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KUCHLER TYPE: Cypress savanna
KUCHLER-TYPE-NUMBER : K091 PHYSIOGNOMY : Open forest to savanna with needleleaf deciduous trees, some broadleaf evergreen trees or shrubs. Trees and shrubs also in groves surrounded by open grasslands [31]. OCCURRENCE : Kuchler's description of cypress savanna is similar to Davis' description [7] of cypress swamps in southern Florida. Davis classified southern Florida by physiographic regions and recognized four vegetation types containing baldcypress (Taxodium distichum) or pondcypress (T. d. var. nutans) in southern Florida. Three of these correspond to generally recognized types of stillwater cypress swamps in southern Florida: dwarf cypress savanna, cypress domes (also called cypress heads or cypress ponds), and cypress strands [6,7,17]. In addition, mixed swamps (also called mixed hardwood swamps or mixed swamp forests) contain cypress, although cypress is usually not dominant [7,17]. Kuchler's description and species list for cypress savanna appears to include the three cypress-dominated types. Throughout this writeup, where information pertains to all three types the term cypress swamp will be used. Where information is specific to a particular type, the abovementioned type names will be used. Kuchler's definition of cypress savanna appears to include cypress domes, strands, and dwarf cypress savanna that occurs in southern Florida in Big Cypress Swamp, Fakahatchee State Preserve, Corkscrew Swamp Sanctuary, and Everglades National Park. Cypress domes also occur in Okefenokee Swamp and within pine flatwoods in northern Florida. These domes are not on Kuchler's map of cypress savanna; whether this is because of size or because he did not include them in the definition is unknown. This writeup will focus on cypress domes, strands, and dwarf cypress savanna occurring in southern Florida. The cypress domes occuring in northern Florida and in Okefenokee Swamp are slightly different from those described by Kuchler, but they contain many of the same species and have similar ecological processes and are thus included in this writeup [8,31]. Information derived from cypress domes elsewhere will be included where it is deemed relevant since many of the species and ecological processes appear to be similar. This writeup does not include cypress swamps on alluvial soils which occur over the southeastern Atlantic Coastal Plain from the Carolinas to Louisiana. STATES: FL COMPILED BY AND DATE : Janet Sullivan, November 1994 LAST REVISED BY AND DATE : NO-ENTRY AUTHORSHIP AND CITATION : Sullivan, Janet. 1994. Cypress savanna. In: Remainder of Citation
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PHYSIOGRAPHY : Big Cypress Swamp is a large, basin-like area, with low limestone outcrops, numerous sloughs, shallow ponds, and prairies on sand or marl (unconsolidated calcitic clay). It forms a discrete hydrological unit [6,11]. Big Cypress Swamp drains south and west through sloughs, strands, and culverts under the Tamiami Trail. During periods of low water levels, water is impounded in numerous ponds [6]. Elevation ranges from 12 to 40 feet (3.6-12 m) above mean sea level in the northern part of Big Cypress to sea level where it grades into mangrove (Rhizophora mangle, Avicennia germinans, or Laguncularia racemosa) swamps [6]. The formation of cypress domes, strands, and dwarf cypress savannas is driven primarily by hydrology. Cypress domes develop in depressions within large watersheds. The characteristic circular shape is probably created by the slow dissolution of the underlying limestone over years of acid water percolation [19]. The water level in cypress domes normally fluctuates dramatically once or twice during the year [17]. In Big Cypress Swamp and in the Everglades, cypress domes are underlain by marl and limestone bedrock [8]. Cypress strands form where high water level and sufficient flow generate a depression channel. At low water there may be no discernible flow [17]. Corkscrew Swamp, a cypress strand, is an elongate depression in the mineral soil with very little relief (less than 4 inches/mile [6cm/km]). The ground surface in Corkscrew Swamp is more irregular in the deeper parts of the strand due to mounds formed by stumps, logs, litter, burned-out holes in peat, and root wells [9]. Dwarf cypress savannas are open stands of stunted cypresses (also called hatrack cypress) with an understory of grasses. They occur on sites with a medium-length hydroperiod (6-9 months). Rainfall is the most important source of water for dwarf cypress savannas [6,17]. CLIMATE : The climate of southern Florida is moist and mild; it is frost-free nearly all year. Mean annual precipitation is around 60 inches (1,524 mm), 80 percent of which falls from May to October, creating distinct wet and dry seasons [3]. Precipitation received in 1 year ranges from 30 to 100 inches (762-2,540 mm) [3]. Occasionally, drought in summer (normally the wet season) can result in complete cypress defoliation which normally does not occur until fall [17]. SOILS : In southern Florida, cypresses are found on a variety of soils including organics, sands, marls, and rock lands [7], with pH usually in the range of 6 to 8. Cypress swamp soils are characterized by Coultas and Duever [43]. Soils in Big Cypress Swamp are mostly derived from Tamiami limestone and quartz sands. The soils are usually pure sand, marl, or mixtures, ranging from 2 to 24 inches (5-61 cm) deep on limestone bedrock. Dwarf cypress savanna occurs on thin marl, 3 to 6 inches (7.6-15 cm) deep [6] Outcrops of dense, fine-grained limestone are scattered within the cypress savanna ecosystem [6,23]. VEGETATION : There is no standard nomenclature for the vegetation associations of southern Florida [42]. Cypress swamps occur in the oak-gum-cypress Forest and Range Ecosystem. Garrison and others [23] described the alternating wetlands and drylands west of the Everglades as cypress savanna. Craighead [6] defined a number of physiographic provinces for southern Florida. His descriptions include cypress domes, strands, and dwarf cypress and are similar to those of Davis [7]. Wade and others [42] listed 10 vegetation associations for southern Florida and included cypress as a single type. Long and Lakela [32] described five physiographic provinces for southern Florida, listing Big Cypress as a single province. The Florida Natural Areas Inventory [22] included three natural community types that correspond to cypress swamps. Baldcypress dominates alluvial floodplain forests, and pondcypress is dominant in cypress domes [37]. It has been indicated by some authors, however, that both species occur in cypress domes and strands [6,9,17,]. It is thought that distributional differences may be due differences in tolerance to acidity [34], or because pondcypress is more drought tolerant than baldcypress [1]. Many authors do not attempt to discriminate between the two taxa, partly because of controversy over their taxonomic status and partly because of the difficulty of distinguishing them in the field. Throughout this writeup, where baldcypress and pondcypress have not been differentiated by the authors or where the information is applicable to both taxa, the term cypress will be used. Plant associates in cypress swamps vary with depth, duration, and frequency of flooding, soil type, geographic location, and stand density [17]. Cypress domes and strands have few associates in the center; most of those present are shade-tolerant species of epiphytic bromeliads, orchids, ferns, and nettles, many of which do not occur in the United States outside of Florida [6,17,42], or aquatic macrophytes such as tall flag (Thalia geniculata) and arrowhead (Sagittaria spp.) [7,42]. Ground cover is sparse in the center [42]. On the perimeter of cypress domes and strands, associates include buttonbush (Cephalanthus occidentalis), redbay (Persea borbonia), sweetbay (Magnolia virginiana), magnolia (Magnolia spp.), cocoplum (Chrysobalanus icaco), dahoon (Ilex cassine), myrsine (Myrsine florida), southern bayberry (Myrica cerifera), Coastal Plain willow (Salix caroliniana), and Florida poisontree (Metopium toxiferum) [6]. Slash pine (Pinus elliottii) is occasionally present in cypress domes, particularly in northern Florida [17,19,20]. Water tupelo (Nyssa sylvatica var. biflora) is also present in northern cypress domes [10,20]. Red maple (Acer rubrum), pond apple (Annona glabra), strangler fig (Ficus aureus), water ash (Fraxinus caroliniana), swamp bay (Persea palustris), and paurotis palm (Paurotis wrightii) are frequent in cypress domes and strands [6,7,17,44]. Spanish moss (Tillandsia usneoides) is common in more northerly cypress domes. Other species of Tillandsia are more common in central and southern Florida. Resurrection fern (Polypodium polypodioides) is another common epiphyte [5]. In open stands grading to dwarf cypress savanna, sawgrass (Cladium jamaicense) may be an important component on the higher layers of peat. Occasionally palmettos (Sabal palmetto or Serenoa spp.) are present, and greenbriars (Smilax spp.) are conspicuous. Herbs present in cypress domes and strands include sweet rush (Cyperus haspan), maidencane (Panicum hemitomon), beakrushes (Rhynchospora spp.), spikerushes (Eleocharis spp.), arrowweeds (Pluchea spp.), eupatorium (Eupatorium spp.), fingergrasses (Chloris spp.), swamp fern (Blechnum serrulatum), small yellow bladderwort (Utricularia juncea), and leather ferns (Achrostichum spp.) [6,7,31,42]. Algal mats often cover the soil surface in the wet season [42]. WILDLIFE : Cypress swamps play a unique role in animal ecology. They do do not have a distinct fauna, sharing many species with adjacent plant comunities. Most species and individuals spend only part of their lives in the swamp [17,23,28]. Benthic invertebrates form the base of the food chain. A high diversity of invertebrates has been recorded for cypress domes and dwarf cypress savannas. In cypress domes, immature chironomids (Diptera) dominate the benthic fauna [21]. Wild turkeys (Meleagris gallopavo) and other predators feed on crayfish in cypress swamps [17] In summer, reptiles and amphibians dominate cypress swamp vertebrate communities; in winter the vertebrate fauna is dominated by birds. High temperatures allow herpetiles to remain active through the winter [28]. Jetter and Harris [45] trapped 23 species of reptiles and amphibians in three cypress domes from 1974 to 1976. The most numerous group was ranid frogs, particularly southern leopard frog (Rana utricularia). Other common species included cricket frogs (Acris spp.), oak toad (Bufo quercicus), green treefrog (Hyla cinerea), and cottonmouths (Agkistrodon spp.). Undisturbed cypress domes in Bradford County, Florida, contained (in addition to southern leopard frog) little grass frog (Pseudacris ocularis), central newt (Notophthalmus viridescens louisianensis), pine woods treefrog (Hyla femoralis), striped mud turtle (Kinosternon baurii), and slender dwarf siren (Pseudobranchus striatus spheniscus). Amphibian species outnumbered reptiles [28]. Bird densities are higher in cypress swamps in winter than during breeding season, largely due to the presence of wintering birds that breed elsewhere. Birds using cypress swamps include wild turkey, ibis (Plegadis spp.), double-crested cormorant (Phalacrocorax auritus), herons (Ardea herodias, Butorides striatus, Egretta caerulea, Nycticorax nyticorax, and Nyctorax violaceus), snowy egret (Egretta thula), great egret (Casmerodius albius), anhinga (Anhinga anhinga), and belted kingfisher (Ceryle alcyon) [23]. The limpkin (Aramus guarauna) is a characteristic occupant of cypress swamps due to the availability of snails. Canopy-feeding passerines are common, but there are usually only a few mid-story species such as red-bellied woodpecker (Melanerpes carolinus), red-headed woodpecker (M. erythrocephalus), pileated woodpecker (Drycopus pileatus), tufted titmouse (Parus bicolor), and great crested flycatcher (Myiarchus crinitus). The wood duck (Aix sponsa) is common in the larger cypress domes that contain sufficiently large trees [28]. Swainson's warbler (Limnothlypis swainsonii) and prothonotary warbler (Protonotaria citrea) are also typical members of the cypress swamp avian community. Endangered bird species occurring in cypress swamps include Bachman's warbler (Vermivora bachmanii) and southern bald eagle (Halieetus leucocephalus leucocephalus). Mississippi kite (Ictinia mississippiensis) and American swallow-tailed kite (Elanoides forficatus) breed only in swamps, including cypress swamps [17]. The globally endangered (extinct in the United States) ivory-billed woodpecker (Campephilus principalis) once inhabited cypress swamps [23,28]. Most mammals occurring in cypress swamps occupy ecotones. Mammals include white-tailed deer (Odocoileus virginiana), gray fox (Urocyon cinereoargenteus), fox squirrel (Sciurus niger), raccoon (Procyon lotor), Virginia opossum (Didelphis marsupialis), striped skunk (Mephitis mephitis), eastern cottontail (Silvilagus floridana), and swamp rabbit (S. aquaticus) [23,28]. River otter (Lutra canadensis) and bobcat (Lynx rufus) are common residents of large cypress domes [28]. There are many rodents and shrews [23] including southeastern shrew (Sorex longirostris), short-tailed shrew (Blarina brevicauda), cotton mouse (Peromyscus gossypirus), hispid cotton rat (Sigmodon hispidus), and golden mouse (Ochrotomys nutalli). The latter nests in trees to avoid floodwaters [17,28]. Away from the center of cypress domes small mammals include marsh rabbit (Silvilagus palustris) and wood rats (Neotoma spp.) in addition to species mentioned previously. Arboreal mammals include southern flying squirrel (Glaucomys volans), eastern gray squirrel (Sciurus carolinensis), and several species of bats [28]. Rare and endangered species, and species whose ranges formerly included cypress swamps, include mangrove fox squirrel (Sciurus niger avicenna), elk (Cervus canadensis), black bear (Ursus americanus), Florida panther (Felis concolor), mink (Mustela vison), and gray wolf (Canis lupus) [17,23,28]. ECOLOGICAL RELATIONSHIPS : Successional Processes: Cypresses are dependent on regular water level fluctuation for successful germination and establishment [17]. Cypress stands become established on open sites during periods of drought [27,42,44]. Conditions that exist following crown-killing fires that do not consume the surface peat layer are conducive to cypress establishment [27]. In the absence of fire, cypresses are succeeded by hardwoods. Cypresses are not the most rapidly growing trees in swamps; hardwoods, particularly water ash and red maple, produce more wood (basal area) relative to their biomass than cypresses [9]. Cypress dome establishment within sawgrass marshes (i.e., the Everglades) can proceed on batteries of peat: peat masses that float loose from the substrate and provide a relatively dry site suitable for colonization. Peat batteries may be initially colonized by buttonbush, hurrahbush (Lyonia lucida), or dahoon. These shrubs stabilize the mass and increase the rate of peat formation, favoring subsequent invasion by bays (Persea spp.) and cypresses. The resulting community is similar to that found in cypress domes in closed depressions. These communities are usually referred to as tree islands or cypress islands and usually succeed to hardwoods in a short period of time [5]. Community Structure: Distinct seasonal communities dominate cypress swamps. The wet season flora dominants include species adapted to growing under flooded conditions; a different group of species grows during periods when there is no standing water [9]. Dwarf cypress savannas do not usually have standing water in late winter, but are wet the remainder of the year [6]. Duever and others [11] reported a hydroperiod for dwarf cypress savanna of 120 days, with a maximum water depth of 6 to 8 inches (15-20 cm). On a dwarf cypress savanna study site in Big Cypress Swamp, the water table was above ground level for approximately 4 months between July and November. Maximum depth of surface water was 4 inches (10 cm). In March of the dry season, maximum depth to the water table was 40 inches (103 cm) [3]. In dwarf cypress savannas, cypress seedlings become established in wet years but grow very slowly [6]. Trees are usually less than 12 feet (3.4 m) tall and have disproportionately large buttressed trunks. Many of these small trees are over 100 years old; trees the same size in nearby cypress domes may be only 25 years old. Trees are often 50 to 65 feet (15-20 m) apart [6]. Cypress domes are small, roughly circular, forested wetlands that occur in poorly drained depressions [5,7,17,42]. In cypress domes and strands, decomposition rates are slow and peat accumulates [17]. In some cypress domes the peat layer is up to 20 feet (6 m) deep; a record depth of 96 feet (29 m) was reported for one cypress dome [7,11]. Most of the cypress domes in southern Florida have a central pond, surrounded by the tallest trees [6]. The cypress canopy may be up to 60 feet tall (18 m) around the central pond with almost complete crown closure [42]. Further away from the pond, the trees are shorter. Some authors report that the shorter trees are younger [7]. However, Craighead [6] stated that the trees in the middle are not much older than the trees on the periphery; in some cases, the trees on the shallow soil of the periphery are older. This occurs for two reasons: the lower fertility of the soil on the periphery and the lower likelihood of severe fire due to a smaller quantity of fuel. Trees are killed in the center during peat-consuming fires. The higher fertility in the center contributes to relatively rapid growth rates for new trees established after the fire. Further discussion of fire and community structure is in FIRE EFFECTS ON VEGETATION [6,9,42]. Cypress domes sometimes expand and grow together to form meandering cypress strands [11]. Cypress strands are found along major drainageways, mostly oriented north-south. Cypress strands have a well developed layer of peat, up to 6 feet (2 m) deep [11]. Cypress strands have a hydroperiod of 8 months or more [10], and are sometimes flooded year-round [42]. Normal wet season water levels in Corkscrew Swamp may fluctuate from 1.6 to 5 feet (0.5 to 1.5 m) deep and there is usually a measurable but slow flow rate. Water level may be relatively stable in a wet year (with dry season rainfall) but may drop as much as 6.6 feet (2 m) in a drought year [9]. The trees in the middle of cypress strands are larger than those in domes and may be 100 to 115 feet (30-35 m) tall and 6 feet (1.8 m) in diameter above the butt swell. Strands intergrade with dwarf cypress savanna on the edge. Trees near the periphery of the strand are usually smaller than trees in the center, probably for reasons similar to those for similar structure in cypress domes. The best correlation between tree size and environmental factors was found to occur with peat depth; larger trees tended to be found on deeper peat [9]. The relationship of tree size and fire is discussed further in FIRE EFFECTS ON VEGETATION. In Gordon Swamp (a small strand on the edge of Corkscrew Swamp Sanctuary), cypresses near the center of the strand grew 50 percent faster than trees near the edge for the first 50 years of growth. Trees over 150 years old had ring widths that were similar in all parts of the strand. It was hypothesized that young trees were better able to take advantage of improved site quality in the center of the strand than older trees [9]. In cypress domes and strands hardwood species contribute to the formation of a thick layer of peat. With the gradual reduction of cypress in older stands due to lack of regeneration, hardwoods increase in dominance, and the stand becomes a mixed swamp (a transitional stand), and then a bay head (hardwood swamp stand dominated by bays [Persea spp.]) [6]. Nutrient Status and Community Productivity: Nutrient inflow to dwarf cypress savanna is extremely low. For example, total phosphorus inflow, which is solely via rainfall, is approximately 0.1 g/sq m/year [1]. Nutrient concentrations in dwarf cypress savanna surface water were reported: Nitrate nitrogen ranged from 0.07-0.18 mg/L and total phosphorus ranged from 0.005-0.013 mg/L [3]. An increase in nutrient flow can occur when lower water levels allow more rapid decomposition and nutrient mineralization [17]. Spanish moss extracts nutrients from incident rainfall and may play an important role in nutrient cycling [5]. There is not an appreciable contribution by nitrogen-fixation [17]. Gross primary productivity and net primary productivity are low in cypress swamps, which are supplied with nutrients primarily by rainfall and have little nutrient storing capacity. Dwarf cypress savanna is rated the lowest in productivity, and cypress domes and strands are slightly higher. Estimated gross primary productivity for southern Florida cypress strands is approximately 6.74 g of carbon/sq m/day; and for southern Florida dwarf cypress savanna gross primary productivity is 1.82 to 2.41 g of carbon/sq m/day. Aboveground biomass productivity is also low: 0.5 kg/sq m/year [17]. Mean basal area increment for dwarf cypress savanna trees ranged from 7.2 to 12.7 sq cm/year [3]. Biomass, productivity, and water relations were discussed for Fakahatchee Strand [4].


KUCHLER TYPE: Cypress savanna
FORESTRY VALUES : Most old-growth cypress swamps (and other swamps) in Florida were logged by 1950. The most valuable product was the old-growth baldcypress which contained large volumes of very durable hardwood. Current logging is in second-growth, relatively young cypress, which is less durable. It is primarily used for chipping; until recently cypress was primarily used for specialty items such as crab traps and ladders. [17,33]. RANGE VALUES : WILDLIFE VALUES : Cypress swamps provide food, nesting sites, hibernation sites, and cover for a variety of species. Rare and endangered birds and mammals are more likely to be found in cypress swamps and mixed hardwood swamps than in other kinds of swamps. Cypress swamps are amoung the few areas not colonized by humans in southern Florida [17]. In Big Cypress Swamp, 6 of 19 bird species listed as rare, endangered, or threatened use cypress swamps for breeding and feeding [8]. Cypress swamps commonly contain rookeries of wood storks (Mycteria americana), herons (nine species and subspecies), and double-crested cormorants [17]. The wood stork is dependent on Florida wetlands. It nests in cypress stands or mangrove stands, and its range is now largely restricted to Florida [15]. Other rare birds found in cypress swamps include short-tailed hawk (Buteo brachyurus), southern bald eagle [36], and osprey (Pandion haliaetus). The roseate spoonbill (Ajaja ajaja) is found in dwarf cypress savannas [17]. The Florida panther, which is endangered, is restricted to Big Cypress Swamp and the Everglades [15]. Ewel [17] subjectively rated the contributions of various cypress swamp parameters to bird and mammal habitat. All three types of cypress swamps were rated low in canopy insect production, low in edible seed and fruit production, and low in vegetative density. Cypress domes and strands were rated high in cavity density, but dwarf cypress savannas were rated low. All three types were rated high for presence of water. Dwarf cypress savannas are important for their sparse cover; this community provides perching sites for raptors and wading birds that search for small organisms in an open environment [17]. OTHER VALUES : Cypress swamps recharge groundwater and play a role in regional flood control. Cypress domes in northern Florida are under study for wastewater treatment areas [18,19]. Nutrient enrichment from wastewater dumping result in immediate response among floating aquatics and herbaceous species. Net photosynthesis in cypresses growing in sewage-treated domes increased over a 6-year study, but there was no net increased tree growth [5,40]. Other uses of cypress swamps include harvesting of peat and phosphate mining. Reclamation after phosphate mining usually converts the site to planted grass pasture [17]. A detailed review of cypress swamp values has been published [18]. MANAGEMENT CONCERNS : Currently, most of the extant cypress swamps in southern Florida are under management or protection in Big Cypress National Preserve, a 920 square mile (2,300 sq km) area. Other preserves in which cypress swamps occur include Corkscrew Swamp Sanctuary, Fakahatchee State Preserve, and Everglades National Park [19]. Southern Florida has the youngest flora of any area in the United States; perhaps 3,000 to 5,000 years have passed since establishment of vegetation on newly emerged land. It has been hypothesized that since the flora is so young, there are many unfilled niches which therefore facilitate invasion by exotic species [41]. Melaleuca (Melaleuca quinquenervia) invasion affects groundwater levels in cypress swamps through increased transpiration [17]. Other impacts of melaleuca invasion are discussed in FIRE MANAGEMENT CONSIDERATIONS. Many cypress swamps have been drained, logged, impounded, or experienced other disturbances. Drainage allows invasion by species with low flood tolerance, and often results in significant decrease in primary productivity [17]. Drainage of cypress swamps causes a shift in the frequency distribution of reptile and amphibian species but no differences in abundance or richness [28]. Cypress swamps in northern Florida had poor cypress regeneration, increased shrub density, hardwood invasion, and increased fire potential after drainage [17]. Logging activities change drainage patterns due to the construction of dikes, roads and tramways, and the loss of trees (with concomitant lower transpiration rates). Regeneration after logging in cypress swamps is usually dominated by hardwoods. Water level changes and lack of cypress seed are responsible for the lack of cypress regeneration [17]. Structural characteristics change with logging of large old-growth cypress and affect bark-gleaning birds, reptiles, amphibians, and large arboreal mammals such as raccoons. These changes include loss of nest sites for cavity nesting species. Species that forage in open areas are temporarily benefited. Logging slash favors certain species of herpetiles, small mammals (i.e., cotton mouse), and birds (i.e., rufous-sided towhee). Breeding birds respond favorably to edges created by clearcutting pinelands that surround cypress swamps. Individual density of animals, number of species, species diversity and density are all higher in sharply defined edges than in ecotones [28]. Alexander and Crook [46] documented vegetation changes in southern Florida over the last 16 to 30 years, on 100 mile square (160 km square) quadrats. They concluded that the natural ecosystem will continue to lose its diversity and ability to maintain itself with any resemblance to the pre-1940 condition unless ways are found to return and properly distribute more water to wild habitats, control exotics, and manage fire.


KUCHLER TYPE: Cypress savanna
FUELS, FLAMMABILITY, AND FIRE OCCURRENCE : Fuels: On the driest sites, rapid decomposition and occasional flooding prevent organic matter accumulation [17]. Most of Big Cypress Swamp, however, is covered with sufficient fuel to carry fire. Occasional freezes may influence fuel characteristics by killing back the cold-sensitive vegetation and increasing litter [39]. Mean standing stock of litter in a dwarf cypress savanna was 344 (+/- 94) grams per square meter [3]. In years of average precipitation, fires may burn in grasslands and dwarf cypress savanna, but will not burn very far into the middle of cypress domes or strands. Conversely, in a drought year fire may be more severe in the centers of domes and strands than on the edges. When peat is dry enough to burn, the larger amount of litter and organic material in the center will support a more severe fire than on the edges. Fires that consume peat may kill trees due to root damage and loss of structural support. Fires are rarely severe enough to kill trees in dwarf cypress savanna because of sparse organic matter [3,6]. Flammability: In cypress swamps, ericaceous evergreen shrubs are particularly flammable community members [17]. Cypress domes and strands and hardwood swamps probably constituted natural barriers to the spread of wildfire in presettlement times; fire burning in grasslands and savannas around cypress domes would burn into the dome only as far as soil and vegetation moisture levels would permit. Drainage of these wetlands has increased the flammability of cypress domes [42]. Fire During Presettlement Human Occupation: Human occupation occurred within a few hundred years of vegetation establishment in southern Florida [39,24]. Most cypress swamps in southern Florida contain evidence of past fires [42]. It is generally assumed that lightning fires have always been an important factor in southern Florida fire regimes. Prior to European settlement, human-caused fire may have substantially increased fire frequency over the background frequency of lightning fires [14,38,39]. It is likely that most Native American-caused fires were set in the dry season, as soon as fuels could carry fire [39]. Fire in the Postsettlement Period: The frequency of fires increased with European settlement of southern Florida. Fires were set for numerous reasons and many fires were started through carelessness or accident. Drainage of swamps increased the frequency and intensity of fires, with concomitant severe damage to peat exposed by drainage [41]. Fire Frequency: The Everglades and Big Cypress National Preserve are the most fire-prone ecosystems in the National Park Service system. Southern Florida experiences 70 to 90 thunderstorm days per year, the highest number of any area in the United States. The mean annual number of ground strikes is from 4 to 12 per square kilometer. Most of the cloud-to-ground strikes occur during the wet season and 87 to 93 percent of lightning fires occur from May to August; lightning fires have been recorded in all months except January and February. Fewer strikes but more lightning fires occur in the Everglades than in Big Cypress Swamp [39,41]. Taylor [41] estimated that the interval between extreme fire years (years in which a very large number of fires or large number of acres burned) in the Everglades is 5.8 to 7.5 years, and that there is a moderate fire season every 3.2 years. In southern Florida, widespread and severe fires occurred in more than one-third of the years between 1900 and 1952. In very dry years, organic soil fires occasionally burned from one dry season to the next, through the normally wet summer months. Between 1948 and 1979, there were 682 fire reports and 451,082 burned acres in the Everglades. For the first 21 months of fire records for Big Cypress (starting in 1979), there were 131 reported fires, which burned 40,370 acres (16,716 ha). Many fires in Big Cypress burn out before discovery. Both Parks have been disturbed by drainage, logging, farming, off-road vehicle use, and invasion by exotic species, all of which affect fire frequency to some extent. Other changes in fire intensity, frequency, and type have come from fire suppression, prescribed fire programs, and incendiary fire. Wildfires and incendiary fires are extinguished [41]. Wet-season prescribed fires have replaced dry season lightning fires in many grasslands. Drainage has decreased hydroperiods and increased flammability of peat [5,41]. In Big Cypress Swamp, lightning-caused fire is minor relative to human-caused wildfire and prescribed fire. Human-caused wildfires occur in the dry season, and are strongly correlated with moisture conditions [39]. Of the 1,068 fires recorded in Big Cypress Swamp from 1979 to 1988, about four-fifths were incendiary or accidental [39]. The number of fires is positively correlated with peak backcountry use during wild turkey season in March and concides with dry conditions. "Sunday is a particularly popular day for starting fires" [39]. The average fire return interval for all community types in Big Cypress is approximately 12 years [39]. Cypress in dwarf cypress savanna show fire scars and may reveal fire frequency in the dwarf cypress type; however, the slow growth of the trees makes interpretation of rings difficult [41]. Snyder [39] estimated the fire return interval for dwarf cypress savanna as 24 years. Wade and others [42] reported an estimate of seven to nine fires per century in dwarf cypress savanna. Cypress-mixed hardwood strands and cypress domes have a fire return interval of 110 years according to Snyder [39], but Ewel [17] stated that they experience three to five fires per century. Strands appear to have longer fire rotations than domes or dwarf cypress savanna [42]. FIRE EFFECTS ON SITE : Peat based habitats such as cypress domes and strands can support both ground and surface fires. Fire may remove aboveground vegetation only or may also consume peat. FIRE EFFECTS ON VEGETATION : The effects of fire on southern Florida ecosystems are compounded by changing hydrologic relationships. Presettlement fires had different effects than current fires due to human-caused changes in drainage patterns, fire frequency, and the introduction of exotic species [42]. Cypresses survive surface fires better than competing hardwoods [20]. Low-severity surface fires therefore tend to maintain monospecific cypress stands by suppressing hardwood regeneration and killing young hardwoods [9,17]. For example, in Okefenokee Swamp, woody species invade peat islands between fires but most are killed by fire [12]. Young cypress trees (less than 200 years old) may sprout after fire if damage to roots is slight. Sprout production and viability decline with tree age [5]. The characteristic rounded shape of cypress domes and strands appears to be related to peat depth, fire frequency, and site conditions. In Corkscrew Swamp, trees are generally larger and older on sites with deep peat. The deeper peat is in contact with water for longer periods and dries out more slowly. Fires are therefore more frequent and more severe on the edges of domes and strands than in the center where peat is deepest. The less frequent the fire on a microsite, the more likely cypresses will survive to larger size. Outside the strand, hydrologic conditions favor fire that is frequent enough to kill young cypress trees before they can grow large enough to survive low-severity fire; the community is therefore maintained as grassland. In Gordon Swamp, a small cypress strand on the edge of Corkscrew Swamp Sanctuary, a different pattern was recorded. Peat depth increased towards the center of the strand, but tree size was uniform (rather than increasing in the center) and trees were younger in the center of the strand. This situation was probably caused when fire killed trees in the center but not the edges. Following fire, newly established cypresses grew faster than the trees on the edges of the strand. Better site conditions in the center may have contributed to faster tree growth; relative sizes of trees may be partly due to the poor condition of fire-damaged trees on the strand edge. Strands and domes probably expand and contract in response to fire occurrence and hydrologic conditions. During extreme drought, even large strands and domes may experience stand-replacing fire [12]. In a north-central Florida cypress dome, a wildfire killed most shrubs, hardwoods and pines, but killed less than 50 percent of the dominant cypress trees. By 3 years after the fire, the study site flora was composed almost completely of cypress, but the site was being colonized by hardwoods and shrubs [20]. Gunderson [25] proposed a conceptual succession model for southern Florida swamps that shows the effects of logging and fire on cypress and mixed hardwood swamps. After logging, mixed hardwoods and remnant cypress are maintained in the absence of fire. With a severe fire, willow (mostly Coastal Plain willow) colonizes open areas. Willows are maintained by repeated fire, but in the absence of fire succeed to mixed hardwoods. After logging, stands of mixed hardwoods and remnant cypress may experience low-severity fires, which favor cypress over hardwoods, and could return the stand to conditions approximating the prelogging conditions. In the absence of fire, cypress regeneration returns the stand to a mixed swamp (cypress and hardwoods, but with continued lack of fire succession to mixed hardwoods occurs). Cypress and mixed hardwood stands may be converted to monospecific cypress forests by surface fires; in the absence of fire, monospecific cypress stands are invaded by hardwoods. With severe fire, monospecific cypress stands and cypress and mixed hardwood stands are converted to willow and remnant cypress stands. Willow-remnant cypress stands may develop to monospecific cypress stands with occasional surface fires, if cypress regeneration occurs. More frequent or more severe fire that eliminates cypress regeneration will maintain a willow-remnant cypress stand [25]. Since cypress seed dispersal in limited, recovery from severe fires that consume peat may be very slow, particularly where water flow is slow to negligible. In areas where severe fires have burned old-growth cypress stands, there is little sign that communities are recovering to prefire conditions [5]. FIRE EFFECTS ON RESOURCE MANAGEMENT : Prescribed fire in and near cypress swamps is used mainly to enhance wildlife habitat, particularly for endangered species [39]. FIRE USE CONSIDERATIONS : Dye [13] reported on firing techniques in the Everglades and Big Cypress Swamp. In the past, prescribed fire in Big Cypress was usually conducted in the cooler months: November to April [39]. The current trend is to conduct prescribed fire in May and June during the "natural" lightning fire season, which fulfills the goal of approximating original conditions. Most fires are set as headfires; burn plots are small (600-1,000 ac [243-405 ha]); blacklines are wide; and multiple firing patterns are employed. Dye [13] recommended staggered headfires after the establishment of backing and single flankfires within zone perimeters. Plowing lines and the use of firing patterns leading to development of broad coalescing fire fronts are not recommended. FIRE MANAGEMENT CONSIDERATIONS : Low-severity fires do no apparent damage to dwarf cypress savannas and prevent invasion by hardwood species. Fuel build-up could result in crown scorch and cypress mortality. It is unlikely that wet season lightning fires ever occurred with any regularity in presettlement times; fuels were either inundated or too sparse to carry fire. Lightning fires were probably common whenever the water table dropped. Low-intensity, prescribed headfires are recommended for this type at approximately 10-year intervals [42]. Fire management in cypress domes and strands will be an incidental effect of prescribed fire in dwarf cypress savanna. Some domes will be moist enough to prevent fire spread, while others (in the same year) will carry fire. Prescribed fire in dwarf cypress savanna will, therefore, create a patchwork of fire that burns into domes to some extent. Because of reduced hydroperiods, however, some prescribed fires may burn severely in cypress domes. Fire management in cypress swamps must be "fine-tuned". Fire cannot be excluded, but cypress is vulnerable to severe fire [42]. In northern Florida, periodic fire in seasonally dry cypress domes will perpetuate cypress at the expense of pine [20]. An intense fire every 2 or 3 decades kills most pine and hardwoods, but leaves cypress. The removal of a thin layer of organic soil appears to be tolerated by cypresses and other species. It is possible that more frequent, low-intensity fires (every 2-3 years) could be used to achieve the same goal, although this has not been demonstrated [42]. The nature of fires in both northern and southern Florida has changed [17]. Drainage of southern Florida has resulted in soils that are dry early in spring which contributes to wildfires that are more destructive than fires that occurred before widescale drainage. Under current conditions, fires consume large amounts of organic soil and kill the roots of most fire-adapted plant species (as well as fire sensitive species) [42]. Fire in logged cypress swamps is more severe than in unlogged swamps due to remaining slash and the presence of dense regrowth. Fires burn hotter, destroy cypress seeds and roots in the soil, and eliminate cypress sprouts. This situation favors the replacement of cypresses by willows which are followed by mixed hardwoods [25]. The detrimental effects of severe fires in cypress swamps include smoke production (hazardous to health), loss of organic soils, loss of vegetation, damage to wildlife habitat, loss of esthetic value, and scars and damage caused by suppression efforts [42]. Melaleuca invasion occurs on sites that have been drained and severely burned. The flammability of melaleuca increases the probability of crown fires in cypress stands, particularly where melaleuca has invaded the ecotone between cypress stands and pine flatwoods [39,41]. Fire management programs in southern Florida will probably be designed to bring about desired effects rather than recreate a specific fire regime. Prescribed fire use is limited by public tolerance; visible smoke is produced, and high intensity fires are feared [39]. REHABILITATION OF SITES FOLLOWING WILDFIRE : NO-ENTRY

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