrens, in which pines are few or absent, wiregrass ground cover is sparse and patchy, and the dominant overstory tree is Quercus laevis In many cases this arrangement undoubtedly resulted from removal of pines by logging after which pines failed to regene rate, but some turkey oak barrens are probably natural (Myers 1990). Of the150 ha of sandhill at Tiger Creek Preserve, ve ry little is classic sandhill; much of it is turkey oak barrens or shrubby ecotonal hybrids between sandhill, scrub, or scrubby flatwoods, or is becoming dominated by Quercus geminata and beginning to look more like xeric hammock than sandhill. Logging of longleaf pines from the preserve in the 1940s, along with a history of cattle grazing, fera l hog rooting, and fire suppression, has no doubt helped shape these habitats, but much of thei r current appearance is probably the natural consequence of the intricate mosaic of undulati ng uplands and swales cr eating a historically variable fire and ecotone pattern in the area (C hristman 1988a). Characterizing the sandhills at Tiger Creek is thus difficult, a nd choosing the state to which to restore and manage them may be somewhat arbitrary. At Tiger Creek, the few areas of classic sandhill have the typical stru cture, with scattered Pinus palustris Quercus laevis, Q. geminata, and in some areas Q. incana, with abundant wiregrass, and species such as Chamaecrista fasciculata Croton argyranthemus, Garberia heterophylla, Licania michauxii, Opuntia humifusa, Pity opsis graminifolia, Polanisia tenuifolia, Pteridium aquilinum subsp. pseudocaudatum Quercus minima, Stillingia sylvatica Stylisma villosa Tephrosia chrysophylla, Tragia urens, Yucca flaccida, and the rare endemic Clitoria fragrans. Grasses are abundant, and include Andropogon floridanus, A. glomeratus var. glomeratus, A. ternarius var. cabanisii, A. virginicus var. decipiens, Aristida stricta, A. 44


purpurascens, A. condensata, Muhlenbergia capillaris, Piptochaetium avenacioides, Schizachyrium scoparium, Sporobolus junceus and Tridens flavus. Sedges include Bulbostylis ciliatifolia, B. warei, Cype rus retrorsus, C. croceus, and Rhynchospora megalocarpa. Less frequent are Asclepias humistrata Berlandiera subacaulis Carphephorus corymbosus Dalea carnea, D. feayi, D. pinnata, Lespedeza hirt a, Liatris tenuifolia, Lygodesmia aphylla, Physalis arenicola P. walteri, and Rubus cuneifolius. Areas of turkey oak barrens and other ecotonal sandhills include many of these species, but scattere d much more sparsely, with frequent areas of relatively bare ground. Christman (1988a) noted that certain rare endemic species are actually more common in ecotones and hybrid sandhill than they are in classic sandhill or scrubs, and these do occur in such habitats at Tiger Creek ( e.g., Clitoria fragrans, Eriogonum longifolium var. gnaphalifolium, Nolina brittoniana, Polygala lewtonii, and Prunus geniculata). At Saddle Blanket, the few hectares of sandhill remnants present are severely overgrown, appearing in two separate areas, featuring a few longleaf pines a nd turkey oaks, with sand pines and scrub vegetation mixed in with patches of wiregrass and a few sandhill herbs. Their appearance resembles changes in sandhill due to longtime fire exclusion as described in a study by Myers and White (1987), suggesting that sand hill vegetation once existed there. Several endemic, mostly scrub, species are found in these places, including Bonamia grandiflora, Chionanthus pygmaeus, Conradina brevifolia, Ilex opaca var. arenicola, Paronychia chartacea, Polygonella myriophylla, and Prunus geniculata. Longleaf pine sandhills once covered a pproximately 37 million hectares across southeastern North America, but human encroachment has reduced this ecosystem to about 3% of its original range, making it one of the mo st endangered habitats in the world (Condon and Putz 2007; Landers et al. 1995). Ma nagers of the Preserves place a high priority on restoring and 45


maintaining sandhill, with the hope that some of todays xeric hammock and turkey oak barrens could be restored to a more classic longleaf pi ne and wiregrass associ ation. They are using prescribed springtime restoration fires and ma nual tree removal to reduce the dominance of Quercus geminata in xeric hammocks that were once sandh ills, and prescribed fires every three to ten years in other sa ndhill areas (TNC 2006a). Xeric Hammock Xeric hammock is an oak-dominated hardw ood forest occurring on deep, well-drained sandy soils (FNAI 1990). Hammock, a term mean ing hardwood forest, has been in use in the southeastern United States for at least a century (Harper 1905). The species composition of xeric hammocks varies, and depends on how the associati on formed. It has long been understood that sandhill and scrub vegetation, in th e long-term absence of fire, w ill succeed to a form of xeric hammock dominated by Quercus geminata and other oaks (Abraham son et al. 1984; Menges et al. 1993; Myers 1985). On the Lake Wales Ridge, the shift to xeric hammock from sandhill or scrub is mostly just a shift in dominance, rather than a qualitative change in species composition; Quercus geminata is already part of these ecosystems, a nd without fire it can attain tree size, forming an overstory that partially shades out understory vegetation (Menges et al. 1993). Preserve managers list 210 ha of xeric hammo ck at Tiger Creek, and 34 ha at Saddle Blanket (TNC 2006a, 2006b). At Tiger Creek there is an overstory of Quercus geminata with a variable understory of scattered low shrubs and sparse grasses, whose species composition suggests in most cases a sandhill origin. Pres erve managers consider most of this acreage to be overgrown sandhill, not a separate community in its own righ t, and are actively working to remove the oak overstory through fire and mechanical means. Th e goal is to convert most of it to sandhill and scrubby flatwoods, from which it is presumed deri ved. At Saddle Blanket, some of the xeric hammock features a closed canopy of mature Quercus geminata, Q. virginiana, along with some 46


sand pines that have invaded, with a very sparse understory including Serenoa repens and a few scattered grasses, and is assumed to be successional to sandhill and scrub. Mesic Flatwoods Pine flatwoods, considered the most widespr ead natural Florida ecosystem, occurs on low, flat ground, on poorly drained, acidic sandy soil s often underlain by a relatively impermeable hardpan layer which creates conditions for standing water during wet periods (Abrahamson and Hartnett 1990). Flatwoods are a py rogenic association, with a fire re turn interval of one to five years (FNAI 1990). Flatwoods often occur in zones of transition be tween wetter habitats such as swamps and marshes, and drier ones such as scrubby flatwoods, sandhills, and scrub (Myers 1990). Following FNAI (1990), preserve manage rs recognize two kinds of pine flatwoods: mesic and wet, whose species composition varies due to hydrol ogical differences. Mesic flatwoods at Tiger Creek (258 ha) a nd Saddle Blanket (24 ha) feature an open overstory of Pinus palustris (on slightly higher elevations) or Pinus elliottii, with an understory dominated by shrubs such as Serenoa repens, Ilex glabra, Lyonia lucida, L. fruticosa, Vaccinium myrsinites, Quercus minima, and Myrica cerifera var. pumila. Certain of these flatwoods areas tend to have a relatively abundant herb and grass cover, including Aristida stricta, A. purpurascens, A. spiciformis, Andropogon brachystachyus, A. glomeratus var. glaucopsis, A. virginicus, Dichanthelium spp., Paspalum setaceum, Schzachyrium scoparium, Sporobolus floridanus; various sedges, including Cyperus retrorsus, C. croceus, Rhynchospora ciliaris, R. pineticola, R. plumosa, Scleria triglomerata, and S. reticularis; herbs including Asclepias pedicellata, Galactia elliottii, Piloblephi s rigida, Pterocaulon pycnostachyum, Sabatia brevifolia, S. difformis, and in a few cases, the rare endemic Panicum abscissum (cutthroat grass). At Tiger Creek, most of the mesic flat woods understory is dominated by dense thickets of Serenoa repens, and sometimes abundant Rubus argutus and Vitis rotundifolia, while herbs 47


and grasses are sparse or absent. This conditi on is probably due to a history of grazing, logging, and hog-rooting, followed by long-te rm fire exclusion, as well as perhaps nutrient runoff from citrus groves on the west side of the Preserve. Where flatwoods are adjacent to wetlands, fire exclusion resulted over the decades in the encr oachment of bayhead hardwood vegetation into considerable portions of the flatwoods, a succes sional pattern documented in similar habitats by Menges et al. (1993). Preserve managers ha ve introduced prescribed burning, which has eliminated the bayhead species, opening up th e flatwoods. However, once hardwoods have invaded, they may cause changes that compromise the integrity of the hardpan soil, altering the hydrology, making it possibly difficu lt for these flatwoods to r ecover their pre-alteration composition (Abrahamson and Hartnett 1990). Lowland Communities Lowland communities occur at both Tiger Creek and Saddle Blanket Preserves. At Tiger Creek they are quite extensive due to the va st floodplain areas surroundi ng the creek system. They include wet pine flatwoods, bayhead, seepage slope, depression marsh (or wet prairie), floodplain forest, and blackwater stream. Wet Flatwoods At Tiger Creek, at lower elevations on the we st side of the preser ve and bordering creek floodplain areas near the confluence of Tiger and Patrick Creeks, are about 31 ha of wet flatwoods, characterized by a variably dense canopy of Pinus elliottii. Presumably because it is more often flooded, the understory lacks palmetto s, but often supports some bayhead species such as Gordonia lasianthus Magnolia virginiana and Persea palustris with scattered shrubs including Ilex glabra, Lyonia ligustrina var. foliosiflora L. lucida, Myrica cerifera var. cerifera, and a variable ground cover of Osmunda cinnamomea Woodwardia virginica, Polygala rugelii, Rhexia mariana, and a few sedges and grasses, including Cyperus retrorsus, Eleocharis 48


baldwinii, Scleria triglomerata, Andropogon glomeratus var. glaucopsis, Aristida spiciformis, Dichanthelium spp., and Sporobolus floridanus Bayhead A bayhead, or baygall, is a forested, acidic, peat-filled seepage wetland located at the base of sandy slopes (FNAI 1990). Tiger Creek has 65 ha, and Saddle Blanket has 16 ha, of bayheads. The dominant tree species of th ese bayheads, forming a dense canopy, are the evergreen broadleaved Gordonia lasianthus and Magnolia virginiana, with a subcanopy of Ilex cassine and Persea palustris Scattered shrubs include Itea virginica, Myrica cerifera var. cerifera, Vaccinium corymbosum, and Viburnum nudum, with a ground cover dominated by ferns ( Blechnum serrulatum, Osmunda cinnamomea, O. regalis, and Woodwardia virginica). In more open areas, one can find Peltandra saggitifoli a, P. virginica, and a few sedges including Eleocharis baldwinii, Rh ynchospora fascicularis, and Scleria triglomerata. Tillandsia setacea, T. simulata, and T. utriculata are common epiphytes on tree trunks. Around the edges, Rhododendron viscosum, Smilax laurifolia, Gelsemium sempervirens, and Vitis rotundifolia may be found. Bayheads are seasonally inundated, fe d by seepage from surrounding uplands, and most of the time they are too wet to burn, so their fire interval is long, on the order of 50-100 years (FNAI 1990). Their hydrology is dependent on the surficial water ta ble; there is some concern that continued water withdrawals fr om deeper aquifers by surrounding land uses, especially citrus irrigatio n, could endanger the bayhead community (TNC 2006b). Preserve managers are using prescribed fire to burn into the bayheads, as long as they remain wet enough to prevent smoldering peat fires which could pe rmanently damage their vegetation, thus keeping bayhead species from invading surro unding habitats (TNC 2006a, 2006b). 49


Seepage Slope A seepage slope is an acidic open wetland at the base of a sandy slope, fed by seepage so that it is saturated but seldom inundated (FNAI 1990). Species composition can vary between shrubby and grassy. These habitats at Tiger Cr eek and Saddle Blanket are dominated by the rare endemic Panicum abscissum (cutthroat grass), along with a few other graminoids ( e.g., Andropogon glomeratus, Aristida stricta, Rhynchos pora fascicularis, Scleria triglomerata, and Xyris ambigua ), and are called cutthroat seeps, a type of seepage slope found only in the region of the Lake Wales Ridge (TNC 2006a; FNAI 1990) These wetlands are very small and few in number at the Preserves, comprising less than 2 ha. Managers are using prescribed growingseason fires every one to four years, which stimulate cutthroat grass to bloom and prevent invasion of shrubs. Hog rooting is a problem in these areas. Monitoring over several years has shown that cutthroat grass does not readily recolonize hog-rooted areas, suggesting that such damage is long lasting. Hog-rooted cu tthroat seeps often become dominated by Lachnanthes caroliana. Some of the cutthroat seeps at Tiger Creek have been fenced to exclude hogs. Depression Marsh Depression marshes (also known as ephemera l ponds, or wet prairies) are shallow, somewhat rounded depressions in sandy soils, char acterized by grassy an d herbaceous vegetation in concentric bands, with soil organic matter in creasing toward the cente r, inundated with water for 50 to 200 days per year (FNAI 1990). Tiger Cr eek Preserve has 40 ha of depression marshes, most occupying swales between dunes in the undulat ing uplands of the preserve. Saddle Blanket has about 4.9 ha of depression marshes. At Tige r Creek, typical species in clude an outer ring of Serenoa repens, then a broad area of graminoids, including Amphicarpum muhlenbergianum, Coelorachis tuberculosa, Cyperu s lecontei, Fimbristylis puberu la, Fuirena scirpoidea, Panicum hemitomon, Rhynchospora cephalantha, R. ciliaris, R. chapmanii, R. fascicularis, R. latifolia, 50


Scleria reticularis, Xyris brevifolia, X. elliottii, and X. ambigua. Herbs are diverse, and include Aletris lutea, Drosera capillaris, Eriocaulon decangulare, Lachnanthes caroliana, Lachnocaulon beyrichianum, L. minus, Lycopodie lla alopecuroides, L. appressa, Pinguicula pumila, Pluchea baccharis, P. foetida, Polygala cymosa, P. rugelii, P. nana, Proserpinaca pectinata, Rhexia petiolata, Sabatia grandiflora, and Syngonanthus flavidulus. There are usually a few scattered shrubs, including Cephalanthus occidentalis, Hypericum fasciculatum, and H. myrtifolium. Panicum abscissum also occurs in a few depression marshes at Tiger Creek. In one particular depression marsh, near some flatwoods south of th e preserves undulating central highlands, I found the endemic species Hartwrightia floridana and Lilium catesbei. At Saddle Blanket, the depression marshes are si milar, but less rich in species. Floodplain Wetlands Floodplain wetlands comprise by far the larg est proportion of Tige r Creek Preserve, occupying more than 971 ha, surrounding the cr eek system. This category is a complex patchwork of floodplain wetland types considered as an aggregate for purposes of this study because of the difficulty of access for surveying a nd characterizing in detail its various aspects. It most closely resembles black water floodplain forests as desc ribed by Ewel (1990). In those areas I visited and surveyed, it ap pears that vast portions of th ese floodplains are forested and swampy, supporting swamp and bottomland forest vegetation and experiencing inundation for substantial periods each year. In cer tain areas of the floodplain, charring on Sabal palmetto trunks indicates that fire has reac hed these areas at times in the past. A few areas of slightly higher elevation, especially around the steeper edges of the wetla nd, have a more or less narrow band of drier forests, which resemble mesic hammocks. The floodplain is bordered by large swaths of pine flatwoods on the west side of th e preserve, and south and east of the confluence of the two creeks. There are also some more ope n areas supporting a more marsh-like vegetation. 51


Before the hurricanes and unusual flooding of 2004 and 2005, these open areas were few, but afterwards, in considerable por tions of the floodplain, the whole aspect changed. Many trees were blown down by the hurricanes, and the floo ding lasted long enough to essentially drown the forest in places. These areas present a jumb le of fallen trees with newly opened gaps, and herbaceous colonizing vegetation is rampant, ma king passage difficult in the mucky terrain. The nature of future changes in the floodplain habitat will depend on rainfall patterns in the coming decades. Aerial photos from 1922 show that a huge area around the creek confluence was grassy marsh, possibly a Cladium jamaicense (sawgrass) marsh; today there are still scattered patches of Cladium jamaicense in the forest understory. There is one area of intact sawgrass marsh northeast of Tiger Creek near the Red Gum Trail ar ea. Preserve managers have speculated that continued decades of high wate r may favor the return of sawgrass marsh over large areas, whereas a return to dry conditions will favor re-establishment of forested wetlands. Differences in hydroperiod, fire frequency, and so il organic matter accumulation all have effects on the species makeup of swampy wetlands (Ewel 1990). Preserve managers want to study these wetlands further to clarify the relative importa nce of such factors before crafting specific management plans for the floodplains. Currently, pres cribed fire is used to burn into the edges of the wetlands to encourage developm ent of healthy ecotones (TNC 2006a). In the lower lying areas, typical canopy species include Acer rubrum Liquidambar styraciflua Magnolia virginiana Nyssa biflora Pinus elliottii, Quercus nigra Q. laurifolia, Sabal palmetto and Ulmus americana. Subcanopy and shrub species include Baccharis halimifolia, Cephalanthus occidentalis Cornus foemina Ilex cassine, Itea virginica, Persea palustris, Sambucus nigra subsp. canadensis, and Viburnum nudum. At slightly higher elevations, species such as Ampelopsis arborea, Asimina parviflora, Callicarpa americana, 52


Carya glabra, Diospyros virginiana, Gelsemiu m sempervirens, Prunus serotina, Psychotria nervosa, P. sulzneri, Quercus virginiana, R hododendron viscosum, Rubus argutus, Sabal minor, Serenoa repens, Smilax bona-nox, S. laurifolia, S. pumila, S. walteri, Toxicodendron radicans, and Vitis rotundifolia are found. There is one area near th e Red Gum Trail area north of Tiger Creek with a population of Rhapidophyllum hystrix which is not common on the Lake Wales Ridge. Herbaceous species, some becoming more abundant in recently opened areas, include Apios americana, Arisaema triphyllum, Boehmeria cylindrica, Encyclia tampensis (epiphytic), Eupatorium capillifolium, Habenaria odontopetala, Hypericum hypericoides, H. mutilum, Hydrocotyle umbellata, H. verticil lata, Ludwigia lanceolata, L. leptocarpa, L. peruviana, L. repens, Mikania scandens, Peltandra virginic a, P. sagittifolia, Pe rsicaria punctata, P. hydropiperoides, Saururus cernuus, and the ferns Blechnum serrulatum, Dryopteris ludoviciana, Nephrolepis exaltata, N. cordifo lia, Osmunda cinnamomea, O. regalis, Thelypteris interrupta, T. kunthii ,Woodwardia virginica, and W. areolata. Grasses and sedges include Arundinaria gigantea, Carex comosa, C. vexans, Chasmanthium laxum, Cyperus tetragonus, C. virens, Dichanthelium spp., Echinochloa muricata, Eleocharis baldwin ii, Oplismenus hirtellus, Panicum rigidulum subsp. rigidulum, P. hemitomon, Rhynchospor a corniculata, R. fascicularis, R. miliacea, Sacciolepis striata, Scirpus cyperinus, Scleria triglomerata, S. muhlenbergii, Setaria magna, Tripsacum dactyloides, and Urochloa maxima. Epiphytic bromeliads ( e.g., Tillandsia setacea, T. simulata, T. usneoides, and T. utriculata) are common in places. The dominant canopy species varies from place to place, with Magnolia virginiana in some places, and Acer rubrum, or Sabal palmetto, in others Blackwater Stream Tiger Creek is a blackwater stream originating in deep sandy lowlands where extensive wetlands with organic soils collect and store water, dischargin g it slowly. The water is tea53


colored with dissolved tannins (FNAI 1990), and it s acidity and dark colo r inhibit growth of submersed aquatic vegetation. Emergent and floating plants are co mmon, however, in the shallower places and near the banks. Typical species are Azolla caroliniana, Cephalanthus occidentalis, Cicuta maculata, Eichhornia crassipes, Eleocharis baldwinii, Hydrocotyle umbellata, H. verticillata, Landoltia punctata, Lemna aequinoctialis, Ludwigia peruviana, Micranthemum glomeratum, Nuphar advena, Persicaria hydropiperoides, P. punctata, Pontederia cordata, Sagittaria lancif olia, S. latifolia, Salvinia minima, and Typha domingensis. Exotic invasive species seen in and along the creeks include Eichhornia crassipes, Ludwigia peruviana, and the possibly native and often invasive Pistia stratiotes. Tighe (1987) considered the creek system relatively undisturbed (by humans), although nutrient runoff from surrounding lands, especially citrus groves around Lake Patrick, the source of Patrick Creek, may be having an impact on water quality. Ruderal There are few disturbed (ruderal) areas at Tiger Creek; they include the Preserve boundaries, and a small stretch of roadsides where Murray and Pfundstein Roads cut through the preserve. Various common roadside weeds are found here, as well as most occurrences of certain invasive exotic pest plants. Roadside weeds include such species as Bidens alba, Desmodium incanum, Emilia fosbergii, E. sonchifolia, Gomphrena serrata, Lantana camara, Medicago lupulina, Oxalis corniculata, Paspalum notatum, Richardia brasiliensis, Salvia lyrata, Spermacoce remota, Zeuxine strateumatica, and others (see Chapter 4) most of which are not found within the preserve. Invasive exotic species that are being actively monitored and eradicated when possible include Abrus precatorius, Imperata cylindrica, Lygodium japonicum, L. microphyllum, Melaleuca quinquenervia, Melinis repens, Schinus terebinthifolius, and 54


Solanum viarum With the exception of Imperata cylindrica, which is spotted occasionally within the preserve, these species are mainly found along its edges. At Saddle Blanket, disturbed areas include th e Preserve edges and certain areas recently used as cattle pasture. Species of the edge s are many of the same ones found on Tiger Creek roadsides, and within the Preserve are some non-native grasses such as Eremochloa ophiuroides, Melinis repens, and Paspalum notatum With the exception of Imperata cylindrica and Melinis repens, which are found occasionally, the Preserve interior appears to be relatively free of nonnative or ruderal species. 55


Figure 2-1. Vegetation map of Tiger Creek Preserve (Data source: The Nature Conservancy2007). 56


57 Figure 2-2. Vegetation map of Saddle Blanket Scrub Preserve (Data source: The Nature Conservancy 2007).


CHAPTER 3 FLORISTIC METHODS AND RESULTS Methods Field work was conducted in Tiger Creek and Saddle Blanket Scrub Preserves from September 2004 to February 2008. Aerial photos, along with vegetation maps prepared by preserve staff, were used to ensure adequate coverage of plant communities. A compass and handheld global positioning unit were of great assistance in marking points where plant collections were made, and in navigating areas of difficult access. Most upland and adjacent wetland areas on Tiger Creek and Saddle Blanket Preserves are crisscrossed by jeep trails, from which access is relatively easy by foot and all-terrain vehicle (ATV). From these trails, numerous forays and transects were initiated into and thr ough surrounding areas. Where accessible, Tiger Creek Preserves extensive bou ndaries were traversed by ATV and on foot, with transects and forays into areas of intere st. Tiger Creeks vast floodplain wetlands were sampled by walking, often with great difficult y, from wetland edges toward the creeks in numerous locations. The creeks themselves were too choked with fallen trees to allow passage of a canoe or kayak, so sampling was done by walking along the creek banks for distances of varying length as the terrain permitted. Plant specimens were prepared following standard field and herbarium techniques ( e.g., Judd et al. 2008), and vouchers were deposited in the University of Florida Herbarium (FLAS). Plants were identified using primarily Wunderlin and Hansen (2003), Tobe et al. (1998), Hall (1978), the University of South Florida Plant Atlas website (Wunderlin and Hansen 2004), as well as keys in the Flora of North Ameri ca volumes (Flora of North America Editorial Committee 1993-2007). Recent taxonomic revisions we re consulted when appropriate, as cited in the species list (Chapter 4). 58


Results From the combined areas of both preserves, 554 species or subspecifi c taxa of vascular plants were documented, representing 306 gene ra and 112 families. Of these species or subspecific taxa, 18 are ferns, 3 are lycophytes, 3 are conifers, and 530 are angiosperms. Of this total, Tiger Creek Preserve yielded 520 taxa, in 303 genera and 112 families, while Saddle Blanket yielded 242 taxa, in 160 genera and 76 familie s. The largest families, in terms of total number of species or subspecific taxa from bot h Preserves, are Poaceae (74 taxa), Asteraceae (63 taxa), Fabaceae (39 taxa), Cyperaceae (36 taxa), Ericaceae (16 taxa), Euphorbiaceae and Fagaceae (13 taxa each), Rubiaceae (12 taxa ), Polygonaceae (11 taxa), Hypericaceae and Onagraceae (10 taxa each), and Lamiaceae, Planta ginaceae, and Rosaceae (8 taxa each). The largest genera are Rhynchospora (15 taxa), Andropogon (13 taxa), Quercus (13 taxa), Hypericum (10 taxa), Ludwigia and Dichanthelium (9 taxa each), Polygala (7 taxa), Asclepias, Panicum, and Smilax (6 taxa each), and Chamaesyce, Cyperus, Eupatorium, Polygonella, and Tillandsia (5 taxa each). Taxa of Special Interest or Concern Quite a number of species or subspecific ta xa found on these preserves are of special interest or concern: 37 taxa are at or near the southern or northern limits of their ranges relative to their distributions in Florida; 24 taxa ar e new Polk County records, having never been vouchered in the county before; 54 taxa are non-native to Florida. Fifty-five taxa are endemic to Florida, of which 27 are listed as endangered or threatened in Florida, while 18 are federally listed as endangered, threatened, or of special management concern. These taxa are discussed in detail below. Taxa at the limits of their ranges: Todays scientific consensus is that global climate is rapidly warming, that human activity is likely a major cause, and that one effect of this climate 59


change is that the ranges of many plant species will be altered in the coming centuries (Field et al 2007). Meta-analyses have documented significant poleward range shifts of many species in the last century, and although other f actors besides climate affect speci es distribution, the fingerprint of climate change is now considered unmistakab le (Lavergne et al 2006; Parmesan and Yohe 2003). Throughout geological history, vegetation ha s migrated in response to changes in climate, but those changes typical ly occurred at a much slower rate than the rapid pace being observed today and projected for the future, sugges ting that plant species will face unprecedented challenges (Davis and Shaw 2001). Conservation of rare species may therefore require human intervention as ranges shift and old habitats become unsuitable (Maschinski et al. 2006). For these reasons, information on range limits is important. Wunderlin and Hansen (2004) was used to de termine species ranges in Florida. For purposes of this study, taxa at the limit of their ranges reach to Polk County from the north or the south, but no further. Taxa near their range lim its occur no more than two counties to the north or south of Polk County. Non-native taxa are marked with an asterisk. Five taxa are at their southern limits: Albizia julibrissin*, Ar undinaria gigantea, Croton argyranthemus, Eryngium prostratum, and Hieracium gronovii. Twenty-eight taxa are near their southern limits: Andropogon glomeratus, Arnoglossum floridanum, Aronia arbutifolia, Asclepias humistrata, Asimina pygmaea, Cl ematis reticulata, Dalea pinnata, Eriogonum tomentosum, Garberia heterophylla, Gaylussacia nana, Gels emium sempervirens, Gratiola virginiana, Krameria lanceolata, Lespedeza hirta, Lycopodi ella alopecuroides, Lyonia ferruginea, Lyonia ligustrina var. foliosiflora, Peltandra saggitifolia, Pe rsicaria hirsuta, Prunus serotina, Prunus umbellata, Quercus incana, Rhapidophyllum hy strix, Rhododendron viscosum, Rumex hastatulus, Smilax glauca, S. walteri, and Viburnum nudum. One taxon, Croton glandulosus var. 60


floridanus, is at its northern limit. Three taxa, Carphephorus odoratissimus var. subtropicanus, Euphorbia polyphylla, and Melaleuca quinquenervia*, are near their northern limits. New records for Polk County: Twenty-four taxa are Polk County records: Albizia julibrissin*, Andropogon tracyi, Arundinaria giga ntea, Catalpa bignonioides, Chamaecrista nictitans, Chamaesyce ophthalmica, Cucumi s anguria*, Desmodium floridanum, Elephantopus carolinianus, Landoltia punctata*, Lechea minor, Matelea floridana, Melochia spicata, Mimosa strigillosa, Oplismenus hirtellu s, Paspalum urvillei*, Polygal a polygama, Pyrostegia venusta*, Quercus minima, Rhynchospora scirpoides, Sabal minor, Smilax walteri, Sporobolus jacquemontii*, and Verbena brasiliensis*. For one study to produce so many new county records suggests that floristic work in th is part of Florida is as yet incomplete. Taxa non-native to Florida: Fifty-four taxa found in the Preserves are non-native to Florida. Many of these are considered invasive (FLEPPC 2007), and Pres erve staff are actively working to control and eradicate some of them ( Abrus precatorius, Imperata cylindrica, Lygodium microphyllum, Melaleuca qui nquenervia, Rhyncheletrum repens, and Solanum viarum). Most of these exotics are found main ly around the Preserve boundaries, although Solanum viarum and Lygodium microphyllum are also found deep within the floodplain wetlands, and Imperata cylindrica is found occasionally in the Preser ve interior. Also subject to these control efforts, but not collected in this study, are Dioscorea bulbifera Lygodium japonicum Schinus terebinthifolius, and Sorghum halepense Other exotic species considered invasive but not being activel y treated at this time are Albizia julibrissin, Alternanthera philoxeroides, Lantana camara, Ludwigia peruviana, Momordica charantia, Nephrolepis cordifolia, Urochloa maxima, and Urena lobata. One non-native species, Pyrostegia venusta, was collected at Tiger Creek, then subseque ntly, as far as is known, eradicated. Pistia stratiotes, 61


an invasive species listed in th is study as native (see Wunderlin and Hansen 2003), is considered non-native by some experts (FLEPPC 2007). For additional non-native ta xa found naturalized on the Preserves, see Chapter 4, where all non-native ta xa are denoted by an aste risk in front of the species name. Taxa endemic to Florida: Fifty-five taxa found on Ti ger Creek and Saddle Blanket Preserves are endemic to Florida. Twenty-seven of these are restricted to Florida peninsular scrubs or scrub/sandhill ecotonal ha bitats (Table 3-1). Of these 27 taxa, 15 are restricted to Floridas central ridges, especia lly to the ancient scrubs on th e Lake Wales, Lake Henry, and Winter Haven Ridges (Christman and Judd 1990). Of th ese, nine species are re stricted to four or fewer counties: Clitoria fragrans, Conradina brevifolia, Crotalaria avonensis, Hypericum cumulicola, Liatris ohlingerae, Polygonella basiramia, P. myriophylla, Prunus geniculata, and Ziziphus celata. The remaining 28 taxa are Florida endemics not restricted to scrubs (Table 3-2). Of all 55 endemics, 27 are listed as endangered or threatened in Florida, and 18 are listed federally as endangered, threatened, or of ma nagement concern (Coile and Garland 2003; USFWS 2007; see Tables 3-1 and 3-2). One of these species, Ziziphus celata, was not found growing naturally on either of the Preserves, bu t it is being deliberately re-introduced at Tiger Creek as part of a recovery effo rt in an experimental framework. Six species found on the Preserves are considered nearly endemic, with at least 90% of their distribution being within Floridas state boundaries: Andropogon floridanus, A. brachystachyus, Callisia ornata, Pa lafoxia integrifolia, Pinus clausa, and Sisyrinchium xerophyllum (Muller et al. 1989). Six non-endemic species found on the Preserves are protected by listings at the state level: Coelorachis tuberculosa threatened; Tillandsia utriculata endangered; and Encyclia 62


tampensis, Osmunda cinnamomea, O. regalis, and Rhapidophyllum hystrix commercially exploited (Coile and Garland 2003). Endemism on Floridas Ancient Sand Ridges Of all the abovementioned taxa of special inte rest, the endemics are remarkable for their sheer numbers in such a small study area. Florida, as a whole, contains numerous endemic plant species; why this is so has been the subject of much study over the ye ars. Harper (1949), tabulating the discoveries of botanists since colonial times, attr ibuted 427 endemics to Florida. A more recent study suggests a figure of 235 endemic vascular species, and 40 nearly endemic, with ranges extending slightly beyond Floridas polit ical boundaries (Muller et al. 1989). Harper (1949) attributed Floridas high ende mism to three characteristics that make it unique: first, it is a peninsula; second, it projects southward to wher e there are no large land masses nearby with similar climate; and third, it features sandy soils unlike those of the continent to the north. Floridas endemic species are concentrated in th ree distinct areas or hotspots: the Apalachicola Bluffs and other areas in the pa nhandle, the subtropical tip of th e peninsula, and the Lake Wales Ridge (Ward 1979). The degree of endemism in the Lake Wales Ridge area is high by continental standards, exceeded only by islands su ch as Hawaii (Chaplin et al. 2000; Christman 1988a). Thus it is no surprise that a survey of Tiger Creek and Saddle Blanket Preserves should turn up so many endemics. Human beings are generally fascinated with rare things, so it follows that it is the narrow endemics, the rare plants, which often capture the interest of botanis ts and lay people alike (Kruckeberg and Rabinowitz 1985). Ward (1979) recognized 170 Florida species as rare, and noted that many are becoming rarer due to habitat loss as humans encroach upon the land. Florida scrubs are high in endemics, especially those on the Lake Wales, Lake Henry, and Winter Haven Ridges (Christman and Judd 1990). Stout (2001) identified 56 endemics 63


associated with Florida scrubs, 48 of which he co nsidered rare. Of Stout s 48 rare scrub species, 21 are found on Tiger Creek and Saddle Blanket Pres erves (these are the first 21 taxa listed in Table 3-1). Ward et al (2003) used a variety of criteria to rank Floridas threatened and endangered plants in terms of ra rity, giving each species a numerical rating between one (most rare) and 12 (least rare). Of the species at Tiger Creek and Saddle Blanket for which such a rating was given (see Tables 3-1 and 3-2), most fall somewhere along the less-rare half of the scale (6.0 or greater), except for Ziziphus celata (1.5), Clitoria fragrans (3.5), Crotalaria avonensis (4.5), Polygala lewtonii (4.5), and Liatris ohlingerae (5.5). An endemic, be it narrow or broad, is simply a species of limited or restricted distribution (James 1961). Mason (1946) points out that all plan t species have their areas restricted to some extent by various conditions, and are thus, in some sense, endemics. But it is the narrow endemics that catch peoples attention. A variety of factors work to define the limits of the area that a given species can occupy. Mason (1946) emphasized the importance of an individual plants responses to environmental factors, as determined by its genetic makeup. Environmental parameters, such as climate and edaphic (soil) factors, are geographically distributed. Edaphic factors can occur in small, loca lized, sharply defined areas, whic h can limit the range of a species whose tolerances are narrow (Mason 1946). Hi storical factors are also important to understanding endemism. For example, changes in climate and sea level over geological time can shrink or expand habitats, affecting the di stribution of species (Woodson 1947; James 1961). Stebbins (1942) stressed the importance of popula tion genetics, observing that narrowly endemic rare species often lack genetic va riability and thus cannot adapt to a wide variety of niches as can a more genetically variable, wide spread species. Yet noting, decad es later, that rarity or commonness of a species appears to have no correla tion with genetic variab ility, Stebbins (1980) 64


advocated, in his gene pool / niche interacti on theory, a synthetic approach that considers environmental, genetic, and historical factors. No one factor alone can account for endemism. Habitat specialization is often cited as a factor in rarity and narrow endemism (MaliakalWitt et al. 2005). Kruckeberg and Rabinowitz (1985) asserted that perhaps the ultimate cause of rarity and narrow endemism are geological proces ses that create fractionated landscapes with numerous discrete habitats and unusual environmen tal characteristics. Florida, of course, has undergone many such processes (see Chapter 1), which have altered its climate and landforms through time, shaping features like the Lake Wales Ridge, whose deep, infertile sands were characterized by Mulvania (1931, p. 424) as little more than a bed of silica, to which the term soil is but remotely applicable. Many narrow endemics around the world are edaphic specialists associated with peculiar soils (Kruckeberg and Rabinowitz 1985) Stout (2001) regarded many, perhaps even most, of Floridas rare endemic scrub plants as ed aphic specialists, their ranges limited by the extent of the patches of peculiar so ils (habitat islands) to which they are uniquely adapted. But are these soils so peculiar? Referr ing to the southeastern coastal plain of North America, of which Florida is a part, Sorrie a nd Weakley (2001, p. 69) stat e that the casual observer may be inclined to consider the coastal plain devoid of many char acteristics considered likely to generate evolutionary innovation. The coastal plain has few obvious geographic dividers, has very subdued topography..., is of rela tive geologic youth, has an absence of striking edaphic factors (such as serpentine geology), an d has a relatively homogeneous climate. Yet the coastal plain contains several centers of high endemism, including those in Florida. Some of these areas, including the Fall-lin e sandhills of the Carolinas and Georgia, the Carrizo sands of Texas, and Floridas Lake Wales Ridge, featur e deep, porous, coarse sa ndy soils (Sorrie and Weakley 2001). Other notable area s of high local endemism feat uring deep, nutrient-poor sandy 65


soils are the white sandlands of southwest Pinar del Rio and the Isle of Youth in Cuba (Alain 1953; Borhidi and Muniz 1986), and areas of Amazoni an forest featuring white-sand patches in otherwise nutrient-rich clay soils near Iquitos, Peru (Fine et al. 2006). But in the Cuban and Amazonian examples, the white-sand areas ar e surrounded by very diss imilar soils, creating obvious edaphic islands. In Florid a, in contrast, the soil differen ces are often not so great. But other factors, such as differences in fire regime, hydrology, and genetically determined predispositions of a plant, can act to limit the ability of a given species to disperse beyond a particular area. Here are some examples: Flam mable longleaf pine sandhill islands persist in Ocala National Forest amid a sea of less flammable sand pine scrub, even though the soils differ little between the two habitats (Kalis z and Stone 1984; Myers 1985, 1990). Areas of rosemary scrub, featuring rare e ndemic plants that depend on bare-sand gaps, are often hemmed in by areas of slightly lower elevation where the soil is essentially identical, but the higher water table favors dense thickets of gap-closing shrubs, effectivel y confining the gap-dependent endemics to restricted areas (Menges and Kohfel dt 1995). And some rare scrub endemics appear to be microhabitat specialists, unable, for perhaps genetically determined reasons, to effectively colonize areas that their habita t-generalist congeners can inhab it easily (Maliakal-Witt et al. 2005). Recent work by Fine et al. (2006) has show n that herbivory, too, can be a selective force that encourages the evolution of habitat specialization. It would seem, perhaps, that although Floridas sandy upland soils may seem homogeneous there are many factors at work, other than edaphic factors, that can create habitat islands upon which narrow ly endemic species can evolve, and to which they can remain restricted. Many of Floridas endemic species appear to be what Stebbins (1942) calls insular species, i.e., species adapted to restricted habitats (either true islands or habitat islands) that 66


have diverged from, and are closely related to, more widespread species located on adjacent land areas. Some may be neo-endemics, having diverg ed from a relatively recent common ancestor, perhaps during times of high sea level when th e Lake Wales Ridge likely was a refuge for populations effectively isolated from their relatives (Huck et al. 1989). A good number of the endemics found on Tiger Creek and Saddle Blanket Preserves fit this description, appearing to be specialized variants of more common southeastern North American species. These include Asclepias curtissii (related to A. purpurascens, Woodson 1954), Chionanthus pygmaeus ( C. virginicus, Hardin 1974), Conradina brevifolia ( C. canescens, Kral and McCartney 1991), Crotalaria avonensis ( C. rotundifolia, DeLaney and Wunderlin 1989), Hypericum cumulicola ( H. gentianoides, Adams 1962), Ilex opaca var. arenicola ( Ilex opaca var. opaca, Wunderlin 1982), Osmanthus megacarpus ( O. americanus, Hardin 1974), Persea humilis ( P. borbonia, Wofford 1973), Polygonella basiramia ( P. ciliata, Nesom and Bates 1984), P. robusta ( P. fimbriata, Nesom and Bates 1984), Prunus geniculata ( P. maritima, Shaw and Small 2005), Quercus inopina ( Q. myrtifolia, Johnson and Abrahamson 1982), Sabal etonia ( S. palmetto, Zona and Judd 1986 ; Zona 1990), and Stylisma abdita ( S. aquatica, Myint 1966). These species, having diverged from their more widespr ead ancestors, have beco me habitat specialists. They have remained more or less isolated in th eir habitat islands, often being poor dispersers, and their distinctive genotypes have persisted (Zona and Judd 1986). Historical factors, too, are an important to understanding Florida scrub endemism. Although at first glance the south eastern coastal plain flora ma y seem geologically young, it is probable that parts of the coastal plain have b een habitable by plants since the Eocene (ca. 50 MYBP), and parts of Florida since at least the middle Miocene (Sorrie and Weakley 2001). During that immense time span, the area has had climates ranging from tropical to cool 67


temperate, wet to dry, and has received inputs from older floras in Appalachia, the Caribbean, and western North America (Sorrie and Weakle y 2001; Webb 1990). Not all Florida endemics appear to have the same biogeographic origin. St ebbins (1942) defined another kind of endemic, the depleted species. These are species that we re once more widespread, but have become rare due to shrinkage of available habitat. Such speci es may lack close living relatives, or else their closest relatives are disjunct and geographically distant. Also known as paleo-endemics, these species are often relics, ancient holdovers from the geological past (S tebbins and Major 1965). Certain scrub endemics found on the Preserves seem to fit this pattern, and they appear to have their closest relatives in southweste rn North America. These include Bonamia grandiflora (Myint and Ward 1968), Carya floridana (Hardin and Stone 1984), Eriogonum longifolium var. gnaphalifolium (Horton 1972), Nolina brittoniana (Judd and Hall 1984), Palafoxia feayi (Turner and Morris 1976), and Ziziphus celata (Judd and Hall 1984; Islam a nd Simmons 2006). During Pleistocene glacial maxima, and probably in the Tertiary period before that, Florida was likely connected to southwestern Nort h America by a continuous corridor of xeric habitat. As sea levels rose and climate moistened, these member s of a once widespread xeric flora underwent a reduction in available suitable ha bitat, becoming cut off by areas of unsuitable habitat from their now far-away relatives. The Lake Wales Ridge, a bove sea level since at least the Pliocene, has likely served as a refugium for such plant sp ecies when much of the rest of Florida was inundated or otherwise inhospitable. Several other endemics found on Tiger Creek and Saddle Blanket Preserves were considered by James ( 1961) also to be possible paleo-endemics, whose distributions suggest they had been reduced to central ridge refugia durin g high sea-level stands, but have now expanded back somewhat from these refugia. These include Asclepias feayi, Garberia heterophylla, Hartwrightia floridana (a wetland species), and the genus Phoebanthus, 68


comprised of two endemic Fl orida species, one of which, P. grandiflorus, is found on the preserves. A study of insect endemism on the Florida scrub ridges (Deyrup 1990) reveals patterns similar to those seen with plants: some species have western affinities, and the highest diversity of endemics occurs on the southern Lake Wales Ridge, with endemic diversity decreasing northwards on the ridges. Changes in climate asso ciated with glacial cooling episodes may have reduced biodiversity on the more northern pa rts of the peninsular ridge system. Although people speak of rare species as ge nuine entities in natu re, their designation depends upon taxonomic judgments made by human beings. A rare species attracts admiration and attention, as well as efforts directed toward its preservation, ye t one taxonomic opinion can change its status, reducing it to a mere synonym, relegating it overnight to the quotidian obscurity of the unremarkable and commonplace. A species collected at Saddle Blanket for this study provides a case in point. Conradina brevifolia, first described by Shinners (1962), is one of several shrubby mints endemic to Florida scrubs Over the decades it has enjoyed both state and federal protection as an endangered spec ies of extremely narrow distribution (Polk and Highlands Counties). But Wunderlin (1982) incl uded it within the more widespread and nonendemic C. canescens, where it remains, at the time of this writing, in certain widely consulted references ( i.e., Wunderlin and Hansen 2003, 2007). Despite this taxonomic ambiguity, its listed status as a protected endemic has not changed (Coile and Garland 2003; USFWS 1996, 2007). Meanwhile, systematic / phylogenetic work on Fl orida scrub mints supports the recognition of C. brevifolia as a separate species (Edwards 2007). This rare endemic scrub mint will likely keep its protected status, and its place in the public awareness. Two othe r rare taxa collected in this study also illustrate this issue. Crataegus lepida, currently included by some taxonomists within 69


the more common C. michauxii, and a scrub variant of Sideroxylon tenax, once recognized as Bumelia lacuum, await further study to clarify their taxonomic status (see Chapter 4). The existence of rare endemic species, and thei r evolution and persiste nce, are particular facets of the larger questions of evolutionary bi ology, so the study of rare species will continue to be important (Stebbins 1980). Molecular studi es, in particular, are beginning to enrich our understanding of rare plants, and of endemism in general. Many molecular studies have been directed toward assessing genetic variability with in populations of rare taxa, with an eye toward conservation planning ( e.g., Dolan et al. 1995; Evans et al. 2000; Lienert et al. 2002). A study of the genus Prunus in North America using chloroplast DNA revealed that the closest living relative of the scrub endemic Prunus geniculata is P. maritima (Shaw and Small 2005). And in a study of a type that has great potential for el ucidating processes leading to endemism, Maskas and Cruzan (2000) used chloro plast DNA and phylogeographic anal yses to infer historical processes leading to the diversification of a complex of morphotypes of Piriqueta cistoides subsp. caroliniana in Florida and the Bahamas. More st udies such as these are needed. As human development continues to destroy the habita ts of rare species, the need for systematic research becomes ever more urgent. It is hoped that this study will have contributed in some small way to that ongoing effort. 70


Table 3-1. Endemic vascular pl ant species found on Tiger Creek and Saddle Blanket preserves that are restricted to Flor ida peninsular scrubs or sc rub/sandhill ecotones. Species marked with an asterisk are restricted to the Lake Wales, Lake Henry, and Winter Haven Ridges. Dashes indicate no listing or designation available. Listed statusb Species Preservea FNAI FL USA Rarityc Asclepias curtissii SB G3/S3 E 8.0 *Bonamia grandiflora TC, SB G3/S3 E T 7.0 *Chionanthus pygmaeus TC, SB G3/S3 E E 6.0 *Clitoria fragrans TC G3/S3 E T 3.5 *Conradina brevifolia SB G2/S2 E E 7.5 *Crotalaria avonensis SB G1/S1 E E 4.5 Eriogonum longifolium var. gnaphalifolium TC, SB G4T3/S3 E T 8.5 Garberia heterophylla TC, SB T 10.0 *Hypericum cumulicola SB G2/S2 E E 7.5 Lechea cernua SB G3/S3 T 10.0 *Liatris ohlingerae SB G3/S3 E E 5.5 *Nolina brittoniana TC, SB G2/S2 E E 7.0 *Paronychia chartacea TC, SB G2/S2 E T 8.0 Polygala lewtonii TC G3/S3 E E 4.5 *Polygonella basiramia SB G3/S3 E E 8.0 *Polygonella myriophylla SB G3/S3 E E 11.0 *Prunus geniculata TC, SB G3/S3 E E 8.0 *Schizachyrium niveum TC, SB G1/S1 E MC 7.0 Stylisma abdita TC E 7.0 *Warea carteri TC G3/S3 E E 6.5 *Ziziphus celata TC (intro) G1/S1 E E 1.5 Carya floridana TC, SB Ilex opaca var. arenicola TC, SB Osmanthus megacarpus TC Persea humilis TC, SB Quercus inopina TC, SB Sabal etonia TC, SB aPreserve: TC = Tiger Creek; SB = Saddle Blanket. bStatus key: FNAI: Gx/Sx = Global and State endangerment rankings for Special Element sp ecies of the Florida Natural Areas Inventory, from 1 (critically imperiled) to 5 (demonstrably secure). T = rank for taxonomic subgroup. FL: E, T = Listed by the State of Florida as Endange red or Threatened. USA: E, T = Protected by federal law as Endangered or Threatened. MC = federally designated as being of Management Concern. (Chafin 2000, Coile and Garland 2003, Christman and Judd 1990). cRarity: Numerical ratings denoting relative rarity of listed species: 1 = most rare, 12 = least rare (Ward et al. 2003). 71


72 Table 3-2. Endemic vascular pl ant species found on Tiger Creek and Saddle Blanket preserves that are not restricted to peninsular scrubs. Listed statusb Species Preservea FNAI FL USA Rarityc Carex vexans Centrosema arenicola TC G2/S2 E 8.0 Dichanthelium chamaelonche subsp. breve SB Hartwrightia floridana TC G2/S2 T 9.0 Lechea divaricata TC G2/S2 E 7.0 Lilium catesbaei TC T 12.0 Matelea floridana TC E 7.0 Panicum abscissum TC, SB G3/S3 E MC 8.5 Arnoglossum floridanum TC Asclepias feayi TC Asimina obovata TC, SB Asimina reticulata TC, SB Berlandiera subacaulis TC Callisia ornata TC Carphephorus odoratissimus var. subtropicanus SB Chapmannia floridana TC, SB Chrysopsis subulata TC Crataegus lepida TC Crocanthemum nashii TC, SB Croton glandulosus var. floridanus TC Palafoxia feayi TC, SB Phoebanthus grandiflorus TC, SB Piptochaetium avenacioides TC, SB Polygala rugelii TC, SB Polygonella robusta TC, SB Rhynchosia cinerea TC, SB Scutellaria arenicola TC, SB Tillandsia simulata TC, SB aPreserve: TC = Tiger Creek; SB = Saddle Blanket. bStatus key: FNAI: Gx/Sx = Global and State endangerment rankings for Special Element sp ecies of the Florida Natural Areas Inventory, from 1 (critically imperiled) to 5 (demonstrably secure). T = rank for taxonomic subgroup. FL: E, T = Listed by the State of Florida as Endange red or Threatened. USA: E, T = Protected by federal law as Endangered or Threatened. MC = federally designated as being of Management Concern. (Chafin 2000, Coile and Garland 2003, Christman and Judd 1990). cRarity: Numerical ratings denoting relative rarity of listed species: 1 = most rare, 12 = least rare (Ward et al. 2003).


CHAPTER 4 ANNOTATED LIST OF VASCULAR PLANTS Just as this floristic inventory represents a momentary snapshot of a dynamic flux of organisms and environments, so also does its pr oduct a list of plant names represent a snapshot of current taxonomy, itself a some what dynamic and changing thing. A taxonomic classification aims for stability, for too much ch ange would render it useless, especially for information retrieval purposes. Yet who can deny th at the best system of classification is one that reflects our best underst anding of phylogeny, based upon all the evidence we can muster (see Judd et al. 2008)? New eviden ce, especially from molecular phylogenetics, is pouring in, nudging inertia-bound plant taxonomy into a reluctant state of motion. The following species list is therefore a frozen frame, in which I have hopefully captured a sense of our current understanding. It should be kept in mind that each species name is a taxonomic statement, representing a hypothesis of lin eage delimitation among the organisms being studied, while each generic name is a hypothesis of phylogenetic relationships. As our understanding improves, species and generic names will continue to change. For family names and circumscriptions, I have followed the Angiosperm Phylogeny Group (APG II 2003, Stevens 2007) for flowering plants. For ferns, I have followed Smith et al. (2006), and for lycophytes and gymnosperms, Kram er and Green (1990). For species names, (including common names), I have followed, wherev er possible, those given in the Flora of North America (Flora of North America Editorial Committee 1993-2007). For taxa not covered in the Flora of North America, I have follo wed Wunderlin and Hansen (2003, 2004). In some cases I have deviated from the above authorities, citing recent (o r more convincing) sources in the species list. Author abbreviations follow Brummitt and Powell (1992). Taxa are listed alphabetically by family, then by genus and species. 73


Each species name is followed by an author abbreviation, a preferred common name, one or more habitat designations denoting where th e species occurs, an a bundance designation, and specimen numbers. Habitat desi gnations are as follows: SC (scrub, including yellow sand scrub); SF (scrubby flatwoods); SH (sandhill); XH (xeric hammock); MF (mesic flatwoods); WF (wet flatwoods); BH (bayhead); SS (seepag e slope); DM (depression marsh or ephemeral pond); FP (floodplain wetlands); BW (blackwater stream); and RU (ruderal, or disturbed areas). Estimates of the abundance of each spec ies within these communities are based upon my observations of the plants in the field, and are therefore subjective. A bundance of occurrence is denoted by the following abbreviations and numeric al values: R (rare) 1-4 occurrences; I (infrequent) 5-9 occurrences; O (occasional) 10-24 occurrences; F (frequent) 25 or more occurrences; A (abundant) when a species is do minant in its community and occurs more or less continuously throughout. Specimen numbers given are my own collection numbers except where otherwise noted. Numbers beginning with TC denote specimens collected at Tiger Creek; those beginning with SB were collected at Saddle Blanket Preserve. When a different collectors specimen is cited, the Preserve is specified by (T C) or (SB) before the specimen number. Nonnative taxa are denoted by an asteri sk in front of the species name. For Tiger Creek Preserve, an unvouchered checkl ist of plant species was prepared in 1990, but the identity of the preparer is unknown. A number of species from that checklist were not collected in the present study and do not appear in the list below. But some of these species have been collected by others in areas near the Preserve containing comparable habitats; therefore it is reasonable to suppose that they might still occur on the Preserve. For this reason I mention them here: Baccharis glomeruliflora Pers., Carphephorus paniculatus (J.F. Gmel.) H. Hebert, Ceanothus microphyllus Michx., Cirsium horridulum Michx., Crinum americanum L., 74


Cynanchum scoparium Nutt., Digitaria ciliaris (Retz.) Koeler, Digitaria longiflora (Retz.) Pers., Eryngium aromaticum Baldwin, Eustachys petraea (Sw.) Desv., Fimbristylis caroliniana (Lam.) Fernald, Fraxinus caroliniana Mill., Lachnocaulon anceps (Walter) Morong, Luziola fluitans (Michx.) Terrell & H. Rob., Orontium aquaticum L., Panicum dichotomiflorum Michx., Panicum tenerum Beyrich ex Trin., Paspalum praecox Walter, Paspalum repens P.J. Bergius, Pediomelum canescens (Michx.) Rydb., Quercus pumila Walter, Rhynchospora cephalantha A. Gray, Rhynchospora divergens Chapm. ex M.A. Curtis, Rhynchospora pusilla Chapm. ex M.A. Curtis, Rubus trivialis Michx., Sarcostemma clausum (Jacq.) Schult., Stylisma patens (Desr.) Myint, Thalia geniculata L., Thelypteris palustris Schott, and Vittaria lineata (L.) Sm. ACANTHACEAE Ruellia carolinensis (J.F.Gmel.) Steud.: Wild petunia. SH, XH O. TC111. ADOXACEAE Sambucus nigra L. subsp. canadensis (L.) Bolli: Elderberry. FP, Wetland edges O. TC044. Viburnum nudum L.: Possum haw. BH, FP O. TC200, TC465, SB251, SB277, SB293. AGAVACEAE Yucca flaccida Haw.: Adam's needle. SH, SC, XH F. TC346, SB156. (Ward 2006). ALISMATACEAE Sagittaria isoetiformis J.G. Sm.: Quillwort arrowhead. DM O. TC401, SB284. Sagittaria lancifolia L. subsp. lancifolia : Bulltongue arrowhead. FP F. TC252. Sagittaria latifolia Willd.: Duck potato. FP O. TC546. ALTINGIACEAE Liquidambar styraciflua L.: Sweetgum. FP F. TC197. AMARANTHACEAE Alternanthera philoxeroides (Mart.) Griseb.: Alligator weed. BW I. TC603. Dysphania ambrosioides (L.) Mosyakin & Clemants: Mexican tea. RU F. TC528, SB100. Froelichia floridana (Nutt.) Moq.: Cottonweed. SH, RU O. TC169, SB102. Gomphrena serrata L.: Globe amaranth. RU F. TC226. Iresine diffusa Humb. & Bonpl. ex Willd.: Juba's bush. RU O. TC540. ANACARDIACEAE Rhus copallinum L.: Winged sumac. SH, SC, MF, XH F. TC198, SB148. Toxicodendron radicans (L.) Kuntze: Poison ivy. FP, XH, RU F. TC397, SB235. ANNONACEAE Asimina obovata (Willd.) Nash: Flag pawpaw. SC, SH, SF O. TC074, SB276, SB287. Asimina parviflora (Michx.) Dunal: Smallflower pawpaw. FP O. TC556. 75


Asimina pygmaea (W. Bartram) Dunal: Dwarf pawpaw. SH R. SB012. Asimina reticulata Shuttlew. ex Chapm.: Netted pawpaw. SH, SC F. TC070, SB005. APIACEAE Centella asiatica (L.) Urb.: Coinwort. DM, FP F. TC406, TC500, SB225. Cicuta maculata L.: Water hemlock. BW, FP F. TC191. Eryngium prostratum Nutt. ex DC.: Creeping eryngo. FP I. TC387. Oxypolis filiformis (Walter) Britton var. filiformis : Water dropwort. FP I. TC445. Ptilimnium capillaceum (Michx.) Raf.: Mock bishopsweed. FP O. TC386. APOCYNACEAE Asclepias curtissii A. Gray: Curtiss's milkweed. SC, SF F. SB046. Asclepias feayi Chapm. ex A.Gray: Florida milkweed. SH I. TC112. Asclepias humistrata Walter: Sandhill milkweed. SH I. TC509. Asclepias pedicellata Walter: Savannah milkweed. MF I. SB294. Asclepias tomentosa Elliott: Velvetleaf milkw eed. SH I. (TC) Abbott 22681. Asclepias tuberosa L.: Butterfly milkweed. SH, XH O. TC106, SB105. Catharanthus roseus (L.) G. Don: Madagascar periwinkle. RU O. TC524. Matelea floridana (Vail) Woodson: Florida milkvine. XH O. TC218, TC249. AQUIFOLIACEAE Ilex ambigua (Michx.) Torr. var. ambigua: Sand holly. SC, XH O. TC133, TC258, SB049. Ilex cassine L.: Dahoon holly. FP, BH F. TC012, SB174. Ilex glabra (L.) A.Gray: Gallberry. MF, WF, DM F. TC029, SB038, SB078. Ilex opaca Aiton var. arenicola (Ashe) Ashe: Scrub holly. SC F. TC476, SB144. ARACEAE Arisaema triphyllum (L.) Schott: Jack-in-the-pulpit. FP I. TC557. Landoltia punctata (G. Mey.) Les & D.J. Crawford: Dotted duckweed. BW, FP F. TC552. (Les and Crawford 1999). Lemna aequinoctialis Welw.: Lesser duckweed. BW, FP F. TC569, SB265. Peltandra sagittifolia (Michx.) Morong: Sp oonflower. FP O. TC503. Peltandra virginica (Michx.) (L.) Schott: Green arum. FP O. TC392. Pistia stratiotes L.: Water lettuce. BW O. TC547. [Experts disagree as to whether this species is native or introduced. William Bartram (1791) observed well-established populations in Florida.] ARALIACEAE Hydrocotyle umbellata L.: Marsh pennywort. DM O. TC499, SB298. Hydrocotyle verticillata Thunb. var. verticillata : Whorled marsh pennywort. FP O. TC422. ASTERACEAE Ageratina jucunda (Greene) Clewell & Wooten: Hammock snakeroot. XH, SH O. TC351. Ambrosia artemisiifolia L.: Ragweed. RU F. TC166. Ampelaster carolinianus (Walter) G.L. Nesom: Climbing aster. FP O. TC349. Arnoglossum floridanum (A. Gray) H. Rob.: Florida Indian-plantain. SH O. TC174. Baccharis halimifolia L.: Groundsel tree. DM, RU, Wetland edges F. TC031, TC459. Balduina angustifolia (Pursh) B.L. Rob.: Honeycomb-head. SH, XH F. TC007, SB203. 76


Berlandiera subacaulis (Nutt.) Nutt.: Greeneyes. SH O. TC120. Bidens alba (L.) DC. var. radiata (Sch. Bip.) R.E. Ballard ex Me lchert: Beggarticks. RU F. TC227. (Ballard 1986). Bidens laevis (L.) Britton et al.: Bu r marigold. FP O. TC430. Bidens mitis (Michx.) Sherff: Smallfruit beggarticks. DM I. SB250. Carphephorus corymbosus (Nutt.) Torr. & A. Gray: Florida paintbrush. SH, XH F. TC018, SB186. Carphephorus odoratissimus (J.F. Gmel.) H. Hebert var. subtropicanus (DeLaney et al.) Wunderlin & B.F. Hansen: False vanillaleaf. MF R. SB215. Chrysopsis scabrella Torr. & A. Gray: Coastalplain goldenaster. SH O. TC451. Chrysopsis subulata Small: Scrubland goldenaster. MF I. TC523. Cirsium nuttallii DC.: Nuttall's thistle. RU O. TC381. Conyza canadensis (L.) Cronquist var. pusilla (Nutt. ) Cronquist: Dwarf horseweed. RU F. TC019, SB142. (Nesom 1990). Eclipta prostrata (L.) L.: False daisy. FP, RU O. TC202. Elephantopus carolinianus Raeusch.: Carolina elephant's foot. FP I. TC309. Elephantopus elatus Bertol.: Tall elephant's foot. SH, XH F. TC206, SB121. Emilia fosbergii Nicolson: Florida tasselflower. RU O. TC442, SB161. Emilia sonchifolia (L.) DC.: Lilac tasselflower. RU O. TC224. Erechtites hieraciifolius (L.) Raf. ex DC. var. hieraciifolius : Fireweed. FP, DM, MF F. TC352, SB032, SB220. Erigeron quercifolius Poir.: Oakleaf fleabane. RU F. TC383. Erigeron vernus (L.) Torr. & A. Gray: Early whitetop fleabane. DM I. TC403. Eupatorium capillifolium (Lam.) Small ex Porter & Britton: Dog-fennel. DM, FP, RU F. TC348, SB224. Eupatorium compositifolium Walter: Yankeeweed. SH O. TC371, SB205. Eupatorium leptophyllum DC.: False fennel. DM I. TC013, SB211. Eupatorium mohrii Greene: Mohr's thoroughwort. MF, DM O. TC093, SB306. Eupatorium rotundfolium L. var. rotundifolium : Roundleaf thoroughwort. DM, MF I. TC254, SB209. Euthamia caroliniana (L.) Greene ex Porter & Britton: Flattop goldenrod. DM, MF, SH F. TC021, SB222. Gamochaeta antillana (Urban) Anderberg: Narrowl eaf everlasting. DM I. TC426. Gamochaeta pensylvanica (Willd.) Cabrera: Pennsylvania everlasting. FP I. TC491. Gamochaeta purpurea (L.) Cabrera: Purple cudweed. FP I. TC471. Garberia heterophylla (W. Bartram) Merr. & F. Harper: Garberia. SH, SC, XH F. TC027. Hartwrightia floridana Gray ex S. Wats.: Florid a hartwrightia. DM O. TC336. Helianthus angustifolius L.: Swamp sunflower. MF I. TC338. Heterotheca subaxillaris (Lam.) Britton & Rusby subsp. subaxillaris : Camphorweed. RU I. SB204. Hieracium gronovii L.: Queen-devil. SH R. SB187. Hieracium megacephalon Nash: Coastalplain hawkweed. SH I. TC071, SB145, SB279. Krigia virginica (L.) Willd.: Virginia dwarf dandelion. MF R. TC378. Liatris chapmanii Torr. & A. Gray: Chapman's blazing star. SH O. TC242. Liatris ohlingerae (S.F. Blake) B.L. Rob.: Scrub blazing-star. SC F. SB111. 77


Liatris laevigata Nutt.: Blazing star. SH, XH F. TC005, TC294, SB185. Liatris tenuifolia Nutt.: Blazing star. SH, XH O. TC315. Lygodesmia aphylla (Nutt.) DC.: Rose-rush. SH O. TC347, SB289. Mikania scandens (L.) Willd.: Climbing hempvine. FP, BH F. TC245, TC468. Oclemena reticulata (Pursh) G.L. Nesom: Pinebarren aster. MF I. TC497, SB286. Palafoxia feayi A. Gray: Feay's palafox. SH, XH, SF, SC F. TC003, SB035, SB080. Palafoxia integrifolia (Nutt.) Torr. & Gray: Coasta l-plain palafox. SH I. TC331. Pectis prostrata Cav.: Spreading cinchweed RU, O. TC612, TC633. Phoebanthus grandiflorus (Torr. & A.Gray) S.F. Blake: Florid a false sunflower. SH O. TC150, SB129. Pityopsis graminifolia (Michx.) Nutt. var. aequilifolia F.D. Bowers & Semple: Goldenaster. SH, XH F. TC004, SB202. Pluchea baccharis (Mill.) Pruski: Marsh fleabane. DM F. TC141. Pluchea foetida (L.)DC.: Stinking camphorweed. DM I. TC207. Pluchea odorata (L.) Cass. var. odorata : Annual marsh fleabane. FP O. TC272, TC369, TC427, SB301. Pterocaulon pycnostachyum (Michx.) Elliott: Blackroot. SH, MF O. TC116, SB055. Sericocarpus tortifolius (Michx.) Nees: White-top aste r SH, XH, MF O. TC025, TC255, SB221. Solidago fistulosa Mill.: Pinebarren goldenrod. DM, MF O. TC020, SB201. Solidago odora Aiton var. chapmanii (A.Gray) Cronquist: Chapman's goldenrod. SH, XH F. TC168, SB109. Symphyotrichum dumosum (L.) G.L. Nesom: Ricebutton aster. SH I. TC332. Symphyotrichum simmondsii (Small) G.L. Nesom: Simmonds's aster. Wetland edge I. TC365. Verbesina virginica L.: White crownbeard. XH F. TC312. Youngia japonica (L.) DC.: Oriental false hawksbeard. RU O. TC441. BIGNONIACEAE Catalpa bignonioides Walter: Southern catalpa. SH R. TC593. (Probably resprouted from orna mental planting by a previous property owner.) Pyrostegia venusta (KerGawl.) Miers: Fl ame vine. FP R. TC046. (Probably extirpated subsequent to collection of specimen.) BLECHNACEAE Blechnum serrulatum Rich.: Toothed midsorus fern. BH, FP F. TC035, SB233. Woodwardia areolata (L.) T. Moore: Netted chain fern. BH, FP O. TC094, SB256. Woodwardia virginica (L.) Sm.: Virginia chain fern. BH, FP, WF F. TC045, TC222, TC390, SB232. BRASSICACEAE Lepidium virginicum L. : Poorman's pepper. RU F. TC483. Polanisia tenuifolia Torr. & A. Gray: Slenderleaf clammyweed. SH, XH F. TC109, SB300. Warea carteri Small: Carter's Warea. SF I. TC316. BROMELIACEAE Tillandsia recurvata (L.) L.: Ball-moss. SH, XH, SC, FP F. TC205, SB140. Tillandsia setacea Sw.: Southern needleleaf. FP F. TC057, SB139. Tillandsia simulata Small: Florida air plant. BH, FP F. TC201. 78


Tillandsia usneoides (L.) L.: Spanish moss. SH, FP, XH, SC F. TC204, SB141. Tillandsia utriculata L.: Giant air plant. FP, BH I. TC377, SB183. CACTACEAE Opuntia humifusa (Raf.) Raf. var. humifusa : Prickly-pear. SH, SC, XH F. TC480, SB292. Opuntia humifusa (Raf.) Raf. var. ammophila (Small) L.D. Benson: Prickly-pear. SH, SC, XH F. TC600. CAMPANULACEAE Lobelia paludosa Nutt.: White lobelia. DM I. TC090. CANNACEAE Canna flaccida Salisb.: Bandana-of-the-E verglades. FP O. TC223, TC470. CARYOPHYLLACEAE Paronychia americana (Nutt.) Fenzl ex Walp.: American nailwort. SC, RU F. SB051, SB091, SB208. Paronychia chartacea Fernald var. chartacea: Papery whitlow-wort SH R TC009. SB095. Paronychia hemiarioides (Michx.) Nutt.: Coastalplain nailwort. SH I. TC525. Stipulicida setacea Michx. var. setacea : Pineland scalypink. SH, SC, XH F. TC040, SB021. CHRYSOBALANACEAE Licania michauxii Prance: Gopher apple. SH, SC, XH F. TC099, SB042. CISTACEAE Crocanthemum corymbosum (Michx.) Britton: Pinebarren frostweed. SH O. TC477, SB025. (Arrington and Kubitzki 2003). Crocanthemum nashii (Britton) Barnhart: Florida scru b frostweed. SH, SC F. TC088, SB014. (Arrington and Kubitzki 2003). Lechea cernua Small: Nodding pinweed. SC O. SB119, SB191. Lechea deckertii Small: Deckert's pinweed. SH O. TC156, SB108, SB128. Lechea divaricata Shuttlew. ex Britton: Drysand pinweed. MF R. (TC) Christman 1933. Lechea sessiliflora Raf.: Pineland pinweed. SC, SH O. TC299, TC310, TC341, SB192. Lechea torreyi (Chapm.) Legg. ex Britton: Pi edmont pinweed. MF O. TC613, SB147, SB212. COMMELINACEAE Callisia ornata (Small) G.C. Tucker: Florida scrub roseling. SC, SH O. TC144, SB288. Commelina diffusa Burm. f. var. diffusa : Common dayflower. RU, FP O. TC439. Commelina erecta L.: Whitemouth dayflower. SH, XH, SF F. TC001, SB024. Tradescantia roseolens Small: Longleaf spiderwort. SH, XH I. TC510, SB082. CONVOLVULACEAE Bonamia grandiflora (A. Gray) Hallier f.: Scrub morning glory. SC, SH O. TC508, SB040. Calystegia sepium (L.) R. Br. subsp. limnophila (Greene) Brummitt: Hedge false bindweeed. FP I. TC478. Dichondra carolinensis Michx.: Carolina ponyfoot. RU, FP O. TC385. Ipomoea pandurata (L.) G. Mey.: Man-of-the-earth. XH R. TC267. 79


Stylisma abdita Myint: Hidden dawnflower. SH I. TC513. Stylisma villosa (Nash) House: Hairy dawnflower. SH, XH, SC F. TC006, SB041, SB136. CORNACEAE Cornus foemina Mill.: Swamp dogwood. FP F. TC449, TC464. Nyssa biflora Walter: Swamp tupelo. FP F. TC160, SB236. (Ward 2001). CRASSULACEAE Kalanchoe delagoensis Eckl. & Zeyh.: Chandelier plant. RU I. TC626. CUCURBITACEAE Cucumis anguria L.: Gooseberry gourd. FP I. TC467. Momordica charantia L.: Balsampear. RU F. TC376. CYPERACEAE Bulbostylis barbata (Rottb.) C.B. Clarke: Waterg rass. SH, XH, MF F. TC587. Bulbostylis ciliatifolia (Elliott) Fernald: Capillary hairsedge. SH, XH F. TC011, TC129. Bulbostylis stenophylla (Elliott) C.B. Clarke: Sandyfield hairsedge. SC O. SB150. Bulbostylis warei (Torr.) C.B. Clarke: Ware's hairsedge. SH F. TC119, TC130. Carex comosa Boott: Longhair sedge. FP O. TC474. Carex longii Mack.: Long's sedge. DM O. SB303. Carex vexans F.J. Herm.: Florida hammock sedge. FP O. TC493. Cladium jamaicense Crantz: Sawgrass. FP O/F. TC507. Cyperus croceus Vahl: Baldwin's flatsedge. SH, XH, MF F. TC281. Cyperus lecontei Torr. ex Steud.: Leconte's flatsedge. DM O. TC185, SB018, SB127. Cyperus retrorsus Chapm.: Pinebarren flatsedge. SH, XH, MF F. TC148, TC531, SB029, SB067. Cyperus tetragonus Elliott: Fourangle flatsedge. FP O. TC248. Cyperus virens Michx.: Green flatsedge. FP F. TC416, TC461. Eleocharis baldwinii (Torr.) Chapm.: Road-grass. DM F. TC318, TC405, TC431, SB162. Eleocharis microcarpa Torr.: Smallfruit spikerush. DM I. TC319. Fimbristylis puberula (Michx.) Vahl: Hairy fimbry. DM F. TC117, TC414. Fuirena scirpoidea Michx.: Southern umbrella sedge. DM, Pond margins F. TC286, SB028. Rhynchospora chapmanii M.A. Curtis: Chapman's beaksedge. DM O. TC233, TC340. Rhynchospora ciliaris (Michx.) C. Mohr: Fringed beaksedge. DM, SS, MF F. TC114, TC280. Rhynchospora corniculata (Lam.) A. Gray: Shortbristle horned beaksedge. FP F. TC415, TC436, TC458. Rhynchospora fascicularis (Michx.) Vahl: Fascicled beak sedge. DM, BH, FP F. TC98, TC359, TC622, SB126, SB167, SB259. Rhynchospora inundata (Oakes) Fernald: Narrowfruit horned beaksedge. DM F. TC147, TC597. Rhynchospora latifolia (Baldwin) W.W. Thomas: Giant white-top sedge. DM O. TC208, TC410. Rhynchospora megalocarpa A. Gray: Sandyfield beaksedge. SH, XH, SC F. TC083, TC118, SB022, SB154. 80


Rhynchospora microcarpa Baldwin ex A. Gray: Southern beaksedge. DM, MF F. TC522. Rhynchospora microcephala (Britton) Britton ex Small: Bunched beaksedge. DM O. SB304. Rhynchospora miliacea (Lam.) A. Gray: Millet beaksedge. FP F. TC469, TC567. Rhynchospora pineticola C.B. Clarke: Pinebarren beaksedge. MF O. SB059. Rhynchospora plumosa Elliott: Plumed beaksedge. MF I. SB068, SB280. Rhynchospora rariflora (Michx.) Elliott: Fewflower beaksedge. DM F. TC632. Rhynchospora scirpoides (Torr.) A. Gray: Longbeak beaksedge. DM I. TC517. Rhynchospora tracyi Britton: Tracy's beaksedge. DM F. TC515. Scirpus cyperinus (L.) Kunth: Woolgr ass. FP I. TC566. Scleria muehlenbergii Steud.: Nutrush. SS, MF, FP O. TC279. Scleria reticularis Michx.: Netted nutrush. DM F. TC516. Scleria triglomerata Michx.: Tall nutgrass. MF, WF, FP F. TC151, SB151, SB241. DENNSTAEDTIACEAE Hypolepis repens (L.) C. Presl: Creeping bramble fern. BH R. TC535. Pteridium aquilinum (L.) Kuhn var. pseudocaudatum (Clute) Clute ex A. Heller: Tailed bracken fern. SH, XH, MF F. TC145, SB034, SB063. DROSERACEAE Drosera capillaris Poir.: Pink sundew. DM, WF F. TC053. DRYOPTERIDACEAE Dryopteris ludoviciana (Kunze) Small: Southern wood fern. FP F. TC565. EBENACEAE Diospyros virginiana L.: Common persimmon. SC, XH, SF, MF F. TC178, SB123. ERICACEAE Bejaria racemosa Vent.: Tarflower. MF, SF F. TC110, SB048. Ceratiola ericoides Michx.: Florida rosemary. SC F. SB056. Gaylussacia dumosa (J. Kenn.) Torr. & A. Gray: Dwarf huckleberry. SH, MF O. TC060, SB003. Gaylussacia nana A. Gray: Blue huckleberry. MF F. TC079, TC621, SB004. (Luteyn et al. 1996). Lyonia ferruginea (Walter) Nutt.: Rusty lyonia. SC F. TC353, SB207. Lyonia fruticosa (Michx.) G.S. Torr.: Staggerbush. SF, SC, MF F. TC028, SB047. Lyonia ligustrina (L.) DC. var. foliosiflora (Michx.) Fernald: Maleberry. BH, WF O. TC162, TC505, TC594, SB227, SB239, SB296. Lyonia lucida (Lam.) K. Koch: Fetterbush. MF, WF, DM, SF F. TC008, TC051, SB009. Monotropa uniflora L.: Indianpipe. XH I. Christman 1930. Rhododendron viscosum (L.) Torr.: Swamp azalea. FP I. TC573, SB249, SB281. Vaccinium corymbosum L.: Highbush blueberry. BH, FP O. TC059, TC184, SB242. Vaccinium cf. corymbosum L.: Hybrid blueberry. MF I. TC617. Vaccinium darrowii Camp: Darrow's blueberry. SH, MF F. TC056, SB008. Vaccinium myrsinites Lam.: Shiny blueberry. SH, SF, MF F. TC050, SB274. Vaccinium stamineum L.: Deerberry. SH, SC, XH F. TC413. Vaccinium corymbosum x V. darrowii : Hybrid blueberry. MF, DM O. TC618, TC625. 81


ERIOCAULACEAE Eriocaulon decangulare L.: Pipewort. DM F. TC139. Lachnocaulon beyrichianum Sporl. ex Koern.: Southern bog button. DM I. TC521, SB291. Lachnocaulon minus (Chapm.) Small: Small's bog button. DM F. TC113, TC501, SB019. Syngonanthus flavidulus (Michx.) Ruhland: Hatpins. DM F. TC054, SB290. EUPHORBIACEAE Chamaesyce hirta (L.) Millsp.: Pillpod sandmat. RU F. TC443. (Some authors, e.g., Steinmann and Porter 2002, Park and E lisens 2000, point out that recognizing Chamaesyce renders Euphorbia paraphyletic, and favor including Chamaesyce within an expanded genus Euphorbia.) Chamaesyce hyssopifolia (L.) Small: Hyssopleaf sandmat. RU O. TC288. Chamaesyce maculata (L.) Small: Spotted sandmat. RU O. TC289. Chamaesyce ophthalmica (Pers.) D.G. Burch: Florida hammock sandmat. RU O. TC555. Chamaesyce prostrata (Aiton) Small: Prostrate sandmat. RU O. TC554. Cnidoscolus stimulosus (Michx.) Engelm. & A. Gray: Tread-softly. SH, SC O. TC072, SB077. Croton argyranthemus Michx.: Healing croton. SH O. TC062. Croton glandulosus L. var. floridanus (A.M. Ferguson) R.W. Long: Vente conmigo. SH R. TC287. Croton glandulosus L. var. glandulosus : Vente conmigo. XH I. TC534. Croton michauxii G.L. Webster: Michaux's croton. SH O. TC407, SB054. Euphorbia polyphylla Engelm. ex Chapm.: Lesser Florida spurge. SC F. SB053, SB149. Stillingia sylvatica L.: Queen's delight. SH F. TC122, SB033. Tragia urens L.: Wavyleaf noseburn. SH O. TC512, SB158. FABACEAE Abrus precatorius L.: Rosary pea. RU F. TC589. Albizia julibrissin Durazz.: Mimosa tree. RU R. TC538. Amorpha fruticosa L.: Bastard false indigo. SC, Streambanks O. TC131. Amorpha herbacea Walter: Clusterspike false indigo. SH I. (TC) Christman 1970. Apios americana Medik.: Groundnut. FP O. TC228. Centrosema arenicola (Small) F.J. Herm.: Sand butterfly pea. SH, SF O. TC588. Centrosema virginianum (L.) Benth.: Spurred butte rfly-pea. SH, XH F. TC190, SB130. Chamaecrista fasciculata (Michx.) Greene: Partridge-pea. SH, XH F. TC167, SB110. Chamaecrista nictitans (L.) Moench var. aspera (Muhl. ex Elliott) H.S. Irwin & Barneby : Sensitive pea. SH, MF, RU O. TC306, SB210. Chamaecrista nictitans (L.) Moench var. nictitans : Wild sensitive-pea. SH, XH, RU O. TC022. Chapmannia floridana Torr. & A. Gray: Alicia. SH, SC F. TC153, SB013. Clitoria fragrans Small: Pigeon-wing. SH, XH O. TC137. Crotalaria avonensis DeLaney & Wunderlin: Avon Park rattlebox. SC O. SB094. Crotalaria pallida Aiton var. obovata (G. Don) Polhill: Smooth rattlebox. RU O. TC024. 82


* Crotalaria retusa L.: Rattleweed. RU O. TC219. Crotalaria rotundifolia J.F. Gmel.: Rabbit bells. SH, XH O. TC082, SB159. Dalea carnea (Michx.) Poir. var. carnea : Whitetassels. MF O. SB196, SB302. Dalea feayi (Chapm.) Barneby: Feay's pr airie-clover. SF, SH I. TC313. Dalea pinnata (J.F. Gmel.) Barneby var. pinnata: Summer farewell. SH O. TC320. Desmodium floridanum Chapm.: Florida ticktrefoil. SH O. TC213, TC580. Desmodium incanum DC.: Zarzabacoa co mun. RU O. TC212. Desmodium tortuosum (Sw.) DC.: Dixie ticktrefoil. RU I. TC213. Desmodium triflorum (L.) DC.: Threeflower ticktrefoil. RU O. TC270. Ente rolobium contortisiliquum (Vell.) Morong: Earpod tree. Former homesite. Single large tree, a few saplings. TC456. Erythrina herbacea L.: Coral bean. XH O. TC244. Galactia elliottii Nutt.: Elliott's milkpea. SH, SC, MF F. TC108, SB017. Galactia regularis (L.) Britton et al.: Eastern m ilkpea. SH, SC, XH F. TC203, SB132. Indigofera caroliniana Mill.: Carolina indigo. SH, SC, XH F. TC123, SB090. Indigofera hirsuta L.: Hairy indigo. RU O. TC256, TC368. Lespedeza hirta (L.) Hornem.: Hairy bush-clover. SH I, TC350. Lupinus diffusus Nutt.: Sky-blue lupine. SH F. TC061. Macroptilium lathyroides (L.) Urb.: Wild bushbean. SH, XH I. TC581. Medicago lupulina L.: Black medick. RU O. TC485. Mimosa quadrivalvis L. var. angustata (Torr. & A. Gray) Barneby: Sensitive briar. SH O. TC579. Mimosa quadrivalvis L. var. floridana (Chapm.) Barneby: Fla. sensitive briar. SC, SF O. TC520, SB076. Mimosa strigillosa Torr. & A. Gray: Powderpuff. RU I. TC601. Phaseolus polystachios (L.) Britton et al. var. sinuatus (Nutt. ex Torr. & A. Gray) Marechal et al.: Thicket bean. SH I. TC269. Rhynchosia cinerea Nash: Brown-haired snoutbean. SH O. TC121, SB115. Tephrosia chrysophylla Pursh: Scurf hoarypea. SH F. TC155, SB143. FAGACEAE Quercus chapmanii Sarg.: Chapman's oak. SH, SF, SC F. TC175, SB011, SB057, SB217. Quercus cf. pumila Walter: Running oak. MF I. SB216. (Ward 2007). Quercus geminata Small: Sand live oak. XH A; SH, SC, SF F. TC125, SB062. Quercus cf. hemisphaerica W. Bartram ex Willd.: Laurel oak hybrid. MF I. TC620. Quercus incana W. Bartram: Bluejack oak. SH F. TC127, SB282. Quercus inopina Ashe: Scrub oak. SF, SC F. TC126, SB044. Quercus laevis Walter: Turkey oak. SH A. TC075, SB093. Quercus laurifolia Michx.: Laurel oak. FP F. TC194. Quercus minima (Sarg.) Small: Dwarf live oak. MF F. TC195, SB195. Quercus myrtifolia Willd.: Myrtle oak. SC, SF, SH, XH F. TC077. Quercus nigra L.: Water oak. FP F. TC196, SB066, SB124, SB246, SB267. Quercus virginiana Mill.: Live oak. High creek banks F. TC176. Quercus laevis x Q. incana : Hybrid oak. MF I. TC619. GELSEMIACEAE Gelsemium sempervirens (L.) Aiton f.: Yellow je ssamine. MF, FP O. TC041. 83


GENTIANACEAE Sabatia brevifolia Raf.: Shortleaf rosegentian. MF, WF, DM O. TC032, SB172. Sabatia difformis (L.) Druce: Lanceleaf rosegentian. MF O. TC135. Sabatia grandiflora (A.Gray) Small: Marsh-pink. DM F. TC402, SB075. GERANIACEAE Geranium carolinianum L.: Cranesbill. RU O. TC066. HAEMODORACEAE Lachnanthes caroliana (Lam.) Dandy: Carolina redroot. DM, WF F. TC142, SB103. HALORAGACEAE Proserpinaca pectinata Lam.: Mermaid weed. DM F. TC389. HYPERICACEAE Hypericum brachyphyllum (Spach) Steud.: Coas talplain St. John's wort. DM O. TC091. Hypericum cistifolium Lam.: Roundpod St. John's wort. FP, WF, DM O. TC023, SB071. Hypericum cumulicola (Small) W.P. Adams: Highlands scrub St. John's wort. SC O. SB092. Hypericum fasciculatum Lam.: Sandweed. DM F. TC596, SB027. Hypericum gentianoides (L.) Britton et al.: Pineweed. DM O. TC230. Hypericum hypericoides (L.) Crantz: St. Peter's Wort. FP O. TC370, TC435. Hypericum mutilum L.: Dwarf St. John's wort. FP O. TC419, TC488. Hypericum mytrifolium Lam.: Myrtleleaf St. John's wort. DM O. TC092. Hypericum reductum (Svenson) W.P. Adams: Atlantic St. John's wort. SH O. TC157, SB036, SB096, SB173. Hypericum tetrapetalum Lam.: Fourpetal St. John's wort. MF I. SB097. HYPOXIDACEAE Hypoxis juncea Sm.: Fringed yellow star grass. DM O. TC337, SB086. IRIDACEAE Sisyrinchium nashii E.P.Bicknell: Nash's blue-eyed grass. SC I. SB213. Sisyrinchium xerophyllum Greene: Scrub blue-eyed grass. SC, SH, XH O. TC261, SB175, SB194. ITEACEAE Itea virginica L.: Sweetspire. FP, BH F. TC068, TC163, SB226. JUGLANDACEAE Carya floridana Sarg.: Scrub hickory. SC, SH F. TC085, TC475, SB006. Carya glabra (Mill.) Sweet: Pignut hickory. XH O. TC078, TC578. JUNCACEAE Juncus effusus L. subsp. solutus (Fernald & Wiegand) Hmet-Ahti: Soft rush. FP O. TC171, SB258. Juncus marginatus Rostk.: Grassleaf rush. DM, MF O. TC210, TC623, SB116, SB247. Juncus megacephalus M.A. Curtis: Bighead rush. FP, DM I. TC172. Juncus repens Michx.: Lesser creepi ng rush. WF I. TC502. Juncus scirpoides Lam. : Needlepod rush. DM O. TC164, SB064, SB117. KRAMERIACEAE Krameria lanceolata Torr.: Sandspur. SH O. TC103. LAMIACEAE Callicarpa americana L.: Beautyberry. SC F. TC132, SB050. 84


Conradina brevifolia Shinners: Short-leaved rosemary. SC F. SB001. (Edwards 2007; Shinners 1962). Hyptis alata (Raf.) Shinners: Clustere d bushmint. WF I. TC575. Hyptis mutabilis (Rich.) Briq.: Tropical bushmint. RU O. TC158. Piloblephis rigida (W. Bartram ex Benth.) Raf.: Wild pennyroyal. MF O. SB248. Salvia lyrata L.: Lyreleaf sage. RU F. TC447. Scutellaria arenicola Small: Florida scrub scullcap. SH, XH O. TC533, SB104. Trichostema dichotomum L.: Blue curls. SC, SH O. TC042, TC333, SB184. LAURACEAE Persea borbonia (L.) Spreng.: Red bay. XH, MF O. TC246. Persea humilis Nash: Silk bay. SC F; SH O. TC134, SB043. (Wofford 1973; Christman & Judd 1990) Persea palustris (Raf.) Sarg.: Swamp bay. BH, FP, WF F. TC183, SB58, SB107. LENTIBULARIACEAE Pinguicula pumila Michx.: Dwarf butterwort. DM O. TC339. Utricularia subulata L.: Zigzag bladderwort. DM O. TC317, SB200. LILIACEAE Lilium catesbei Walter: Catesby's lily. DM R. TC602. LINDERNIACEAE Micranthemum umbrosum (J.F. Gmel.) S.F. Blake: Shade mudflower. FP O. TC425. LOMARIOPSIDACEAE Nephrolepis cordifolia (L.) C. Presl: Tuberous sword fern. FP O. TC472. Nephrolepis exaltata (L.) Schott: Wild boston fern. FP O. TC564. LYCOPODIACEAE Lycopodiella alopecuroides (L.) Cranfill: Foxtail clubmoss. DM O. TC047. Lycopodiella appressa (Chapm.)Cranfill: Southern clubmoss. DM O. TC607. LYGODIACEAE Lygodium microphyllum (Cav.) R. Br.: Old-world climbing fern. FP I. TC568. LYTHRACEAE Rotala ramosior (L.) Koehne: Toothcup. DM O. TC400. MAGNOLIACEAE Magnolia grandiflora L.: Southern magnolia. In office yard, possibly planted. TC297. Magnolia virginiana L.: Sweetbay magnolia. BH A; FP F. TC173, SB087. MALVACEAE Hibiscus grandiflorus Michx.: Swamp rosemallow. FP O. TC576. Melochia spicata (L.) Fryxell: Bretonica peluda. RU I. TC229. Sida rhombifolia L.: Indian hemp. RU F. TC159, SB261. Urena lobata L.: Caesarweed. RU, Wetland edges F. TC253, SB253. MELASTOMATACEAE Rhexia mariana L.: Pale meadow beauty. MF, WF, DM, FP F. TC033, TC036, SB073. Rhexia nuttallii C.W. James: Nuttall's meadow beauty. DM I. Orzell & Bridges 16667. Rhexia petiolata Walter: Fringed meadow beauty. MF, DM O. TC165. MELIACEAE Melia azedarach L.: Chinaberry tree. RU R. TC454. 85


MOLLUGINACEAE Mollugo verticillata L.: Indian chickweed. RU I. TC585. MORACEAE Morus rubra L.: Red mulberry. FP O. TC466. MYRICACEAE Myrica cerifera L. var. cerifera : Wax myrtle. FP, BH, WF, SF, MF, XH F. TC076, SB146. Myrica cerifera L. var. pumila Michx.: Dwarf wax myrtle. MF, SF O. TC606. (Ward 2000). MYRTACEAE Melaleuca quinquenervia (Cav.) S.T. Blake: Punktree; melaleuca. Pond edge R. TC296. NARTHECIACEAE Aletris lutea Small: Yellow colic-root. DM, WF O. TC089, TC100. NYMPHAEACEAE Nuphar advena (Aiton) Aiton f.: Spatterdock. BW, DM O. TC179, SB120. Nymphaea odorata Aiton subsp. odorata: White water lily. Ponds, BW F. TC284, SB262. OLACACEAE Ximenia americana L.: Tallowwood. XH, SC, SH F. TC243, SB240. OLEACEAE Chionanthus pygmaeus Small: Pygmy fringe tree. SH I. TC080, SB002, SB106. Osmanthus megacarpus (Small) Small ex Little: Scrub wild olive. SC, XH F. TC192, TC457. ONAGRACEAE Ludwigia erecta (L.) H. Hara: Yerba de jicotea. FP O. TC290. Ludwigia lanceolata Elliott: Lanceleaf primrose willow. FP O. TC506, SB206. Ludwigia leptocarpa (Nutt.) H. Hara: Anglestem primrose willow. FP F. TC292, TC366, TC482. Ludwigia maritima Harper: Seaside primrose willow. DM F. TC335, TC572, SB138. Ludwigia octovalvis (Jacq.) P.H. Raven: Mexican primrose willow. FP O. TC440. Ludwigia peruviana (L.) H. Hara: Peruvian primrose willow. FP, Wetland edges F. TC064, TC367. Ludwigia repens J.R. Forst.: Creeping primrose willow. FP F. TC424. Ludwigia suffruticosa Walter: Shrubby primrose willow. DM F. TC526, SB122. Ludwigia virgata Michx.: Savanna primrose willow. DM O. TC140, TC322. Oenothera simulans (Small) W.L. Wagner & Hoch: Sout hern beeblossom. RU O. TC541. (Wagner et al. 2007). (= Gaura angustifolia Michx.). ORCHIDACEAE Encyclia tampensis (Lindl.) Small: Butterfly orchid. FP I. TC494. Habenaria odontopetala Reichenbach f.: False rein orchid. FP O. TC375, SB255. Oeceoclades maculata (Lindl.) Lindl.: Monk orchid. XH R. TC285. Zeuxine strateumatica (L.) Schltr.: Lawn orchid. RU I. TC444. OROBANCHACEAE Agalinis fasciculata (Elliott) Raf.: Beach false foxglove. SH O. TC209. Buchnera americana L.: Blueheart. RU O. TC136, SB297. 86


Seymeria pectinata Pursh: Piedmont blacksenna. SF, SC, SH F. TC266, TC323, SB088. OSMUNDACEAE Osmudastrum cinnamomeum (L.) C. Presl: Cinnamon fe m. FP, BH, WF F. TC015, SB234, SB268. Osmunda regalis L. var. spectabilis (Willd.) A. Gray : Royal fem. BH, FP O. TC063, SB254. OXALIDACEAE Oxalis corniculata L.: Wood sorrel. RU O. TC065, PALMAE Rhapidophyllum hystrix (Pursh) H. Wendl. & Drude ex Drude: Needle palm. FP F (only in one area). TC374. Sabal etonia Swingle ex Nash: Scrub palmetto. SC F. TC188, SB030. Sabal minor (Jacq.) Pers.: Bluestem palm. FP F. TC473. Sabal palmetto (Walter) Lodd. ex Schult. & Schult. f. : Cabbage palm. FP F. TC527. Serenoa repens (W. Bartram) Small: Saw palmetto. MF A; SF, SH, XH F; Pond margins A. TC189, SB031. PASSIFLORACEAE Passiflora incarnata L.: Purple passionflower. RU F. TC536. PHYLLANTHACEAE Phyllanthus tenellus Roxb.: Mascarene Island l eafflower. RU F. TC615. Phyllanthus urinaria L.: Chamber bitter. RU F. TC616. PHYTOLACCACEAE Phytolacca americana L. var. rigida (Small) Caulkins & Wyatt: Pokeweed. RU, FP O. TC220, TC605, SB015. Rivina humilis L.: Rouge plant. XH, RU I. TC043. PINACEAE Pinus clausa (Chapm. ex Engelm.) Vasey ex Sarg.: Sand pine. SC A; RU O. TC030, SB074. Pinus elliottii Engelm.: Slash pine. MF, WF, SF A. TC399, SB074. Pinus palustris Mill.: Longleaf pine. SH F. TC283, SB170. PLANTAGINACEAE Gratiola hispida (Benth. ex Lindl.) Pollard: Rough hedge-hyssop. SH I. TC115, SB060. Gratiola ramosa Walter: Branched hedge-hyssop. DM I. TC417. Gratiola virginiana L.: Roundfruit hedge-hyssop. FP I. TC487. Nuttallanthus canadensis (L.) D.A. Sutton: Blue toadflax. RU O. TC067. (Crawford and Elisens 2006; Sutton 1988). (= Linaria canadensis (L.) Chaz.) Nuttallanthus floridanus (Chapm.) D.A. Sutton: Apalachi cola toadflax. SH, SC F. TC049, TC182, SB007. (Crawford and Elis ens 2006; Sutton 1988). (= Linaria floridana Chapm.) Penstemon multiflorus (Benth.) Chapm. ex Small: Beard-tongue. SH, SF I. TC101. Scoparia dulcis L.: Sweetbroom. RU, FP F. TC519, SB089. POACEAE Amphicarpum muhlenbergianum (Schult.) Hitchc.: Blue maidencane. DM F. TC275, SB182. Andropogon brachystachyus Chapm.: Shortspike bluestem. MF F. TC326. 87


Andropogon floridanus Scribn. : Florida bluestem. SH, SC F. TC241, TC327, TC343, TC356, TC609, SB197. Andropogon glomeratus (Walter) Britton, Sterns & Poggenb. var. glaucopsis (Elliott) C. Mohr: Purple bluestem. DM, MF O. TC344, TC361, SB264, SB272. Andropogon glomeratus (Walter) Britton, Sterns & Poggenb. var. glomeratus : Bushy bluestem. SH, MF O. TC311. Andropogon glomeratus (Walter) Britton, Sterns & Poggenb. var. pumilus (Vasey) Vasey ex L.H. Dewey: Bushy bluestem. DM O. TC016. Andropogon gyrans Ashe var. gyrans : Elliott's bluestem. SH O. TC437. Andropogon gyrans Ashe var. stenophyllus (Hack.)C.S.Campb.: Elliott's bluestem. DM I. TC342. Andropogon ternarius Michx. var. cabanisii (Hack.) Fernald & Griscom: Splitbeard bluestem. SH, SC F. TC146, TC325, SB101. Andropogon ternarius Michx. var. ternarius : Splitbeard bluest em. SH, SC O. SB190. Andropogon tracyi Nash: Tracy's bluestem. SH I. TC026. Andropogon virginicus L. var. decipiens C.S. Campb.: Broomsedge bluestem SH, MF, RU F. TC265, TC329. Andropogon virginicus L. var. glaucus Hack.: Chalky bluestem. SH, SC F. TC330, SB214. Andropogon virginicus L. var. virginicus : Broomsedge bluestem. SH, MF, RU F. SB169, SB176. Aristida condensata Chapm.: Big threeawn. SH O. TC300, TC308. Aristida gyrans Chapm.: Corkscrew threeawn. SC, SF F. SB178, SB199, SB218, SB230, SB231. Aristida lanosa Muhl. ex Elliott: Woolysheath threeawn. SH I. TC355. Aristida purpurascens Poir. var. purpurascens : Arrowfeather threeawn. SH, MF O. TC328. Aristida purpurascens Poir. var. tenuispica (Hitchc.) Allred: Hills boro threeawn. MF O. TC276. Aristida spiciformis Elliott: Bottlebrush threeawn. DM, MF F. TC324, SB133. Aristida stricta Michx.var. beyrichiana (Trin. & Rupr.) D.B. Ward: Wiregrass. SH A; XH, MF O. TC039, TC263, SB160. Arundinaria gigantea (Walter) Muhl.: Switch cane. FP F. TC362, TC539. Axonopus fissifolius (Raddi) Kuhlm.: Common car pet grass. FP O. TC628. Bambusa multiplex (Lour.) Raeusch. ex Schult. & Schult. f.: Hedge bamboo. Pond edge I. TC302. (Plant eradicated since collection was made.) Cenchrus echinatus L.: Southern sandbur. RU O. TC592. Cenchrus spinifex Cav.: Coastal sandbur. RU, SH O. TC298, SB193. Chasmanthium laxum (L.) Yates var. laxum : Slender woodoats. FP O. TC549. Coelorachis tuberculosa (Nash) Nash: Florida jointt ail grass. DM F. TC599. Cynodon dactylon (L.) Pers.: Bermuda grass. RU O. TC627. Dichanthelium aciculare (Desv. ex Poir.) Gould & C.A. Clark subsp. angustifolium (Elliott) Freckmann & Lelong: Need leleaf panicgrass. XH O. SB135. Dichanthelium acuminatum (Sw.) Gould & Clark.: Tapered witchgrass. SH, MF F. TC301, SB152. 88


Dichanthelium chamaelonche (Trin.) Freckmann & Lelong subsp. breve (Hitchc. & Chase) Freckmann & Lelong: Small-seeded panicgrass. SC, SF I. SB079. Dichanthelium commutatum (Schult.) Gould: Variable witchgrass. FP O. TC186, TC260, TC391. Dichanthelium ensifolium (Baldwin ex Elliott) Gould: C ypress witchgrass. DM, MF O. TC496. Dichanthelium erectifolium (Nash) Gould & C.A. Clark: Erec tleaf witchgrass. DM F. TC514. Dichanthelium laxiflorum (Lam.) Gould: Roughhair w itchgrass. SH, MF O. TC238. Dichanthelium portoricense (Desv. ex Ham.) B.F. Hansen & Wunderlin: Hemlock witchgrass. SH, MF F. TC010, TC237, TC398, TC479, SB069, SB153, SB275, SB305. Dichanthelium strigosum (Muhl. ex ell.) Freckmann.: Roughhair witchgrass. SH, MF F. TC408. Echinochloa muricata (P.Beauv.) Fernald var. muricata : Rough barnyard grass. FP O. TC492. Eleusine indica (L.) Gaertn.: Indian goosegrass. RU, DM O. TC307, TC610. Eragrostis amabilis (L.) Wight & Arn. ex Nees: Feather lovegrass. RU O. TC291. Eragrostis refracta (Muhl.) Scribn.: Coastal l ovegrass. MF O. SB163, SB180, SB198. Eragrostis spectabilis (Pursh) Steud.: Purple lovegrass. MF I. TC631. Eremochloa ophiuroides (Munro) Hack.: Centipede grass. RU O. TC629, SB181. Imperata cylindrica (L.) P. Beauv.: Cogon grass. SH O. TC433, TC543. Melinis repens (Willd.) Zizka: Natal grass. RU, SH F. TC017, TC278, SB083, SB114, SB228. Muhlenbergia capillaris (Lam.) Trin. var. capillaris : Hairawn muhly. SH, XH O. TC305. Oplismenus hirtellus (L.) P. Beauv.: Woodsgrass. FP F. TC373. Panicum anceps Michx. subsp. rhizomatum (Hitchc. & Chase) Freckmann & Lelong: Beaked panicgrass. RU O. SB166, SB179. Panicum hemitomon Schult.: Maidencane. DM, FP F. TC334, TC409. Panicum rigidulum Bosc ex Nees subsp. abscissum (Swallen) Freckmann & Lelong: Cutthroat grass. SS F. TC143, TC262, SB113, SB164, SB177. (= P. abscissum Swallen). Panicum rigidulum Bosc ex Nees subsp. combsii (Scribn. & C.R. Ball) Freckmann & Lelong: Redtop panicum. DM O. TC411. Panicum rigidulum Bosc ex Nees subsp. rigidulum : Redtop panicum. FP O. TC438, TC460, TC550. Panicum verrucosum Muhl.: Warty panicg rass. DM, SS O. TC274. Paspalum conjugatum P.J. Bergius: Sour paspalum. RU F. TC532, TC574. Paspalum notatum Flgg: Bahiagrass. RU F. TC234, TC530, SB118. Paspalum urvillei Steud.: Vaseygrass. RU I. TC396. Piptochaetium avenacioides (Nash) Valencia & Costas: Florida needlegrass. SH O. TC380, TC462, SB283. Saccharum giganteum (Walter) Pers.: Sugarcane plumegrass. DM O. TC345, TC611. Sacciolepis striata (L.) Nash: American cupscale. FP O. TC259, TC303, SB165. 89


Schizachyrium niveum (Swallen) Gould: Pinescrub bluestem. SC I. (TC) Christman 2086. Schizachyrium scoparium (Michx.) Nash var. scoparium : Little bluestem. SH, SF, MF F. TC236, TC273, TC358, SB168, SB229. Setaria corrugata (Elliott) Schult.: Coastal br istlegrass. SH, RU O. TC257, TC277. Setaria magna Griseb.: Giant bristlegrass. FP I. TC577. Setaria parviflora (Poir.) Kergulen: Knotroot foxtail. RU, XH O. TC239. Sorghastrum secundum (Elliott) Nash: Lopsided indiangrass. SH F. TC304, TC357, SB189. Spartina bakeri Merr.: Sand cordgrass. DM O. TC634, SB263. Sporobolus floridanus Chapm.: Florida dropseed. MF O. SB260. Sporobolus jacquemontii Kunth: Smutgrass. RU, MF O. TC211, TC235 SB219. Sporobolus junceus (P. Beauv.) Kunth: Pineywoods dropseed. SH O. TC293, SB244. Stenotaphrum secundatum (Walter) Kuntze: St. Augus tine grass. RU O. TC630. Tridens flavus (L.) Hitchc. var. chapmanii (Small) Shinners: Chapman's purpletop tridens. SH I. TC354. Tripsacum dactyloides (L.) L.: Eastern gama grass. FP I. TC548. Urochloa maxima (Jacq.) R.D. Webster: Guinea grass. RU, FP O. TC247, TC264, TC545, TC591. POLYGALACEAE Polygala cymosa Walter: Tall pinebarren milkwort. DM F. TC105. Polygala incarnata L.: Procession flower. SH I. TC199. Polygala lewtonii Small: Lewton's polygala. SH O. TC052. Polygala nana (Michx.) DC: Wild batchelor's button. DM I. TC097. Polygala polygama Walter: Racemed milkwort. SH O. TC463. Polygala rugelii Shuttlew. ex Chapm.: Yellow milkwort. DM, WF F. TC096, SB072. Polygala setacea Michx.: Coastalplain milkwort. DM O. TC495, SB070. POLYGONACEAE Eriogonum tomentosum Michx.: Wild buckwheat. SH, XH O. TC034. Eriogonum longifolium Nutt. var. gnaphalifolium Gand.: Scrub buckwheat. SH F. TC104. Persicaria hirsuta (Walter) Small: Hairy smartweed. DM F. SB016. (Lamb Frye and Kron 2003). Persicaria hydropiperoides (Michx.) Small: Mild water pepper. FP O. TC418, TC489 SB245. (Lamb Frye and Kron 2003). Persicaria punctata (Elliott) Small: Dotted smartweed. FP F. TC271. (Lamb Frye and Kron 2003). Polygonella basiramia (Small) G.L. Nesom & V.M. Bates: Florida jointweed. SC, SF O. SB125, SB157. Polygonella gracilis Meisn.: Tall jointweed. SH, XH I. TC014. Polygonella myriophylla (Small) Horton: Sandlace. SC F. SB023. Polygonella polygama (Vent.) Engelm. & A. Gray var. polygama : October flower. SH, SC, SF F. TC321, SB188. Polygonella robusta (Small) G.L. Nesom & V.M. Bates: Sandhill wireweed. SH, SF F TC002, SB112. Rumex hastatulus Baldwin: Heartwing dock. FP I. TC412. 90


POLYPODIACEAE Phlebodium aureum (L.) J. Sm.: Golden polypody. FP, Creek banks (epiphytic on Quercus virginiana and Sabal palmetto ) I. TC180. Pleopeltis polypodioides (L.) E.G. Andrews & Windham var. michauxiana (Weath.) E.G. Andrews & Windham: Resurrection fern. FP, XH F. TC187, SB134. PONTEDERIACEAE Eichhornia crassipes (Mart.) Solms: Water hyacinth. BW O. TC250. Pontederia cordata L.: Pickerelweed. FP, DM O. TC087, SB299. PORTULACACEAE Portulaca amilis Speg.: Paraguayan purslane. RU I. TC614. Portulaca pilosa L.: Pink purslane. RU I. TC553. RANUNCULACEAE Clematis reticulata Walter: Leatherflower. SH, SF O. TC102, TC542. ROSACEAE Aronia arbutifolia (L.) Pers.: Red chokeberry. MF O. TC058, SB098. (Kalkman 2004). (= Photinia pyrifolia (Lam.) K.R. Robertson & J.B. Phipps). Crataegus lepida Beadle: Scrub hawthorn. SH I. TC154. [Wunderlin and Hansen (2003) include this taxon within C. michauxii Pers., but Phipps ( e.g., Phipps and Dvorsky 2007) follows Beadle in considering south eastern North America to be an area of extreme species richness for Crataegus. He recognizes several of Beadles species, and has annotated specimens at FLAS as C. lepida. Although one could criticize the naming of hundreds of Crataegus morphotypes, I believe in this case that the distinctive morphology and narrow habita t preference of this taxon justifies recognizing it as a distinct species (Beadle 1901), pending further study. As such, it qualifies as another Fl orida scrub endemic]. Prunus caroliniana (Mill.) Aiton: Carolina laurel cherry. RU I. TC363. Prunus geniculata R.M. Harper: Scrub plum. SH, XH O. TC504, SB273. Prunus serotina Ehrh.: Wild cherry. FP, RU O. TC559. Prunus umbellata Elliott: Flatwoods plum. RU O. TC221, TC481. Rubus cuneifolius Pursh: Sand blackberry. SH, SF I. TC590. Rubus argutus Link: Sawtooth blackberry. FP, WF, MF F. TC069, SB026, SB238, SB271. RUBIACEAE Cephalanthus occidentalis L.: Buttonbush. DM, FP F. TC124, SB307. Diodia teres Walter: Poor Joe. SH, MF O. TC231, SB131. Diodia virginiana L.: Buttonweed. WF I. TC571. Galium hispidulum Michx.: Coastal bedstraw. SH I. TC583. Galium tinctorium L.: Stiff bedstraw. FP I. TC423. Houstonia procumbens (J.F .Gmel.) Standl.: Innocence. SH, XH I. TC073. Mitchella repens L.: Partridge berry. FP, XH I. TC421. Oldenlandia uniflora L.: Clustered mille graines. FP, RU, DM F. TC232, SB137. Psychotria nervosa Sw.: Wild coffee. FP F. TC372, TC560. Psychotria sulzneri Small: Shortleaf wild coffee. FP F. TC561. Richardia brasiliensis Gomes: Mexican clover. RU O. TC215. Spermacoce remota Lam.: Woodland false buttonweed. RU O. TC225. 91


RUSCACEAE Nolina brittoniana Nash: Britton's bear-grass. SH, SF O. TC518, SB155. SALICACEAE Salix caroliniana Michx.: Carolina willow. FP, Wetland edges F. TC388, SB257. SALVINIACEAE Azolla caroliniana Willd.: Carolina mosquito fern. BW, FP F. TC551, TC570. Salvinia minima Baker: Water spangles. BW, FP F. TC434, SB278. SAPINDACEAE Acer rubrum L.: Red maple. FP F. TC382, SB237. SAPOTACEAE Sideroxylon tenax L.: Tough bumelia. SC, SH F. TC107, TC216, TC217, SB045. [At least two of these specimens ( i.e., TC107, SB045 ) may represent the morphologically distinct lacuum entity, once recognized as Bumelia lacuum Small (see Lakela 1963) but now included within S. tenax. More study is needed]. SAURURACEAE Saururus cernuus L.: Lizard's tail. FP F. TC086. SELAGINELLACEAE Selaginella arenicola Underw.: Sand spikemoss. SC, SH F. TC314, SB020. SMILACACEAE Smilax auriculata Walter: Earleaf greenbrier. SH, XH, SC, SF, MF, RU F. TC193, SB010, SB039 Smilax bona-nox L.: Saw greenbrier. FP F. TC455. Smilax glauca Walter: Greenbrier. MF, SH O. TC095. Smilax laurifolia L.: Laurel greenbrier. FP, BH O. TC081, SB085. Smilax pumila Walter: Sarsaparilla vine. FP, XH O. TC452. Smilax walteri Pursh: Coral greenbrier. FP I. TC394. SOLANACEAE Physalis arenicola Kearney: Sand ground-cherry. SH O. TC537, TC582. Physalis walteri Nutt.: Walter's ground-cherry. SH, XH I. TC268, TC511. Solanum americanum Mill.: Common nightshade. RU O. TC364. Solanum viarum Dunal: Tropical soda apple. FP, XH, RU I. TC214, SB252. TETRACHONDRACEAE Polypremum procumbens L.: Rustweed. DM, RU F. TC152, SB052. THEACEAE Gordonia lasianthus (L.) J. Ellis: Loblolly bay. BH A. TC038, SB061. THELYPTERIDACEAE Thelypteris dentata (Forssk.) E.P. St. John: Downy shield fern. FP O. TC563. Thelypteris interrupta (Willd.) K. Iwats.: Hottentot fern. FP F. TC448, TC453. Thelypteris kunthii (Desv.) C.V. Morton: Southe rn shield fern. FP O. TC562. TURNERACEAE Piriqueta cistoides (L.) Griseb. subsp. caroliniana (Walter) Arbo (Hybrid morphotype): Piriqueta. SH I. TC149, TC584. [J.K. Small (1933) recognized four species of Piriqueta, including P. caroliniana of northern Florida sandy uplands, and P. viridis of calcareous soils of southern Florida. Arbo (1995) grouped all four of Smalls species into P. cistoides subsp. caroliniana. Maskas and Cruzan (2000) and Martin and Cruzan (1999) recognize these taxa as distinct morphotypes. The P. viridis 92


93 and P. caroliniana morphotypes interbreed, with a broad hybrid zone in central Florida. These specimens, with erect habi t, oblong leaves, and stellate hairs only, are intermediate between the decumb ent, hirsute, elliptic-leaved P. caroliniana morphotype and the erect, glabrous, linear-leaved P. viridis morphotype]. TYPHACEAE Typha domingensis Pers.: Southern cattail. FP O. TC428. Typha latifolia L.: Common cattail. FP O. TC251. ULMACEAE Ulmus americana L.: American elm. FP O. TC558. URTICACEAE Boehmeria cylindrica (L.) Sw.: Bog hemp. FP F. TC429. VERBENACEAE Lantana camara L.: Lantana. RU O. TC161, SB285. Phyla nodiflora (L.) Greene: Fogfruit. RU F. TC170. Verbena brasiliensis Vell.: Brazilian vervain. RU O. TC384. VIOLACEAE Viola lanceolata L.: Longleaf violet. DM F. TC446, SB269. Viola primulifolia L.: Primroseleaf violet. BH, MF I. TC048, SB270. VISCACEAE Phoradendron leucarpum (Raf.) Reveal & M.C. Johnst.: Oak mistletoe. SH, XH, FP O. TC084. VITACEAE Ampelopsis arborea (L.) Koehne: Peppervine. FP, RU O. TC486. Parthenocissus quinquefolia (L.) Planch.: Virginia cree per. XH, FP, RU F. TC295, SB243. Vitis aestivalis Michx.: Summer grape. RU O. TC484. Vitis cinerea (Engelm.) Engelm. ex Millardet var. floridana Munson: Florida grape. RU, XH O. TC181, TC544. Vitis rotundifolia Michx.: Muscadine grape. SH, MF SF, SC, FP, BH, RU F. TC177, SB037, SB081, SB295. XYRIDACEAE Xyris ambigua Beyr. ex Kunth: Coastalplain yellow-eyed grass. DM O. TC138, TC282. Xyris brevifolia Michx.: Shortleaf yellow-eyed grass. DM O. TC379, TC404. Xyris caroliniana Walter: Carolina yellow-eyed grass. DM O. SB099. Xyris elliottii Chapm.: Elliott's yellow-eyed grass. DM, WF F. TC498, SB084. Xyris jupicai Rich.: Richard's yellow-eyed grass. DM O. TC055, TC608.


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BIOGRAPHICAL SKETCH Paul Corogin was born in Columbus, Ohio. He grew up near the shores of Lake Erie. After earning a Bachelor of Science in zoology from The Ohio State University in 1983, he moved to Gainesville, Florida, where he became a massage therapist and worked in landscaping. Years of working outdoors deepened his interest in plants, and during a stint of volunteer field work in Ocala National Forest, he crystallized a longstanding desire to return to school and study botany. He earned a second bachelo rs degree, in botany, at the Univ ersity of Florida, where he then began his graduate studies, earning a Ma ster of Science in botany in May 2008. 107