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Kuchler Type: Cypress savanna
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].

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Information Courtesy: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory. Fire Effects Information System

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