cause they are tropical plants confinedmostly to south Flnrida- Noneoccur in abundance and only one, whitestopper, is a tree. Isolated populations<strong>of</strong> this species occur north <strong>of</strong>Lake Okeechobee (Little 1978); one ispresent in hydric hammock in TosohatcheeState Reserve. <strong>The</strong> geographicaldistributions <strong>of</strong> several herbaceousplants common to hydric hammocks,including goldfoot fern andwild c<strong>of</strong>fee, extend north from theCaribbean to central Florida.A striking example <strong>of</strong> the influence<strong>of</strong> a species' range on the composition<strong>of</strong> hydric hammock is cabbage palm, thepredominant tree species in many hydric hammocks. From North Carol i na tonorthern Florida and west across thepanhandle, this species is essentiallyrestricted to the coast (Figure 28).South <strong>of</strong> a line from about Cedar Keyon the gulf coast to St. Augustine onthe Atlantic, cabbage palm rangesFigure 28. Distribution <strong>of</strong> cabbage palm inFlorida (from Little 1978). Asterisks on the insetmap indicate isolated populations <strong>of</strong> cabbagepalm outside <strong>of</strong> its continuous range (from Brown1973).across Florida' s peni nsul a. North <strong>of</strong>this line, a few interior populationsare scattered a1 ong rivers. <strong>Hydric</strong>hammocks in the northern region, forexample in San Felasco Hammock StatePreserve (Tab1 e 5), 1 ack cabbage palm.<strong>The</strong> inland Gulf Hammock site that wesampled (Table 5) is situated at theedge <strong>of</strong> cabbage palm's coastal range.Cabbage palm was absent from much <strong>of</strong>the forest, but it was common, especiallyin the understory, in someparts.Numerous studies have demonstratedor inferred that patterns <strong>of</strong> vegetationin southern forested wetlands arestrongly influenced by flooding(Bedinger 1978; Huffman and Forsythe1981; Wharton et a7. 1982; Leitman eta7. 1983). Frequency, duration,depth, and timing <strong>of</strong> flooding are factorsthat affect the species composition<strong>of</strong> a forested wetland. <strong>The</strong> effects<strong>of</strong> flooding on these communitiesare mediated by physical and chemicalchanges in the soils and varying responsesto the alterations by plantspecies.<strong>The</strong> nature and extent <strong>of</strong> the physicaland chemical processes that followsoil inundation largely depend on theduration <strong>of</strong> submergence and on soilproperties (Ponnamperuma 1984). Whena soil is flooded or saturated, gasexchange between the soil and air isgreatly restricted. <strong>The</strong> slowing <strong>of</strong>gas diffusion is particularly great insoils with high clay content. Oxygensupply to the soil is drastically cut,and roots and microorganisms depletethe oxygen in the soil water veryrapidly. Gases produced by soi 1metabol ism (e.g., carbon dioxide) accumulate.Anaerobic conditions inducea number <strong>of</strong> chemical changes in thesoil, many <strong>of</strong> which are detrimental toplant growth. Nitrate is replaced byammonium, <strong>of</strong>ten less preferred for uptakeand assimilation by plantspecies. Oxidized forms <strong>of</strong> iron, manganese,and sulfur are reduced to potentiallytoxic forms, including sulfides. Ethanol, another potent i a1toxin, is a by-product <strong>of</strong> anaerobicrespiration in most plant roots.
A sequence <strong>of</strong> changes in plantmat a hnl i grn and ~hvsi 01 ogi ca9 orocessesfollow the onset <strong>of</strong> anaerobiosis inwater1 ogged soi 1 s (Kozl owski 1984a,b) . Decreased water adsorption andclosure <strong>of</strong> the stomata result in a4 slowed rate <strong>of</strong> photosynthesis. Rootpermeabi 1 i ty is reduced, affecting theImovement <strong>of</strong> ions including nutrients.Leaf chlorosis and abscission, and retardedplant growth (especiallyheight-growth) foll ow. Morphologicalchanqes, such as the formation <strong>of</strong>aerenchyma tissue and the growth <strong>of</strong>adventitious roots, may take place.If flooding is prolonged, the plantdies. Flood-to1 erant plants, characteristic<strong>of</strong> wetlands, possess a variety<strong>of</strong> morphological and physiologicaladaptations (reviewed in Hook andCrawford 1978) that avoid or mitigateflooding stresses. <strong>The</strong>se adaptationscommonly facil ite oxygen flux to theroots and enhance the ability <strong>of</strong> plantroots to respire anaerobical ly withoutharmful effects.Plant responses to flooding arestrongly infl uenced by the duration,timing, and depth <strong>of</strong> inundation, butthe condition <strong>of</strong> the floodwater alsois significant. Flowing water is betterto1 erated than standing water,presumably because <strong>of</strong> the higher concentrations<strong>of</strong> dissolved oxygen in theformer. <strong>The</strong> effects <strong>of</strong> flood durationare clear from the preceding paragraph;changes in plant physiology becomeprogressively more severe withincreased period <strong>of</strong> anaerobiosi s. <strong>The</strong>adverse effects <strong>of</strong> fl ooding are exacerbatedduring the growing season whenoxygen demands are greatest. On theother hand, flooding during the dormantseason has re1 atively 1 i ttl e impacton the physiology and survival <strong>of</strong>most tree species ,(Gill 1970). Depth<strong>of</strong> inundation is critical to plantsurvival because oxygen diffusion tothe roots is greatly slowed by passagethrough water. Water depth with respectto plant height also affectsplant response to flooding. Most Shumardoak and sweetgum seedlings survived60 days when flooded only to theroot collar (Hosner and Boyce 1962),but all died after 20 days <strong>of</strong> completesubmersion (Hosner 1960). Water deepenough to cover most <strong>of</strong> the plant de-creases 1 ight intensity and interfereswith stomata1 function.<strong>The</strong> ability to withstand floodingvaries among plant species, sometimesamong populations, and with plant ageand plant size (Whitlow and Harris1979). Most assessments <strong>of</strong> the relativetolerance <strong>of</strong> woody plants t<strong>of</strong>looding are based on the results <strong>of</strong>experimental inundation <strong>of</strong> seed1 ingsand reservoir flooding <strong>of</strong> establishedforests. Results <strong>of</strong> studies examiningthe responses <strong>of</strong> wetland hardwoodtrees were summarized by Gill (1970),Teskey and Hinckley (1977), Whitlowand Harris (1979), and McKnight et al.(1981). Some tree species common tohydri c hammocks were included. Caremust be taken in applying the results<strong>of</strong> these tests to interpretations <strong>of</strong>natural communities. Reservoir floodingis considerably less erratic thannatural flooding and, in nature, numerousfactors interact with inundationto influence vegetation patterns.Nevertheless, some conclusions fromstudies <strong>of</strong> flood tolerance help in explainingplant distributions withinand among hydric hammocks.McKnight et a1. (1981) assignedsouthern bottomland trees to four toleranceclasses that varied in two regards:length <strong>of</strong> time during thegrowing season that the species canwithstand flooded or saturated soils,and the extent <strong>of</strong> anatomical and physiologi cal adaptations. Only one commonhydri c-hammock species, swamp tupelo, was considered to be tolerant.This species can survive long periods<strong>of</strong> inundation, and its seeds remainviable when submerged in water formonths. Most hydric-hammock species(red maple, sweetgum, Florida elm,loblolly pine, sweetbay, persimmon,green ash, swamp l aurel oak) are moderatelytolerant <strong>of</strong> flooding; somemorphol ogical or physiol ogical adaptationsto flooding may develop (Figure29), but they do not enable the treeto survive flooding for an entiregrowing season. Mature trees <strong>of</strong> sixhydric-hammock species remainedhealthy when flooded for 17%-37% <strong>of</strong>the growing season; red maple withstoodthe longest flooding period, and
- Page 2 and 3: Copies of this publication may be o
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- Page 8 and 9: FIGURESNumber1AL...............Dist
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