Guide to the Natural Communities of Florida - 2010 Edition

Cover Photograph: Mesic flatwoods at Triple N Ranch Wildlife Management Area, Osceola County (Gary Knight) Recommended citation: Florida Natural Areas Inventory (FNAI). 2010. Guide to the natural communities of Florida: 2010 edition. Florida Natural Areas Inventory, Tallahassee, FL. PREFACE In 2007, with funding from the Florida Department of Environmental Protection (FDEP), Division of State Lands, the Florida Natural Areas Inventory (FNAI) began a process of updating the "Guide to the Natural Communities of Florida" (the Guide), which had been only slightly modified since it was first published in 1990 by FNAI and the Florida Department of Natural Resources (now FDEP). The current update includes only the forty-five land-based communities (23 terrestrial and 20 palustrine communities, plus tidal marsh and tidal swamp in the marine and estuarine category), leaving the remaining communities to be updated at a later time, except for the updating of species names. The purpose of the update is to clarify distinctions between communities by listing characteristic species and features distinguishing similar communities, as well as to add information for each community on variations throughout its range (with common variants noted specifically), range, natural processes, management, exemplary sites, and references. The resulting 2010 Guide contains the original marine, estuarine, lacustrine, riverine, and subterranean communities, plus the updated 46 land-based communities, with 9 new community names added - alluvial forest, glades marsh, Keys cactus barren, Keys tidal rock barren, limestone outcrop, shrub bog, slough marsh, upland mixed woodland, and upland pine, and 8 original community names deleted (their names being changed or their concepts being subsumed under other communities) - bog, coastal rock barren, floodplain forest, freshwater tidal swamp, prairie hammock, swale, upland mixed forest, and upland pine forest. A complete crosswalk between the 1990 Guide and the 2010 Guide and vice versa is provided here with further explanation of changes made (Appendix 1). The updated, land-based natural communities are grouped according to a new organization, emphasizing similarities in species composition, structure, and landscape position. Also included in the 2010 Guide are a list of altered landcover types and their definitions (Appendix 2). These are habitats that have been severely impacted by humans and do not fit into FNAI’s Natural Community Classification. Altered landcover types can be used to describe the most common non-natural habitats observed on conservation lands in Florida. ACKNOWLEDGEMENTS 1 INTRODUCTION 2 NATURAL COMMUNITY SHORT DESCRIPTIONS 4 HARDWOOD FORESTED UPLANDS Slope Forest 12 Upland Hardwood Forest 16 Mesic Hammock 19 Rockland Hammock 24 Xeric Hammock 29 HIGH PINE AND SCRUB Upland Mixed Woodland 32 Upland Pine 35 Sandhill 39 Scrub 44 PINE FLATWOODS AND DRY PRAIRIE Wet Flatwoods 49 Mesic Flatwoods 54 Scrubby Flatwoods 58 Pine Rockland 61 Dry Prairie 67 COASTAL UPLANDS Beach Dune 70 Coastal Berm 73 Coastal Grassland 75 Coastal Strand 78 Maritime Hammock 82 Shell Mound 85 SINKHOLES AND OUTCROP COMMUNITIES Upland Glade 88 Sinkhole 91 Limestone Outcrop 94 Keys Cactus Barren 97 FRESHWATER NON-FORESTED WETLANDS PRAIRIES AND BOGS Seepage Slope 99 Wet Prairie 102 Marl Prairie 107 Shrub Bog 110 MARSHES Depression Marsh 113 Basin Marsh 117 Coastal Interdunal Swale 120 Floodplain Marsh 122 Slough Marsh 126 Glades Marsh 129 Slough 134 FRESHWATER FORESTED WETLANDS CYPRESS/TUPELO Dome Swamp 137 Basin Swamp 142 Strand Swamp 146 Floodplain Swamp 149 HARDWOOD Baygall 154 Hydric Hammock 158 Bottomland Forest 162 Alluvial Forest 166 MARINE AND ESTUARINE VEGETATED WETLANDS Salt Marsh 170 Mangrove Swamp 175 Keys Tidal Rock Barren 179 LACUSTRINE Clastic Upland Lake 181 Coastal Dune Lake 182 Coastal Rockland Lake 183 Flatwoods/Prairie Lake and Marsh Lake 183 River Floodplain Lake and Swamp Lake 184 Sandhill Upland Lake 185 Sinkhole Lake 186 RIVERINE Alluvial Stream 187 Blackwater Stream 188 Seepage Stream 188 Spring-run Stream 189 MARINE AND ESTUARINE MINERAL BASED Consolidated Substrate 191 Unconsolidated Substrate 192 FAUNAL BASED Coral Reef 193 Mollusk Reef 194 Octocoral Bed 195 Sponge Bed 195 Worm Reef 196 FLORAL BASED Algal Bed 196 Seagrass Bed 197 COMPOSITE SUBSTRATE Composite Substrate 198 SUBTERRANEAN Aquatic and Terrestrial Cave 199 REFERENCES 201 Appendix 1. Crosswalk Appendix 2. Altered Landcover Types We owe a large debt of gratitude to Dr. O. Greg Brock, bureau chief of the Office of Environmental Services at FDEP, whose support and encouragement made this project possible. We would like to thank the following experts for their helpful comments on the individual accounts: Richard Abbott, Guy Anglin, Wilson Baker, Chris Bergh, Keith Bradley, Susan Carr, Linda Chafin, Pete Colverson, Anne Cox, Melanie Darst, Linda Duever, Mark Deyrup, Janice Dusquesnel, Kathy Ewel, Elise Gornish, Dennis Hardin, Richard Hilsenbeck, Jean Huffman, Michael Jenkins, Erik Johnson, Adam Kent, Kay Kirkman, Helen Light, Matt Love, Eric Menges, Tom Miller, Gil Nelson, Eliane Norman, Dan Pearson, Kevin Robertson, Anne Rudloe, Paul Schmalzer, Bob Simons, James Snyder, Bruce Sorrie, Walter Taylor, Walt Thompson, John Tobe, Greg Walker (and the biologists of the Florida Park Service), and Samuel Wright. INTRODUCTION This document presents the hierarchical classification of 81 Natural Communities, developed by the Florida Natural Areas Inventory (FNAI) and identified as collectively constituting the original, natural biological associations of Florida. A Natural Community (NC) is defined as a distinct and recurring assemblage of populations of plants, animals, fungi and microorganisms naturally associated with each other and their physical environment. The 2010 FNAI natural community classification groups natural communities into categories based on a combination of vegetation, landscape position, and hydrology. The categories, such as “hardwood forested uplands” are meant to be an intuitive grouping of similar natural communities that allows the user to apply a broader level name to an area when the community determination is uncertain. In some cases, an intermediate group (ex. “Cypress/Tupelo”) is identified for further clarification. At the finest level, variants (if any) are described within the community accounts. These variants are optional descriptions that can be used to convey more specific information about an individual community occurrence. For example, a swamp found at the mouth of a river might be designated “floodplain swamp - freshwater tidal swamp variant.” Variants have the same ranking as the natural community of which they are a part. The treatment of each of the 46 communities updated in 2010 follows the outline below. • Description - gives an overall picture of the community with a description of the general vegetation structure and composition, landscape position, soils, hydrology, salinity, and/or fire. • Characteristic Set of Species - short list of typical species. • Rare Species - a discussion of rare species for which the community is critical habitat. This is not meant to be an exhaustive list. • Range - discussion of Florida range and notes on the distribution of similar communities elsewhere in the US or worldwide. • Natural Processes - a discussion of important natural processes including fire, hydrology, etc. • Community Variations - discussion of commonly occurring variation, both regional and within community. ■ Variant - names and short descriptions of notable variants of the natural community. These are informal names that are meant to convey further information about specific community occurrences. Their ranks are the same as that of the entire community. • Associated Communities - addresses similar and/or adjacent communities and gives guidelines for recognition. • Management Considerations - addresses fire recommendations and any other management oriented issues stemming from disturbance of natural processes. • Exemplary Sites - these may be specific place names or managed areas (full name) with county listed in parentheses. • Global and State Ranks - current ranking of community by FNAI. • Crosswalk and Synonyms - crosswalk to other classifications and a list of synonyms commonly found in the literature • References - cited in text The classification of the remaining (primarily aquatic) communities that were not updated remains unchanged. These communities are grouped into Lacustrine, Riverine, Subterranean, and Estuarine and Marine categories A second level of the hierarchy splits the Estuarine and Marine categories into Natural Community Groups, including Faunal, Floral, and Substrate-based. Natural Communities are characterized and defined by a combination of physiognomy, vegetation structure and composition, topography, land form, substrate, soil moisture condition, climate, and fire. They are named for their most characteristic biological or physical feature. Natural Communities could be further subdivided into Plant Communities based on the dominant plant species. This level of subdivision is not presented in this document. FNAI uses several criteria, including area covered and number of occurrences, to determine the relative rarity and threat to each community type; these are summarized into a global and a state rank, the G and S ranks, respectively. Most G ranks for NCs are tentative, pending more information from comparison and coordination with other states and countries. Three NCs are given the highest rarity ranking (G1/S1): upland glade, Keys cactus barren, and pine rockland. The first two, being confined to limestone outcrops in wooded landscapes, are naturally of limited extent; the third occurs in the Miami-Dade urban area and has been largely altered for human use. Twelve NCs are ranked at the second most endangered level (G2/S2). Six of these are of naturally limited extent: slope forest, seepage slope, sinkhole, limestone outcrop, shell mound, and strand swamp, and six have been largely altered for human use: dry prairie, wet prairie, scrub, scrubby flatwoods, upland mixed woodland, and rockland hammock. With the exception of wet prairie and seepage slope which are found in adjacent states, all of these rarest NCs are largely confined to Florida. Several other classification systems are in use in Florida, with four principal ones that cover the entire state. The widely used FLUCCS (Florida Land Use, Cover, and Forms Classification System) system, is a general purpose mapping system, devised by the Florida Department of Transportation, that includes both natural and altered (i.e., residential areas, farmland, pastures, etc.) categories that are distinguishable on aerial photographs. This system is used by the five Florida Water Management Districts to produce periodically updated, digitized maps of the entire state interpreted from current aerial photography. The Florida Land Cover map produced by the Florida Fish and Wildlife Conservation Commission (FWC) uses mechanical interpretation of satellite imagery (2003) to produce a single statewide map using 23 natural categories and 16 human-modified categories. More recently the FWC developed the Florida Land Cover Classification System (2009), a hierarchical classification consisting of more than 200 classes that integrates the FNAI system with FLUCCS. This classification is the basis for Cooperative Land Cover Map (FNAI 2010), a statewide digital map that incorporates FLUCCS, high quality site-based land cover, and improvements to focal community maps through aerial photo review. A finer-scale plant community level of classification called the U. S. National Vegetation Classification (USNVC), developed by The Nature Conservancy (and now maintained by NatureServe) for conservation purposes and continuously updated, uses dominant plant species to define its finest units, called “associations”, of which it has recognized over 350 in Florida.287 NatureServe has also grouped the associations into ecological systems, based on geographic location and landform, of which there are 73 in Florida. The FNAI classification seeks to encompass all natural assemblages at a scale that is useful for mapping natural areas and communication about land management. It differs from the FLUCCS and the FWC landcover systems in its lack of non-natural categories and its greater detail in natural categories. It differs from the associations of the USNVC Classification in not being based on dominant species and from the ecological systems of NatureServe in not being based on geographic area of the state in which the assemblage occurs. The FNAI classification is meant to be relatively stable in order to facilitate its usage in state conservation efforts. However, we believe that the guide should also remain flexible and dynamic to incorporate up-to-date information. Future editions of the Guide will be released as new data and research accumulate. NATURAL COMMUNITY SHORT DESCRIPTIONS HARDWOOD FORESTED UPLANDS - mesic or xeric forest dominated mainly by hardwood trees. Slope Forest (G2?/S1) - steep slope on bluff or in sheltered ravine within the Apalachicola drainage; sand/clay substrate; mesic-hydric; central Panhandle; rare or no fire; closed canopy of mainly deciduous species; American beech, Florida maple, white oak, Ashe's magnolia, southern magnolia, spruce pine, Shumard's oak. Upland Hardwood Forest (G5/S3) - upland with sand/clay and/or calcareous substrate; mesic; Panhandle to central peninsula; rare or no fire; closed deciduous or mixed deciduous/evergreen canopy; American beech, southern magnolia, hackberry, swamp chestnut oak, white oak, horse sugar, flowering dogwood, and mixed hardwoods. Dry Upland Hardwood Forest - on dry slopes or along upper slopes with sand/clay substrate; mesic; temperate; rare fire; closed canopy; laurel oak and/or live oak and/or pignut hickory, southern magnolia, shortleaf pine, loblolly pine, and/or mixed hardwoods. Mesic Hammock (G3/S3?) - flatland with sand/organic soil; mesic; primarily central peninsula; occasional or rare fire; closed evergreen canopy; live oak, cabbage palm, southern magnolia, pignut hickory, saw palmetto. Prairie Mesic Hammock - Isolated stands within a matrix of pyrogenic vegetation; occasional fire; live oak, cabbage palm, saw palmetto. Rockland Hammock (G2/S2) - flatland with limestone substrate; mesic; southern peninsula and Keys; rare or no fire; closed canopy of evergreen mixed tropical hardwoods; gumbo limbo, pigeon plum, stoppers. Thorn Scrub - along ecotones or within openings in rockland hammock; low-statured; dominated by spiny species; saffron plum, blackbead, hog plum, buttonwood, plus other common rockland hammock species. Xeric Hammock (G3/S3) - upland with deep sand substrate; xeric; primarily eastern Panhandle to central peninsula; rare or no fire; closed canopy of evergreen hardwoods; sand live oak, saw palmetto. HIGH PINE AND SCRUB - hills with mesic or xeric woodlands or shrublands; canopy, if present, open and consisting of pine or a mixture of pine and deciduous hardwoods. Upland Mixed Woodland (G2/S2) - upland with loamy soils; mesic-xeric; central Panhandle to extreme northern central peninsula; occasional fire (variable but as little as two up to 20 year interval); open to partially closed canopy over an open understory of mixed herbs and scattered shrubs; mixture of southern red oak, mockernut hickory, and longleaf or shortleaf pine with other mixed hardwoods; wiregrass infrequent. Upland Pine (G3/S2) - upland with sand/clay substrate; mesic-xeric; Panhandle to extreme northern central peninsula; frequent fire (1 -3 years); savanna of widely spaced pines over primarily herbaceous understory; longleaf pine and/or loblolly pine and/or shortleaf pine, southern red oak, wiregrass. Sandhill (G3/S2) - upland with deep sand substrate; xeric; Panhandle to central peninsula; frequent fire (1-3 years); savanna of widely spaced longleaf pine and/or turkey oak with wiregrass understory. Scrub (G2/S2) - upland with deep sand substrate; xeric; statewide except extreme southern peninsula and Keys, mainly coastal in Panhandle; occasional or rare fire (usually 5-20 years); open or dense shrubs with or without pine canopy; sand pine and/or scrub oaks and/or Florida rosemary. Rosemary Scrub - on the driest ridge crests, particularly at the southern end of the Lake Wales Ridge and on Panhandle barrier islands; occasional or rare fire (10-40 years); dominated by Florida rosemary with large areas of bare sand visible between the shrubs. Sand Pine Scrub - on ridges throughout the state; rare fire (20-80 years); canopy of sand pine and an understory of the three shrubby oaks, or less commonly, Florida rosemary. PINE FLATWOODS AND DRY PRAIRIE - mesic or hydric pine woodland or mesic shrubland on flat sandy or limestone subtrates, may have a hard pan that impedes drainage. Wet Flatwoods (G4/S4) - flatland with sand substrate; seasonally inundated; statewide except extreme southern peninsula and Keys; frequent fire (2-4 years for grassy wet flatwoods, 5-10 years for shrubby wet flatwoods); closed to open pine canopy with grassy or shrubby understory; slash pine, pond pine, large gallberry, fetterbush, sweetbay, cabbage palm, wiregrass, toothache grass. Cutthroat Grass Flatwoods - on and near the Lake Wales Ridge; frequent fire (2-4 years); widely scattered pines over cutthroat grass and/or other hydrophytic herbs. Cabbage Palm Flatwoods - on shelly sand or where limestone is near the surface; central to southern peninsula; pine canopy over cabbage palm understory. Mesic Flatwoods (G4/S4) - flatland with sand substrate; mesic; statewide except extreme southern peninsula and Keys; frequent fire (2-4 years); open pine canopy with a layer of low shrubs and herbs; longleaf pine and/or slash pine, saw palmetto, gallberry, dwarf live oak, wiregrass. Scrubby Flatwoods (G2/S2?) - flatland with sand substrate; xeric-mesic; statewide except extreme southern peninsula and Keys; occasional fire (5-15 years); widely scattered pine canopy over saw palmetto and scrub oaks; longleaf pine, sand live oak, myrtle oak, Chapman's oak, saw palmetto, wiregrass. Pine Rockland (G1/s1) - flatland with exposed limestone substrate; mesic-xeric; southern peninsula and Keys; frequent to occasional fire (3-7 years); open pine canopy with mixed shrubs and herbs in understory; south Florida slash pine, palms, mixed tropical and temperate shrubs, grasses, and herbs. Dry Prairie (G2/S2) - flatland with sand soils over an organic or clay hardpan; mesic-xeric; central peninsula; annual or frequent fire (1 -2 years); treeless with a low cover of shrubs and herbs; wiregrass, dwarf live oak, stunted saw palmetto, bottlebrush threeawn, broomsedge bluestem. COASTAL UPLANDS - mesic or xeric communities restricted to barrier islands and near shore; woody or herbaceous vegetation; other communities may also occur in coastal environments. Beach Dune (G3/s2) - active coastal dune with sand substrate; xeric; statewide; rare or no fire; marine influence; open herbaceous vegetation with no canopy; sea oats, railroad vine, bitter panicum, and/or mixed salt-spray tolerant grasses and herbs. Coastal Berm (G3/S2) - old bar or storm debris with sand/shell substrate; xeric-mesic; southern peninsula and Keys; rare or no fire; marine influence; variable vegetation structure; mixed tropical herbs, shrubs, and trees. Coastal Grassland (G3/S2) - coastal flatland behind dunes with stable sand substrate; mesic-hydric; statewide excluding Keys; occasional fire; marine influence; herbaceous vegetation with no canopy; salt-tolerant grasses and herbs; sea oats, bitter panicum, camphorweed, hairawn muhly, Gulf bluestem. Coastal Strand (G3/S2) - stabilized coastal dune with sand substrate; xeric; peninsula; rare fire; marine influence; primarily dense shrubs; saw palmetto in temperate coastal strand or seagrape and/or saw palmetto in tropical coastal strand. Maritime Hammock (G3/s2) - stabilized coastal dune with sand substrate; xeric-mesic; statewide but rare in Panhandle and Keys; rare or no fire; marine influence; evergreen closed canopy; live oak, cabbage palm, red bay, red cedar in temperate maritime hammock; gumbo limbo, seagrape, and white or spanish stopper in tropical maritime hammock. Shell Mound (G2/S2) - small hill of shells deposited by native Americans; mesic-xeric; statewide; rare or no fire; marine influence; closed canopy of mixed hardwoods; soapberry, snowberry, white stopper. SINKHOLES AND OUTCROP COMMUNITIES - small extent communities in karst features or on exposed limestone. Upland Glade (G1/S1) - upland with thin clay soils over limestone outcrops; hydric-xeric; central Panhandle only; sparse mixed grasses and herbs with occasional stunted trees and shrubs that are concentrated around the edge; black bogrush, poverty dropseed, diamondflowers, hairawn muhly, Boykin's polygala, red cedar. Sinkhole (G2/S2) - karst feature with steep walls; mesic-hydric; statewide; variable vegetation structure. Limestone Outcrop (G2/S2) - exposed limestone; mesic-hydric; statewide; often with mosses, liverworts, and a diversity of rare ferns. Keys Cactus Barren (G1/S1) - small openings on flatland with exposed limestone; xeric; restricted to Keys; marine influence; open, herbaceous vegetation with some cacti, agave, and stunted trees; three-spined pricklypear, erect pricklypear, barbed wire cactus, Yucatan fly mallow, Florida Keys indigo, skyblue clustervine, dwarf bindweed. FRESHWATER NON-FORESTED WETLANDS - herbaceous or shrubby palustrine communities in floodplains or depressions; canopy trees, if present, very sparse and often stunted. PRAIRIES AND BOGS - short hydroperiod; dominated by grasses, sedges, and/or titi. Seepage Slope (G2/S2) - on or at base of slope with loamy sand substrate; maintained by downslope seepage, usually saturated but rarely inundated; Panhandle and northern peninsula; frequent fire (1-3 years); dense herbaceous community; wiregrass, wiry beaksedges, flattened pipewort, toothache grass, pitcherplants. Wet Prairie (G2/S2) - flatland with sand or clayey sand substrate; usually saturated but only occasionally inundated; statewide excluding extreme southern peninsula; frequent fire (2-3 years); treeless, dense herbaceous community with few shrubs; wiregrass, blue maidencane, cutthroat grass, wiry beaksedges, flattened pipewort, toothache grass, pitcherplants, coastalplain yellow-eyed grass. Cutthroat Seep - eastern and western edges of the Lake Wales Ridge in central Florida; dominated by the endemic cutthroat grass. Calcareous Wet Prairie - in central and south-central peninsula on calcareous soils; Gulf hairawn muhly typically dominant with other calcium-loving species. Pitcherplant Prairie - in the Panhandle on wetter soils; dense stands of tall pitcherplants. Marl Prairie (G3/S3) - flatland with marl over limestone substrate; seasonally inundated (<4 months); southern peninsula; frequent to occasional fire (2-10 years depending on density of herbs); purple muhly, sawgrass (stunted), spreading beaksedge, black bogrush, Florida little bluestem, and/or mixed grasses, sometimes with dwarf cypress. Shrub Bog (G4/S3) - wetland on organic soil over sand; soil often saturated and mucky, occasionally shallowly inundated; Panhandle to north peninsula; occasional fire (10-20 years); dense stand of shrubs, trees absent or sparse, sphagnum moss common; titi, black titi, fetterbush, large gallberry, laurel greenbrier, pond pine or slash pine. MARSHES - long hydroperiod; dominated by grasses, sedges, broadleaf emergents, floating aquatics, or shrubs. Depression Marsh (G4/S4) - small, isolated, often rounded depression in sand substrate with peat accumulating toward center; surrounded by fire-maintained community; seasonally inundated; still water; statewide excluding Keys; frequent or occasional fire; largely herbaceous; maidencane, sawgrass, pickerelweed, longleaf threeawn, sand cordgrass, peelbark St. John's wort. Basin Marsh (G4/S3) - basin with peat or sand substrate; seasonally inundated; statewide excluding Keys; occasional fire; largely herbaceous; maidencane, sawgrass, bulltongue arrowhead, pickerelweed, Baker's cordgrass, white water lily, coastalplain willow. Lake Bottom - marshes on former lake bottoms of “disappearing” lakes in northern Florida, areas that alternate between lake and marsh when the sinkholes draining them are plugged or reopened; well-known examples are Lake Miccosukee and Paynes Prairie. Coastal Interdunal Swale (G3/S2) - linear wetlands between dunes on sandy barrier islands; inundated by local rainfall events; Panhandle to central peninsula; herbaceous or shrubby; sawgrass, hairawn muhly, broomsedge, seashore paspalum, Baker's cordgrass, saltmeadow cordgrass, wax myrtle, coastalplain willow. Floodplain Marsh (G3/S3) - floodplain with organic/sand/alluvial substrate; seasonally inundated; Panhandle to central peninsula; frequent or occasional fire (ca. 3 years, much less frequent in freshwater tidal marshes); treeless herbaceous community with few shrubs; sawgrass, maidencane, sand cordgrass, and/or mixed emergents. Freshwater Tidal Marsh - river mouth wetland on organic/alluvial substrates; receives pulses of freshwater in response to tides; sawgrass, giant cutgrass. Slough Marsh (G3?/S3?)- broad, shallow channel with sand/peat substrate; seasonally inundated; intermittently flowing water; central to southern peninsula; frequent or occasional fire (3-10 years); sawgrass, maidencane, pickerelweed, and/or mixed emergents. Glades Marsh (G3/S3) - broad, shallow channel with peat/marl substrate directly overlying limestone; seasonally inundated; stagnant or slow flowing water; Everglades basin, Big Cypress region, and Keys; frequent to occasional fire (3-10 years); sawgrass, spikerush, maidencane, beaksedges, mixed emergents. Keys Freshwater Marsh - limestone depression; restricted to Florida Keys; may be saline during dry season; sawgrass. Slough (G3/S3) - broad, shallow channel with peat; inundated except during droughts; flowing water; statewide excluding Keys; rare fire; sparsely canopied or with emergent or floating plants; alligator flag, American white waterlily. Pond Apple Slough - canopied sloughs dominated by pond apple or Carolina ash, often with abundant epiphytes. FRESHWATER FORESTED WETLANDS - floodplains or depressions dominated by hydrophytic trees CYPRESS/TUPELO - dominated entirely by cypress or tupelo, or these species important in the canopy; long hydroperiod. Dome Swamp (G4/S4) - small or large and shallow isolated depression in sand/marl/limestone substrate with peat accumulating toward center; occurring within a fire-maintained community; seasonally inundated; still water; statewide excluding Keys; occasional or rare fire; forested, canopy often tallest in center; pond cypress, swamp tupelo. Gum Pond - underlain by a clay lens; generally occurs in upland pine; mainly Panhandle; longer hydroperiod and lower fire frequency than cypress-dominated dome swamps; dominated by swamp tupelo. Stringer Swamp - narrow linear swamps; occur within a pyrogenic community along intermittent streams that only flow during heavy rainfall; Panhandle; dominated by pond cypress. Basin Swamp (G4/S3) - typically large basin wetland with peat substrate; seasonally inundated; still water or with water output; Panhandle to central peninsula; occasional or rare fire; forest of cypress/tupelo/mixed hardwoods; pond cypress, swamp tupelo. Strand Swamp (G2/S2) - broad, shallow channel with peat over mineral substrate; situated in limestone troughs; seasonally inundated; slow flowing water; vicinity of Lake Okeechobee and southward; occasional or rare fire; closed canopy of cypress and mixed hardwoods; cypress, pond apple, strangler fig, willow, abundant epiphytes. Floodplain Swamp (G4/S4) - along or near rivers and streams with organic/alluvial substrate; usually inundated; Panhandle to central peninsula; rare or no fire; closed canopy dominated by cypress, tupelo, and/or black gum. Freshwater T idal Swamp - floodplain swamp a river mouth where occasional saltwater intrusion significantly affects vegetation composition; receives pulses of freshwater in response to tides; cypress absent or infrequent; closed/open canopy of swamp tupelo, pumpkin ash, sweetbay. HARDWOOD - dominated by a mix of hydrophytic hardwood trees; cypress or tupelo may be occasional or infrequent in the canopy; short hydroperiod. Baygall (G4/S4) - slope or depression wetland with peat substrate; usually saturated and occasionally inundated; statewide excluding Keys; rare or no fire; closed canopy of evergreen trees; loblolly bay, sweetbay, swamp bay, titi, fetterbush. Bay Swamp - large or small peat filled depression; mainly eastern Panhandle to central peninsula; forested; dominated by bay species. South Florida Bayhead - on tree islands in glades marsh on peat substrate; south of Lake Okeechobee in central and southern peninsula; open or closed canopy; swamp bay, sweetbay, dahoon, coastalplain willow, and/or coco plum. Hydric Hammock (G4/S4) - lowland with sand/clay/organic soil over limestone or with high shell content; mesic-hydric; primarily eastern panhandle and central peninsula; occasional to rare fire; diamond-leaved oak, live oak, cabbage palm, red cedar, and mixed hardwoods. Coastal Hydric Hammock - occurring adjacent to coastal marshes; central Panhandle to central peninsula; species composition limited by occasional salt water intrusion; cabbage palm, red cedar, and live oak. Prairie Hydric Hammock - isolated stands of hydric hammock within a pyrogenic community, usually floodplain marsh; shelly sand soils; central and southern peninsula; occasional fire; cabbage palm, live oak, red cedar. Bottomland Forest (G4/S3) - flatland with sand/clay/organic substrate; usually connected or adjacent to a riverine community; occasionally inundated; Panhandle to central peninsula; rare or no fire; closed canopy of mixed hardwoods; deciduous or mixed deciduous/evergreen; tuliptree, sweetbay, water oak, sweetgum, diamond-leaved oak, red maple, loblolly pine, spruce pine, Atlantic white cedar. Alluvial Forest (G4/S3) - floodplain with alluvial substrate of sand, silt, clay or organic soil; inundated yearly during growing season; influenced by disturbance from ongoing floodplain processes (deposition of point bars, creation of “ridge and swale” topography); Panhandle to central peninsula; rare or no fire; closed canopy of mainly deciduous trees; water hickory, overcup oak, diamond-leaved oak, green ash, American elm, water locust, river birch. MARINE and ESTUARINE VEGETATED WETLANDS - intertidal or supratidal zone dominated by herbaceous or woody halophytic vascular plants; salinity >0.5 ppt. Salt Marsh (G5/S4) - estuarine wetland on muck/sand/or limestone substrate; inundated with saltwater by daily tides; statewide; occasional or rare fire; treeless, dense herb layer with few shrubs; saltmarsh cordgrass, needle rush, saltgrass, saltwort, perennial glasswort, seaside oxeye. Salt Flat - salt marsh with much exposed bare soil on slightly higher areas within marsh; high salinity and dry conditions; sparse and stunted cover of succulents and/or shoregrass. Mangrove Swamp (G5/S4) - estuarine wetland on muck/sand/or limestone substrate; inundated with saltwater by daily tides; central peninsula and Keys; no fire; dominated by mangrove and mangrove associate species; red mangrove, black mangrove, white mangrove, buttonwood. Buttonwood Forest - upper tidal area dominated by buttonwood; often transitional to rockland hammock. Keys Tidal Rock Barren (G3/S3?) - flatland with exposed limestone in supratidal zone; restricted to Keys; no fire; open, mainly herbaceous vegetation of upper salt marsh species and stunted shrubs and trees; buttonwood, christmasberry, perennial glasswort, saltwort, seashore dropseed, shoregrass. PONDS and LAKES (LACUSTRINE) - non-flowing wetlands of natural depressions lacking persistent emergent vegetation except around the perimeter Clastic Upland Lake (G3/S2) - generally irregular basin in clay uplands; predominantly with inflows, frequently without surface outflow; clay or organic substrate; Panhandle to northern central peninsula; colored, acidic, soft water with low mineral content (sodium, chloride, sulfate); oligo-mesotrophic to eutrophic. Coastal Dune Lake (G2/S1) - basin or lagoon influenced by recent coastal processes; predominantly sand substrate with some organic matter; Panhandle; salinity variable among and within lakes, and subject to saltwater intrusion and storm surges; slightly acidic, hard water with high mineral content (sodium, chloride). Coastal Rockland Lake (G2/S1) - shallow basin influenced by recent coastal processes; predominantly barren oolitic or Miami limestone substrate; southern peninsula and Keys; salinity variable among and within lakes, and subject to saltwater intrusion, storm surges and evaporation (because of shallowness); slightly alkaline, hard water with high mineral content (sodium, chloride). Flatwoods/Prairie Lake (G4/S3) - generally shallow basin in flatlands with high water table; frequently with a broad littoral zone; still water or flow-through; sand or peat substrate; statewide except extreme southern peninsula and Keys; variable water chemistry, colored to clear, acidic to slightly alkaline, soft to moderately hard water with moderate mineral content (sodium, chloride, sulfate); oligo-mesotrophic to eutrophic. Marsh Lake (G4/S4) - generally shallow, open water area within wide expanses of freshwater marsh; still water or flow-through; peat, sand or clay substrate; statewide except Keys; variable water chemistry, but characteristically highly colored, acidic, soft water with moderate mineral content (sodium, chloride, sulfate); oligo-mesotrophic to eutrophic. River Floodplain Lake (G4/S2) - meander scar, backwater, or larger flow-through body within major river floodplains; sand, alluvial or organic substrate; statewide except extreme southern peninsula and Keys; colored, alkaline or slightly acidic, hard or moderately hard water with high mineral content (sulfate, sodium, chloride, calcium, magnesium); mesotrophic to eutrophic. Sandhill Upland Lake (G3/S2) - generally rounded solution depression in deep sandy uplands; panhandle to southern peninsula; predominantly without surface inflows/outflows; typically sand substrate with organic accumulations toward middle; clear, acidic moderately soft water with varying mineral content; ultra-oligotrophic to mesotrophic. Sinkhole Lake (G2/S2) - typically deep, funnel-shaped depression in limestone base; statewide; predominantly without surface inflows/outflows, but frequently with connection to the aquifer; clear, alkaline, hard water with high mineral content (calcium, bicarbonate, magnesium). Swamp Lake (G4/S3) - generally shallow, open water area within basin swamps; still water or flow-through; peat, sand or clay substrate; statewide except Keys; variable water chemistry, but characteristically highly colored, acidic, soft water with moderate mineral content (sodium, chloride, sulfate); oligo-mesotrophic to eutrophic. RIVERS and STREAMS (RIVERINE) - natural, flowing waters from their source to the downstream limits of tidal influence and bounded by channel banks Alluvial Stream (G4/S2) - lower perennial or intermittent/seasonal watercourse characterized by turbid water with suspended silt, clay, sand and small gravel; Panhandle; generally with a distinct, sediment-derived (alluvial) floodplain and a sandy, elevated natural levee just inland from the bank. Blackwater Stream (G4/S3) - perennial or intermittent/seasonal watercourse characterized by tea-colored water with a high content of particulate and dissolved organic matter derived from drainage through swamps and marshes; statewide except Keys; generally lacking an alluvial floodplain. Seepage Stream (G3/S2) - upper perennial or intermittent/seasonal watercourse with clear to lightly colored water derived from shallow groundwater seepage; panhandle to southern peninsula. Spring-run Stream (G2/S2) - perennial watercourse with deep aquifer headwaters and clear water, circumneutral pH and, frequently, a solid limestone bottom; panhandle to central peninsula. SUBTERRANEAN - twilight, middle, and deep zones of natural chambers overlain by the earth's crust and characterized by climatic stability and assemblages of trogloxenic, troglophilic, and troglobitic organisms. Aquatic Cave (G3/S3) - cave permanently or periodically submerged; often supporting troglobitic crustaceans and salamanders; includes high energy systems which receive large quantities of organic detritus and low energy systems; statewide. Terrestrial Cave (G3/S2) - cave lacking standing water; often supporting bats, such as Myotis spp., and other terrestrial vertebrates and invertebrates; includes interstitial areas above standing water such as fissures in the ceiling of caves; statewide. MARINE and ESTUARINE - subtidal, intertidal, and supratidal zones of the sea, landward to the point at which seawater becomes significantly diluted with freshwater inflow from the land. (The distinction between the Marine and Estuarine Natural Communities is often subtle, and the natural communities types found under these two community categories have the same descriptions. For these reasons they have been grouped together.) MINERAL BASED Consolidated Substrate (G3/S3) - expansive subtidal, intertidal, and supratidal area composed primarily of nonliving compacted or coherent and relatively hard, naturally formed mass of mineral matter (e.g., coquina limerock and relic reefs); statewide; octocorals, sponges, stony corals, nondrift macrophytic algae, blue-green mat-forming algae, and seagrasses sparse, if present. Unconsolidated Substrate (G5/S5) - expansive subtidal, intertidal, and supratidal area composed primarily of loose mineral matter (e.g., coralgal, gravel, marl, mud, sand and shell); statewide; octocorals, sponges, stony corals, nondrift macrophytic algae, blue-green mat-forming algae and seagrasses sparse, if present. FAUNAL BASED Coral Reef (G2/S1) - expansive subtidal area with elevational gradient or relief and occupied primarily by living sessile organisms of the Class Hydrozoa (e.g., fire corals and hydrocorals), Class Anthozoa, abd Subclass Zoantharia (e.g., stony corals and black corals); southern peninsula and Keys; includes deepwater bank reefs, fringing barrier reefs, outer bank reefs and patch reefs, some of which may contain distinct zones of assorted macrophytes, octocorals, & sponges. Mollusk Reef (G3/S3) - substantial subtidal or intertidal area with relief from concentrations of sessile organisms of the Phylum Mollusca, Class Bivalvia (e.g., mollusks, oysters, & worm shells); statewide; octocorals, sponges, stony corals, macrophytic algae and seagrasses sparse, if present. Octocoral Bed (G2/S1) - expansive subtidal area occupied primarily by living sessile organisms of the Class Anthozoa, Subclass Octocorallia (e.g., soft corals, horny corals, sea fans, sea whips, and sea pens); likely statewide; sponges, stony corals, nondrift macrophytic algae and seagrasses spares, if present. Sponge Bed (G2/S2) - expansive subtidal area occupied primarily by living sessile organisms of the Phylum Porifera (e.g., sheepswool sponge, Florida loggerhead sponge and branching candle sponge); statewide; octocorals, stony corals, nondrift macrophytic algae and seagrasses sparse, if present. Worm Reef (G1/S1) - substantial subtidal or intertidal area with relief from concentrations of sessile, tubicolous organisms of the Phylum Annelida, Class Polychaeta (e.g., chaetopterids and sabellarids); southern peninsula (east coast only); octocorals, sponges, stony corals, macrophytic algae and seagrasses sparse, if present. FLORAL BASED (mainly subtidal) Algal Bed (G3/S2) - expansive subtidal, intertidal, or supratidal area, occupied primarily by attached thallophytic or mat-forming prokaryotic algae (e.g, halimeda, blue-green algae); statewide; octocorals, sponges, stony corals and seagrasses sparse, if present. Seagrass Bed (G3/S2) - expansive subtidal or intertidal area, occupied primarily by rooted vascular macrophytes, (e.g., shoal grass, halophila, widgeon grass, manatee grass and turtle grass); statewide; may include various epiphytes and epifauna; octocorals, sponges, stony corals, and attached macrophytic algae sparse, if present. COMPOSITE SUBSTRATE Composite Substrate (G3/S3) - expansive subtidal, intertidal, or supratidal area, occupied primarily by natural community elements from more than one natural community category (e.g., grass bed and algal bed species; octocoral and algal bed species); statewide; includes both patchy and evenly distributed occurrences. Torreya State Park (Liberty County) Photo by Gary Knight HARDWOOD FORESTED UPLANDS - mesic or xeric forest dominated mainly by hardwood trees Slope Forest Description: Slope forest is a well-developed, closed canopy forest of upland hardwoods on steep slopes, bluffs, and in sheltered ravines within the Apalachicola River drainage. Slope forests have extremely high tree and shrub diversity,328 largely because of their mixture of cold temperate and warm temperate elements.145 Tree density is relatively high, inducing much competition for space, water, sunlight and nutrients. The mostly deciduous canopy commonly includes American beech (Fagus grandifolia), Florida maple (Acer saccharum ssp. floridanum), white oak (Quercus alba), tuliptree (Liriodendron tulipifera), Shumard’s oak (Q. shumardii), white ash (Fraxinus americana), black oak (Q. velutina). Several evergreen species are common as well, including southern magnolia (Magnolia grandiflora), spruce pine (Pinus glabra), live oak (Q. virginiana), and laurel oak (Q. hemisphaerica). The diverse understory can be moderately dense to sparse and includes smaller canopy species plus American witchhazel (Hamamelis virginiana), needle palm (Rhapidophyllum hystrix), American holly (Hex opaca), eastern redbud (Cercis canadensis), mountain laurel (Kalmia latifolia), oakleaf hydrangea (Hydrangea quercifolia), basswood (Tilia americana), Florida anisetree (Illicium floridanum), sourwood (Oxydendrum arboreum), Gulf Sebastian bush (Sebastiania fruticosa), white fringe tree (Chionanthus virginicus), flowering dogwood (Cornus florida), horse sugar (Symplocos tinctoria), red buckeye (Aesculus pavia), silky camellia (Stewartia malacodendron), Florida yew (Taxus floridana), Ashe’s magnolia (Magnolia ashei), pyramid magnolia (Magnolia pyramidata), and the historically dominant but now declining Florida torreya (Torreya taxifolia)361 The herbaceous groundcover is often sparse and composed mainly of shade-tolerant species and spring ephemerals such as partridgeberry (Mitchella repens), Florida yam (Dioscorea floridana), woodland pinkroot (Spigelia marilandica), saw greenbrier (Smilax bona-nox), wild blue phlox (Phlox divaricata), sarsaparilla vine (Smilax pumila), prostrate blue violet (Viola walteri), heartleaf noseburn (Tragia cordata), switchcane (Arundinaria gigantea), trilliums (Trillium spp.), Christmas fern (Polystichum acrostichoides), and fringed campion (Silene polypetala). The combination of densely shaded slopes and cool, moist microclimates produces conditions that are conducive for the growth of many plant species that are more typical of the Piedmont and Southern Appalachian Mountains.361 These include mountain laurel, black walnut (Juglans nigra), wild hydrangea (Hydrangea arborescens), sweet-shrub (Calycanthus floridus), burningbush (Euonymus atropurpureus), heartleaf (Hexastylis arifolia), common maidenhair fern (Adiantum capillus-veneris), smooth Solomon’s seal (Polygonatum biflorum), liverleaf (Hepatica nobilis), white baneberry (Actaeapachypoda), perfoliate bellwort (Uvulariaperfoliata), bloodroot (Sanguinaria canadensis), false hellebore (Veratrum woodii), Canadian lousewort (Pedicularis canadensis), wild comfrey (Cynoglossum virginianum), downy rattlesnake plantain (Goodyera pubescens), American bladdernut (Staphylea trifolia), and eastern leatherwood (Dirca palustris). Slope forest occurs in areas with substantial topographic relief. Soils are generally composed of sands, sandy-clays, or clayey-sands with substantial organics and occasionally calcareous components. The Cody Scarp crosses the range of slope forest near its southern extent along the Big Sweetwater Creek.361 The soils above this divide are clayey Miocene soils while the soils to the south are Pleistocene sandy soils. Sandy soils are generally well drained, but clayey soils may shed much of the rainfall and exhibit significant surface water runoff. Thus, soil erosion is often a combination of seepage erosion, which occurs largely from the valley floors up (steepheads), and surface erosion, which occurs largely from the hilltops down.439 Slope forests along the Apalachicola River are included in one of the six biodiversity hotspots in the United States designated by The Nature Conservancy.388 These are relictual forests noted for their admixture of rare plants, coastalplain species, and species more common further north. , Ravines along the Apalachicola River north of the Cody Scarp remained above sea level during the Pleistocene, providing a safe refuge for southward-moving northern species. Since the waters of the Apalachicola River originate in the Appalachian Mountains many northern species had a direct conduit to the south during interglacial periods. The cool microclimate created by the narrow, shaded ravines allowed for their persistence during warm periods. Characteristic Set of Species: American beech, Florida torreya, Florida yew, Ashe’s magnolia, croomia, fringed campion, eastern leatherwood, Shumard’s oak, Florida maple Rare Species: Slope forest is well known for its high diversity of rare plants (approximately 41 species) within a very restricted geographic location. Two Florida endemic tree species, Ashe’s magnolia and Florida yew, are present in slope forest. , Rare plants characteristic of slope forest include fringed campio, Florida torreya, Florida yew, croomia (Croomia pauciflora), burningbush, bay star-vine (Schisandra glabra), Baldwin’s spiny-pod (Matelea baldwyniana), pyramid magnolia, Ashe’s magnolia, eastern leatherwood, narrow-leaved trillium (Trillium lancifolium), liverleaf, wood spurge (Euphorbia commutata), Godfrey’s privet (Forestiera godfreyi), American bladdernut, northern prickly ash (Zanthoxylum americanum), Florida flame azalea (Rhododendron austrinum), and green violet (Hybanthus concolor). Rare animals that occupy slope forest are Apalachicola dusky salamander (Desmognathus apalachicolae), copperhead (Agkistrodon contortrix), hairy woodpecker (Picoides villosus), and worm-eating warbler (Helmitheros vermivorum). Several rare invertebrates are found in Florida slope forests including the Torreya pygmy grasshopper (Tettigidea empedonepia) and floodplain phanaeus scarab beetle (Phanaeus triangularis). The Apalachicola hydroptila caddisfly (Hydroptila apalachicola), is restricted to a single slope forest ravine. Range: Slope forest is restricted to a 35 km stretch along the eastern side of the Apalachicola River in the northern Florida Panhandle and southern Georgia.361 In Florida, slope forest occurs south of Lake Seminole in Gadsden and Liberty counties, from the Georgia state line to just north of Bristol, Florida, roughly following the range of Florida torreya.361 Natural Processes: Succession is generally restricted to single tree canopy gaps. Canopy damage on a larger scale can result from occasional hurricanes and strong storms.19 Fire is rare in these protected mesic forests.83 Slope forest has undergone a drastic change since the 1950s, the near extinction of one of its dominant, understory species, the Florida torreya.361 The decline of the Florida torreya is thought to be caused by a fungal infection, although no specific pathogen has been identified to date.361 It is unclear and probably too early to tell what longterm effects the absence of this tree will have on the community. Community Variations: Slope forest can vary with topographic location. Higher elevations with deep sandy soils and thinner leaf mantles may exhibit nearly xeric soil conditions. Lower elevations on slopes near cool streams, or where seepage is prevalent, tend to be cooler, and soils may be nearly hydric. Slight changes in soil moisture along the slope gradient are often reflected by different plant species.57,224,361 For example, Florida torreya, Florida anise, mountain laurel, and tuliptree tend to be associated with lower slopes in soils with higher moisture and organic content, while mockernut hickory, laurel oak, and live oak tend to be associated with upper slopes of better-drained, drier soils.57,361 The mid-slope is usually dominated by mesic forest species such as American beech, southern magnolia, and Florida maple. Associated Communities: Apart from its narrow geographic range, slope forest may be distinguished from upland hardwood forest by steep slopes, a richer diversity of species, and a diversity of species more common to the Appalachian region further north. Two rare species, Florida torreya and Florida yew are endemic to this community type. Slope forest and upland hardwood forest are mesic communities that can occur in close proximity to one another. Slope forest is often associated with, and grades into, upland pine or sandhill at their upper elevations, and baygall or floodplain communities at their lower elevations. Seepage streams commonly occur along the valley floors of slope forest. Management Considerations: Slope forests are sensitive to direct physical disturbances and to hydrological manipulations that affect seepage and surface water sources. Their steep slopes are highly susceptible to erosion when un-vegetated or damaged. Common disturbances include logging, development, foot or vehicular traffic, and feral hog rooting. Unsightly refuse dumps are frequently located in slope forest ravines and steepheads. This refuse can bury or damage vegetation and impact stream water quality. Impoundments of streams within ravines can also destroy slope forest on adjacent lower slopes. The unique assemblage of slope forest plants and animals attracts many outdoor enthusiasts. Uncontrolled collecting by hobbyists and professionals could, however, significantly impact populations of some plants and animals. Slope forests are very rare, and should be protected diligently from human-related disturbances. It is also important that adjoining upland communities be maintained. Disturbances such as logging in these uplands can lead to accelerated erosion in the slope forest below.385 Invasive exotic species can be a problem even in the highest quality slope forest. Species that often invade these forests include coral ardisia (Ardisia crenata), Chinese privet (Ligustrum sinense), Japanese climbing fern (Lygodium japonicum), heavenly bamboo (Nandina domestica), and silverthorn (Elaeagnuspungens). Exemplary Sites: Apalachicola Bluffs and Ravines Preserve (Liberty County), Torreya State Park (Liberty County) Global and State Rank: G2?/S1 Crosswalk and Synonyms: The 1990 Natural Community Guide126 defined slope forest in a more general sense based primarily on topography. Many forests in the Florida Panhandle and northern peninsula (e.g., Eglin Air Force Base in Okaloosa, Washington, and Santa Rosa counties, and Goldhead Branch State Park in Clay County) that were previously classified as slope forest would be classified as upland hardwood following this update. Davis included in 113/southern floodplain forest 4/mixed hardwoods and pines 12/hardwood forests SCS 5/mixed hardwood and pine 11/upland hardwood hammocks Myers and Ewel SAF FLUCCS Temperate hardwood forests - relict forests 82/loblolly pine - hardwood 431/beech - magnolia 434/hardwood - conifer mixed 438/mixed hardwoods 439/other hardwoods Kuchler 112/southern mixed forest Other synonyms: ravine; steephead224; mesic hardwood hammock, magnolia beech climax forest83; torreya ravines, bluff and slope forests436 Blue Creek Ravine, with seepage stream (Alachua County) Photo by Katy NeSmith Upland Hardwood Forest Description: Upland hardwood forest is a well-developed, closed-canopy forest dominated by deciduous hardwood trees on mesic soils in areas sheltered from fire. It typically has a diverse assemblage of deciduous and evergreen tree species in the canopy and midstory, shade-tolerant shrubs, and a sparse groundcover. Characteristic canopy trees include southern magnolia (Magnolia grandiflora), pignut hickory (Carya glabra), sweetgum (Liquidambar styraciflua), Florida maple (Acer saccharum ssp. floridanum), live oak (Quercus virginiana), laurel oak (Q. hemisphaerica), swamp chestnut oak (Q. michauxii), southern hackberry (Celtis occidentalis), white ash (Fraxinus americana), and loblolly pine (Pinus taeda). Species commonly found in Florida Panhandle and northern peninsula but not farther south include American beech (Fagus grandifolia), white oak (Q. alba), and spruce pine (Pinus glabra). The midstory layer is composed of younger canopy species as well as small trees, and tall shrubs, such as American holly (Ilex opaca), red bay (Persea borbonia), American hornbeam (Carpinus caroliniana), gum bully (Sideroxylon lanuginosum), devil's walkingstick (Aralia spinosa), eastern hophornbeam (Ostrya virginiana), flowering dogwood (Cornus florida), eastern redbud (Cercis canadensis), horse sugar (Symplocos tinctoria), American strawberrybush (Euonymus americanus), silverbells (Halesia spp.), winged elm (Ulmus alata), black cherry (Prunus serotina), basswood (Tilia americana). The groundcover is composed of shade-tolerant herbs, graminoids, and vines, such as partridgeberry (Mitchella repens), Virginia creeper (Parthenocissus quinquefolia), violets (Viola spp.), sedges (Carex spp.), sarsaparilla vine (Smilaxpumila), ebony spleenwort (Asplenium platyneuron), woodsgrass (Oplismenus hirtellus), and longleaf woodoats (Chasmanthium laxum var. sessiliflorum). Trilliums (Trillium spp.) can be found in the groundcover in the Panhandle and northern peninsula. Upland hardwood forest occurs on rolling mesic hills, slopes above river floodplains, in smaller areas on the sides of sinkholes, and occasionally on rises within floodplains. Limestone or phosphatic rock may be near the surface. Soils are generally sandy clays or clayey sands with substantial organic and sometimes calcareous components. These soils have higher nutrient levels than the sandy soils prevalent in most of Florida. The moisture retention properties of clays and layers of leaf mulch conserve soil moisture and create decidedly mesic conditions. The dense canopy and multiple layers of midstory vegetation restrict air movement and light penetration, which maintains high relative humidity within the community. Characteristic Set of Species: flowering dogwood, American beech, southern magnolia, white oak, horse sugar Rare Species: Over fifty species of rare plants occur in upland hardwood forest throughout its range. These include heartleaf (Hexastylis arifolia), pygmy-pipes (Monotropsis reynoldsiae), sweet-shrub (Calycanthus floridus), trout lily (Erythronium umbilicatum), several species of spiny-pods (Matelea floridana, M. alabamensis, M. flavidula), trailing arbutus (Epigaea repens), and the endemic Marianna columbine (Aquilegia canadensis var. australis). Several animal species reach their southern limit within upland hardwood forest in northern Florida: four-toed salamander (Hemidactylium scutatum), copperhead (Agkistrodon contortrix), Louisiana waterthrush (Seiurus motacilla), American redstart (Setophaga ruticilla), worm-eating warbler (Helmitheros vermivorum), and eastern chipmunk (Tamias striatus). Range: Upland hardwood forest occurs throughout the southeastern coastal plain from the Carolinas to east Texas.328 Upland hardwood forest occurs in the Florida Panhandle south to the central peninsula.359,416 Upland hardwood forest most commonly occurs within the inland portions of the state. Natural Processes: Light gap succession is the driving force behind tree recruitment in upland hardwood forest and can happen at many different scales from single tree sized gaps to larger canopy openings.57,328 Localized damage from low intensity, naturally occurring fires that creep into the forest edges from surrounding pyrogenic upland communities (e.g., upland pine, sandhill) appears to be a natural part of the forest dynamics of upland hardwood forest; however, fires rarely burn completely through the understory, and even less frequently lead to crown or devastating fires.19 Hurricanes are another irregular natural process by which the canopy opens and the forest 19 canopy regenerates. Community Variations: Geographic differences in species composition can be noted through the range of this community. Upland hardwood forest of the Panhandle support several herbaceous species that are more common north of Florida, including woodland pinkroot (Spigelia marilandica), bloodroot (Sanguinaria canadensis), heartleaf (Hexastylis arifolia), May apple (Podophyllum peltatum), and several species of trilliums (Trillium underwoodii, Trillium decipiens). In the peninsula, the proportion of evergreens tends to increase and species richness in the canopy decreases as northern deciduous species (e.g. American beech, white oak) reach their southern limits.145,328 Upland hardwood forest is generally considered a mesic forest. However it may occur along a wide range of moisture conditions such as along upper slopes and ecotones along upland communities where it can be dominated by species tolerant of drier conditions (e.g., laurel oak, live oak). Upland hardwood forest occurring in the Mariana Lowlands (e.g., Florida Caverns State Park) is noted for its calcareous slopes and prominent limestone outcrops on rich clayey soils, and abundance of northern spring ephemeral species. , Upland hardwood forest species can invade pyrogenic communities where fire is excluded; however the forests that develop in this manner have a mixed species composition representing both community types.57 Upland mixed woodland may develop in the ecotone of upland hardwood forest and adjacent sandhill or upland pine in the Panhandle and northern peninsula. One common variant of upland hardwood forest occurs within Florida. Variant: Dry Upland Hardwood Forest - Occurring along upper slopes of upland hardwood or slope forest, on sandy ridges within more mesic upland hardwood forest, and mostly found in the northern peninsula where upland hardwood forests tend to be drier. This drier and more evergreen forest is dominated by laurel oak in the canopy and also includes live oak, pignut hickory (Carya glabra), southern magnolia, wild olive (Osmanthus americanus), Carolina laurelcherry (Prunus caroliniana), the rare Arkansas oak (Quercus arkansana), sparkleberry (Vaccinium arboreum), and horse sugar. This variant is distinguished from upland mixed woodland by its closed, evergreen canopy of laurel oak rather than an open mixed canopy of pines and deciduous species such as southern red oak (Quercus falcata) and sand post oak (Quercus margaretta). This variant may have a live oak component in the canopy but is not dominated by live oak; those forests dominated by live oak are usually better classified as mesic hammock. Associated Communities: Upland hardwood forest often is associated with and grades into upland pine, slope forest, bottomland forest, or hydric hammock. Upland hardwood forest and slope forest are very similar community types which share characteristic species (e.g., American beech, southern magnolia, white oak) and forest structure. Upland hardwood forest differs from slope forest primarily by its lack of the abundance and high diversity of rare species characteristic of slope forest (e.g., Florida torreya [Torreya taxifolia], fringed campion [Silenepolypetala], Florida yew [Taxus floridana]; see slope forest). Upland hardwood forest differs from mesic hammock by the presence of northern deciduous species in the canopy and subcanopy (e.g. Florida maple, white oak, white ash, swamp chestnut oak), a lack of cabbage palm (Sabalpalmetto) and saw palmetto (Serenoa repens), and a typically diverse canopy that is not dominated by live oak. Upland hardwood forest located in Central Florida, at the southern end of its range (particularly along the Brooksville Ridge), is often difficult to distinguish from mesic hammock as there is a greater overlap of species in this geographic area. Upland hardwood forest differs from hydric hammocks in that the latter generally occupy lower, flat topography, are dominated by hydrophytic species, and lack American beech and white oak. Upland hardwood forests often contain limestone outcrops. Management Considerations: Upland hardwood forest often occurs near streams, creeks, and rivers and can provide watershed protection.362 Common disturbances include logging, development, foot or vehicular traffic, and feral hog rooting. Unsightly refuse dumps are frequently located in upland hardwood forests. This refuse can bury or damage vegetation and impact stream water quality. Damage from invasive exotic plants and animals is a common problem in upland hardwood forest. Plant species that often invade these forests include coral ardisia (Ardisia crenata), Chinese privet (Ligustrum sinense), camphor tree (Cinnamomum camphora), Japanese climbing fern (Lygodium japonicum), and heavenly bamboo (Nandina domestica). Feral hog (Sus scrofa) foraging (rutting) damages soil and vegetation, may interfere with forest regeneration, and can lead to erosion problems, especially on slopes. Global and State Rank: G5/S3 Crosswalk and Synonyms: Kuchler Davis SCS Myers and Ewel SAF FLUCCS 112/Southern Mixed Forest 4/Mixed Hardwoods and Pines 12/Hardwood Forests 11/Upland Hardwood Hammocks 15/Oak Hammocks Temperate hardwood forests 76/Shortleaf Pine - Oak 431/Beech - Magnolia 434/Hardwood - Conifer Mixed 438/Mixed Hardwoods Exemplary Sites: Florida Caverns State Park (Jackson County), Wakulla Springs State Park (Wakulla County), Ichetucknee River State Park (Columbia County), Withlacoochee State Forest (Hernando County) Other synonyms: upland mixed forest in part 126; southern mixed hardwood forest19; mesic hardwood forest275 Salt Lake Wildlife Management Area (Brevard County) Photo by J. Richard Abbott Mesic Hammock Description: Mesic hammock is a well-developed evergreen hardwood and/or palm forest on soils that are rarely inundated. The canopy is typically closed and dominated by live oak (Quercus virginiana), with cabbage palm (Sabal palmetto) generally common in the canopy and subcanopy. Southern magnolia (Magnolia grandiflora) and pignut hickory (Carya glabra) may be occasional in the subcanopy. These species become less important where they reach their southern extent just north of Lake Okeechobee. South of this region, the overstory may contain a few tropical species such as gumbo limbo (Bursera simaruba) and satinleaf (Chrysophyllum oliviforme). Water oak (Q. nigra) and laurel oak (Q. hemisphaerica) may also be frequent in this community. Other than pignut hickory, only a few deciduous species such as sweetgum (Liquidambar styraciflua) and sugarberry (Celtis laevigata) are found in the canopy and subcanopy layers. Pine trees, particularly slash pine (Pinus elliottii) or loblolly pine (P. taeda), may form a sparse emergent layer. The shrubby understory may be dense or open, tall or short, and is typically composed of a mix of saw palmetto (Serenoa repens), American beautyberry (Callicarpa americana), American holly (Hex opaca), gallberry (I. glabra), sparkleberry (Vaccinium arboreum), hog plum (Ximenia americana), common persimmon (Diospyros virginiana), highbush blueberry (Vaccinium corymbosum), Carolina laurelcherry (Prunus caroliniana), yaupon (I. vomitoria), wild olive (Osmanthus americanus), and/or wax myrtle (Myrica cerifera). Tropical shrubs such as Simpson’s stopper (Myrcianthes fragrans), myrsine (Rapanea punctata), and wild coffee (Psychotria nervosa) are common in more southern mesic hammock. The herb layer is often sparse or patchy and consists of various graminoids, including low panic grasses (Panicum spp.), witchgrasses (Dichanthelium spp.), woodsgrass (Oplismenus hirtellus), longleaf woodoats (Chasmanthium laxum var. sessiliflorum), sedges (Cyperaceae), and whip nutrush (Scleria triglomerata), as well as various ferns and forbs such as bracken fern (Pteridium aquilinum) and partridgeberry (Mitchella repens). Toothpetal false rein orchid (Habenaria floribunda) and other ground orc