Regeneration of cypress, Taxodium distichum and Taxodium ascendens , in logged and burned cypress strands at Corkscrew S...
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Title: Regeneration of cypress, Taxodium distichum and Taxodium ascendens , in logged and burned cypress strands at Corkscrew Swamp Sanctuary, Florida
Physical Description: viii, 88 leaves : ill. ; 28 cm.
Language: English
Creator: Gunderson, Lance H., 1952-
Publication Date: 1977
 Subjects
Subjects / Keywords: Cypress   ( lcsh )
Corkscrew Swamp Sanctuary (Fla.)   ( lcsh )
Botany thesis M.S
Dissertations, Academic -- Botany -- UF
Genre: bibliography   ( marcgt )
non-fiction   ( marcgt )
 Notes
Thesis: Thesis (M.S.)--University of Florida.
Bibliography: Bibliography: leaves 84-87.
Statement of Responsibility: by Lance H. Gunderson.
General Note: Typescript.
General Note: Vita.
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Source Institution: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: aleph - 000011489
oclc - 03403657
notis - AAB3990
System ID: AA00004002:00001

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REGENERATIONOFCYPRESS. TAXODIUNDISTICHill1 AND TAXODIill'[ IN LCX;GEO ANDBURNEDCYPRESSSTRANDSAT CORKSCPE1[SliA!1P SANCTUARY. FLORIDA ByLANCEH.GUNDERSONA THESIS PRESENTEDTOTHEGRADUATECOUNCILOFTHEUNIVERSITYOFFLORIDAIN PAR'IIAL FULFILLHENT OF THEREQUIREr-rENTS FOR THEDEGREE OF HASTER OFSCIENCE UNIVERSITYOFFLORIDA1977

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ACKNOWLEDGEMErITS Theauthorwouldliketoextendthankstomembersofhissupervisorycommittee:Dr.A.E.LugoandDr.JohnEwel,chairman.Dr.Lugoreviewedandcommented onthemanuscript.Dr.Ewelprovidedexpertguidanceandassistancefromtheformulativestagesthroughthefinalproduct.Specialthanksaredue Dr.M.J.Duever,althoughanunofficialmember,actedaspartofthecommitteebyassistinginthefieldwork,criticizingthemanuscriptandprovidingencouragement.The work wassupportedbytheCenterforWetlands,UniversityofFlorida,undercontractsfromtheRANNdivisionoftheNationalScienceFoundationandtheRockefellerFoundationentitledCypressWetlandsforWaterManagement,RecyclingandConservation.SpecialthanksgototheNationalAudubonSocietyandthemanagementofCorkscrew SanctuaryforallowingtheresearchtobeconductedwithintheSanctuary.Manyotherpeopleassistedinvariousportionsofthework.EdCarlson,LarryRiopelle,AliceHumbertandGlenGundersonhelpedwithfieldwork.LindaDuever commented onthe artHork andmanuscript.Darrell Bro,m developedandprintedthephotographs.Mrs. FaySchattnerassistedintypingthefinaldraftofthemanuscript.ii

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TABLEOFCONTENTSA CKNOHLEDGEMENTS. ..................................iiLISTOFTABLES..............................,ivLISTOFFIGURES.........................................vABSTRACT.....................................viiINTRODUCTION.......................................1Logging................................2Fire....................5CypressAutecology..............6Objectives.,.,.9STUDYSITE...........................................11 ....................................1?VegetationDescription...........1?Seedlings..................20Seeds21WaterLevels....................2JAerialPhotographs..........2JRESULTS.......................................25VegetationInventory..............25WaterLevels.............J8Seedlings42Seeds49AerialPhotographs.,60DISCUSSION..................................68CypressRegeneration...........68SuccessionalStatusofCypressForests........??Management 81LITERATURECITED..., ; .,.,.,.,.,.,84BIOGRAPHICAL SKETCH ....................88iii

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LIST OF TABLES1.Importancevaluesforoverstoryspeciesattheburned.above-the-dikestudysite.............262.Importancevaluesforoverstoryspeciesatthelogged-and-burned,above-the-dikeandbelow-the-dikestudysites27 3. Importancevaluesforoverstoryspeciesatthelogged,above-the-dikestudysite......................................294.Importancevaluesforoverstoryspeciesatthelogged,below-the-dikestudysite...................................... 305. Importancevaluesofspeciesineachharvestofunderstoryplantsattheburned.above-the-dikestudysite 32 6.Importancevaluesofspeciesineachharvestofunderstoryplantsatthelogged-and-burned,above-the-dikestudysite..... 33 7.Importancevaluesofspeciesineachharvestofunderstoryplantsatthelogged-and-burned.below-the-dikestudysite.... J4 8.Importancevaluesofspeciesineach harvest ofunderstoryplantsatthelogged.below-the-dikestudysite 35 9.Importancevaluesofspeciesineachharvestofunderstoryplantsatthelogged.above-the-dikestudysite............... 36 10.Above groQnd understorybiomassduringthestudyperiod 37 11.Numberoflivingseedlingsattheburned,above-the-dikestudysite.5012.Numberoflivingseedlingsatthelogged-and-burned,above-the-dikestudysite......... 5213. Numberoflivingseedlingsatthelogged-and-burned,below-the-dikestudysite......................................5214. NWlwer oflivingseedlingsatthelogged,above-the-dikestudysite..................................................... 54 15.Numberoflivingseedlingsatthelogged,below-the-dikestudysite55iv

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LISTOFFIGURES1.LocationofCorkscrewSwamp insouthernFlorida...122.Habitattypesandrecent ofstudysiteswithinCorkscrew SWaJTJ.p Sanctuary................................................143.PhotographsofstudysitesatCorkscrewSwampSanctuarytakeninDeoember. 1976164.Schematicmapofsamplingprocedureateachstudysite...185. Hater levelsattheburned, above-the-dikestudysite.....................................................396. Hater levelsatthelogged-and-burned,above-the-dikestudysite.....................................................407. Hater levelsatthelogged-and-burned,below-the-dikestudysite......................................................418. Hater levelsatthelogged,but-not-burned,above-the-dikestudysiteo439. Hater levelsatthelogged,but-not-burned,beloH-the-dikestudysite.............................................. 1+410.HeightincreasesoftransplantedseedlingsatthestUdysites4611.NumberoftransplantedseedlingssurvlVlngtotheendofthestUdyperiodatthestUdysites.......4712.Numberoftransplantedseedlings Hith leavesfrom 21t Februaryto23October1976ateachstudysite....4813.Meangerminationofpondcypressandbaldcypressseeds SO,ill ateachstudysite................"5714.Germinationofpondcypressandbaldcypressseed SO,ill ingreenhouseflatsfollowingsoakingatstudysites...... 58 15.Cumulativegerminationofpondcypressand seedsinincubatorfollowingacidscarification.....5916.Vegetationmapconstructedfromaerialphotographstakenin1953.priortologgingor burning ..........62v

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17.Vegetationmapconstructedfromaerialphotographstakenin1963.nineyearsafterloggingandsixmonthsafterburning.6418.Vegetationmapconstructedfromaerialphotographstakenin1972,eighteenyearsafterloggingandnineyearsafterburning..................................................6619.Meansurvivorshipofcypressseed so,m atstudysites7120.Fateoftransplantedseedlingsandseedlingsgerminatedfromseedsownatstudysites.................................. 73 21.Summaryofcompetitioneffectsongrowthandsurvivaloftransplantedseedlings......................................7522.GeneralizedsuccessionschemeinsouthFloridaswamps.........78vi

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AbstractofThesisPresentedtotheGraduateCounciloftheUniversityofFloridain Partial FullfillmentoftheRequirementsfortheDegreeof i'faster ofScienceREGENERATIONOFCYPRESS. TAXODIUI1 DISTICHUMAND TAXODIUI1 ASCENDENS,INLOGGEDANDBURNEDCYPRESSSTHANDSAT CORKSCllEHSHANP SANCTUAllY,FLOllIDAByLanceH.Gunderson August 1977Chairman:JohnEwelMajorDepartment:BotanyRegenerationoftwospeciesof Taxodiumdistichum(L.)Richard(baldcypress)and TaxodiQm ascendensBrogn.(pondcypress),wasinvestigatedinsuccessionalseresresultingfromlogging (1954) andburning (196J) ofcypressstrandsatCorkscrew S1;amp Sanctuary,near naples, }i'lorida.Theeffectsofadikeconstructedin1969toimpoundwatertothenorthwerealsoexamined.SitesWereestablishedinareasthatwere:1)loggedandnorthofthedike,2)loggedandsouthofthedike,J)logged-and-burnedandnorthofthedike,4)logged-and-burnedandsouthofthe'dike,and5)burnedandnorthofthedike.Seedlingsof 1. ascendensweretransplantedin JanuarJ, 1976amongexistingvegetationandintoclearedareasateachstUdysite,andseed-lingheightsweremeasuredthroughOctober,1976.Seedsof 1. distichumand 1. ascendenswere sownateachstudysite.Germinationofseedwasmonitoredinthefieldandundergreenhouseandlaboratory conditions.Hater levelsweremonitoredateachsitefromOctober,1975throughOctober,1976.Vegetationmaps Here dra,ffi from sequential setsofaerial photographs todetermineratesofchangeofthevegetationaftervii

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disturbancesofloggingandburning.Therewas nosignificantdifferenceinthe ratesofcypressseedlingsamongthefivesites.Nodetectabledifferencewasmeasuredbetweenseedlingsgrowingamongexistingvegetationandseedlingsgrowinginclearedareas.Maximummeanheightincreaseofseedlingsmeasuredwas 80%. Survivaloftransplantedseedlings Has6&fo. Maximumgerminationof SOVillc;,rpress seedswas4.8%.Averagegermi-nationinthefieldwas 2.1%;lessthan 5% ofthosewhichgerminatedsurvivedthroughthewetseason.AttheloggedsiteshardHoodssuchasIlexcassine,Acerrubrum,Perseapalustris, ]>Iyri@ cerifera,andMyrsineguianensisarepresentintheoverstoryandareactivelyregenerating.Somecypressregenerationbycoppicingandseedisoccurringatthesesitesalso.Thelogged-and-burnedandburnedsitesaredominatedbySalixcaroliniana.Thereisnocypressregenerationatthelogged-and-burnedsites,butsomeregenera-tionisoccurringbycoppiceandseedattheburnedsite.Measurementsofwaterlevelsindicatelongerperiodsofinundationattheloggedandlogged-and-burnedsitesthanattheburnedsite.Thedikecreateslowerwaterlevelstothesouth,butdoesnotchangetheperiodofinundation.Vegetationmapsindicatethathardwoodspersistfollowinglogging.Colonization caroliniana follovls destructionofthehardwoodsbyfire.Lackofseedsources,immobilityofcypressseed,lowviability,andcrucialwaterrequirementsfor gerciQqtion seemtoberetardingcypressregeneration.Onceestablished,cypressisabletocompetewithexistingvegetation. __Chairman viii

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INTRODUCTIONSouthFloridacypressforests,dominatedbybaldcypressTaxodiumdistichum(1.)RichardandpondcypressTaxodiumascendensBrogn.,havebeendescribedbySmall(1920),Harper(1927),Davis(1943),andCraighead(1971).Davis(1943)describedthreetypesofcypressforests:domes,strandsandheads.Cypressdomesarefoundinshallow,circularly concave depressionsinthelimestonesubstrate(Craighead,1971).Cypressstrandsarefoundindeeper,elongateddepressions,andcypressheadsarecypressdomescontiguouswithstrandsnearthestrand Theperiodofinundationinacypressecosystemisafunctionoftopography,substrateandrainfall.The topographYofmostofsouthFloridaisveryflat(gradientsof8to16cm/km)andsurfacewaterflowisslow(0.8km/day)(Carteretal.,1973).Elevationsincypresssloughsmaybe1.5mlowerthansurroundingpineland.Peatsubstratespredominateincypressstrandsandtheincreasedwaterholdingcapabilityofpeatoverthewell-drainedsoilsofthesurroundingpinelandsandwetprairiesprolongsthehydroperiodinthestrands.Meanannualrainfallfor the BigCypressareaisfrom1400to1500mmwith60to 65% fallingfromJunetoAugust(Carteretal.,1973).Dueveretal.,(1975)monitoredcypressstrandecosystemsatCorkscrewSwampSanctuaryandfoundthatinundationusuallycommenceSinearlyJuneandlastsfrom 200to250days.Cypressstrandsareconsideredbysometobesubclimaxcommunities1

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2tomesic,mixed hardwoodforests(Davis,194};Craighead,1971)dominatedbytropicalspeciesinsouthernsouthFloridaandtemperatespeciesinthenorthernpartsofsouthFlorida(AlexanderandCrook,1973).Thetransitiontoclimaxstatusisaccomplishedbytheslowprocessofpeataccumulation which altersedaphicconditionspermittinghardwoodstobecomeestablishedandthriveinthemoremesicconditionsaroundthebasesofthecypresstrees(Penfound,1952).Duetothelonglifespanofcypresstreesandperiodicpeatremoval (byfireandwaterflow),however,cypressforestsusuallyremainascypressforestsforlongperiodsoftime.Thestrandecosystemsaresubjecttochange by amyriadoffactors,bothnaturalandanthropogenic.Naturalforceswhichbringaboutchangeincludefire,lightning,hurricanes,frostsandalligators(Davis,194};Craighead,1971).Man-induced changesthatinitiatesecondarysuccessionincludelumbering,andcanalandleveebuilding(Davis, 1943; AlexanderandCrook,1973).Thevalueofbaldcypressaslumberledtoextensiveloggingwhich removedalmostallofthelarge(greaterthan0.5mdiameter)treesfromsouthFlorida.Logging LoggingofbaldcypressintheBigCypressSwampareaofFloridawasinitiatedinthe1920's.Harvestingofthecypresspeakedinthemid1940'sandtaperedoffinthe1950's,whenmostofthelargetreeshadbeenremoved(Betts,1960).Hardwoodspecies,which weresubcanopytreesintheunloggedforest,dominatefollowinglumbering(AlexanderandCrook,1973).Nocypressreforestationprograms wereestablishedinFlorida.Inthe1940'stheuseofrailroadstocarrycypresslogstosawmillsbecamewidespreadinsouthFlorida.Theoperationbeganwithconstructionofearthendikesparalleltothelongaxisoftheswamp.Perpendicular

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3 spurswerebuiltintoadjacentcypressareasatintervalsof0.5km.Whiletherailswerebeinglaidonthesedikes,a crewofworkers Hould girdlethecypresstrees.Thegirdledtrees Hould beleft s"Ganding forseveralmonthspriortofelling,sothatthelogs Hould dryoutandfloat Hhen cut.Severedegradationofthelumber Has causedbyambrosiabeetleattacksduringthisperiod(Craighead,1971).Thetreeswerefelledduringhigh Hater andcutinto10mlengths Hhich Here skiddedtotherail Hay carsusinganoverheadcable.Largeskidtrails Here leftfromremovalbythismethod(Prestridge,1947),andmanyofthetramroadsandspursarestillapparenttoday.Reforestationofcut-overcypresswetlands(transplantingnursery grolID cypressseedlings)beganinLouisianain1948(BUll,1949)andcontinuedthrough 1951 (Rathborne,1951).Rathborne(1951)estimateda70-yearregrowthperiodfollowingthisoperation.Stubbs(1972)statedthatmostloggedcypress-tupeloforestshavenaturallyregenerated,buttupelo(NyssasPp.)maydominateinsteadofcypress.Stumpsproutsareimportantintheregenerationcypress-tupelo (Johnson,1972).Supplyingseedmaybenecessary,asuncutseedtreesandcoppicemayproduceinadequateamountsofseed(Stubbs,1972).Sternitzke(1971)suggestedthatthestandingstockofcypressinthesoutheasternU.S.increased 2% peryearthrough1968,mostofwhichis fromcutoverforests.ThesupplyofmarketabletimberinFloridaislargelypondcypress(T.ascendens):notregrowthofbaldcypress(T.distichum)(Sternitzke,1972).SuccessionfollowingLoggingSuccessionfollowinglumberingmaybedeterminedbytheseverityofthephysicaleffectsoftheloggingoperation.DeBelletal.(1968)

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4foundthat 60% ofallsaplingsand8%ofallsmalldiametertreesofoverstoryspecieswerekilledbyfellingandskiddingduringaselectivelumberingofbottomlandsinLouisiana.Remainingseedtreesandstumpsproutingmayresultinrapidregrowth follolfing cutting(Oosting, 19.56). Vegetationreproductionsuchasstumpsprouts,adventitiousshoots,andepicormicbranchesmayresultinrapid regrolrCh, butthe issusceptabletofungalinfectionandthe gr01"th formofthetreeswhichareleftisusuallynotgood(Smith,1962).Ifcullsaretheonlyremainingtrees,theproportionofundesirablegeneticstockinthecommunitymaybelargerthaninthepreloggingcommunity.Regenerationofwetlandforestsmaybe slolf if felf seedsourcesareleftandifseedgermi-nationrequiresveryspecificconditions(Stubbs,1972).Exposureoforganicsoils,andsubsequentsubsidenceofthesoillevel, was onere-sultoflogginginsouth Florida cypressstrands(AlexanderandCrook,1973).Thesechangesinthesoilduetologgingmayalterthesuccessionalpatterns.Mueller-DomboisandEllenberg's(1975)explanationofsuccessionafterlogging(usingEgler's (1954) termsof"relayfloristics"and"initialfloristiccomposition")maybeapplicabletopatternsobservedinsouth Florida swamps. Therelaypatternofspeciesreplacementisaresultofmodif'icationsofasiteby agroupofspeciesinsuchawaythattheirownsurvivalislimited,andthesurvivalofasucceedinggroupofspecieisenhancedwitharrivalofthesucceedingspecies'propogules.Theothertheory,"initialfloristiccomposition",attributesthepatternofspeciesreplacementtodifferencesingrowthratesandlifespansofthegroupsofspecies,withamajorityofpropogulesbeingpresentatthebeginningofthesere.Mostorallofthecypressseedsourcesmaybeexhaustedfollowingalumberingoperation.Followingthedisturbance,

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5secondarysuccessionproceedsrapidlythroughastagedominatedbythosefast-growinghardwoodspecieswhichremainaftertheloggingoperation.Succession may stagnateuntilproperconditionsandarrivalofcypressseed(relayfloristicsmechanisms)resultinthereestablishmentofcypressdoninatedcommunities.FireFireinnaturalforestedwetlandsmayoccurifthesubstratebecomesdry.Peat,evenwithitshigh retentioncapacity,becomesdryduringyearsof 10H rainfall,leavingthesystemvulnerabletofire(Craighead,1971).Firecouldbepartlyresponsibleformaintainingthecypressforestsinasubclimaxstageofsuccession(AlexanderandCrook,1973)bytheremovalofhardl,oods.Thehighwaterretentionpropertiesofpeatusuallycreateaneffectivebarriertofireduringthewetseasonandearlyintothe dry season(Craighead,1971),yetsurfaceburnsmayoccurearlyintothedryseason(Robertson,1953).Ifthemoisturecontentofthepeatgoesbelow30percent,thepeatmayignite,initiatingaslow-burningpeatfire(Craighead,1974).Theheatfromthefiremaydryoutunderlyinglayersofpeat Hhich mayinturnbe (AlexanderandCrook,1973).Thepeatfiremaycontinuetosmolderuntilitisextinguishedbyrainsormeetsanon-combustiblebarriersuchasthewatertable(Robertson,1953)orsandlayer(Cypert,1961).Succession folloHing BurningSuccessionafterafireinsouthFloridamayberelatedtothetimeoftheyearandseverityoftheburn.Fastmovingsurfacefiresmayconsumeunderstoryandshrubbyspecies(Cypert,1961),aswellasseedlingsandsaplingsofoverstoryspecies(Penfound,1952). HardHood speciesand

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6 upland conifersmayalsobeweededoutbysurfacefires.EwelandMitsch(1975),forexample, fOULdthat cypresssurvivedasurfaceburnina north centralFloridacypressdomebetterthanslashpine,Pinusel1icttii,andtupelo,Nyssaspp. gro,dng inthesame dome. Thespeciescompositionofa isdrasticallyalteredafteradeep peat burn.Adeepburn therootsoflargecypressandbaytrees(Cypert,1961)andeliminatesrecoverybysprouting(AlexanderandCrook,1973).Inopenwaterareas,Hhichmaybearesultofpeatremovalby the fire,bothrootedandfloatingaquaticplantsthenbecomepartofthesuccessionalvegetation(Penfound,1952).Thecharred stlli"pS arefociforcolonizationbynon-aquaticherbaceousinvaders(Beaven,1939).Fastgrowingshrubssuchastiti, C,Tilla racemiflora,andbuttonbush, Ce"h2. 1anthus occidentalis,areconspicuousinnorthernFlorida Sl,amps untilregenerationoftheoverstorycypressandbaytreesoccurs(Cypert,1973).FrequentfiresinsouthFloridaallowstandsofwillow,Salix toprevail(Robertson,1953;Loveless,1959;Craighead,1971).Fireinaloggedcypress swamp isoftenmoredevastatingthanfireinanunloggedswamp(Cypert,1961;AlexanderandCrook,1973)becausethe remining slashanddense undergro\,-th thatfollowsalumberingoperationburnsmorehotlythandoesthesparse undergr01fth ofuncutswamps(Cypert,1961).Theshallowrootsystemsofthe second-grolrl,h forestsare Eore likelytobedamaged by thehotterfires.Thesevereburnsmaydestroyseedsourcesandvegetative regro,rth, deterringsuccessionuntil pr'Jpogules arriveandproperconditionsarereestablishedforthedevelopmentofpredisturbancedominantspecies. Cyuress Autecology isagenusintheTaxodiaceae,a family intheConiferales.

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7Taxodiwndistichwnis commonly knoHn asbaldcypress,cypress,southerncypress,gulfcypress, tideHater redcypressand yelloHcypress(FoHells, 1965).Thenamebaldcypressisusedbecausethetreeisannuallydeciduous (Mattoon, 1915).T.distichwnrangesonthesoutheasterncoastalplainfrom toTexas,andintheMississippiRivervalleytoIndianaandIllinois(Mattoon,1915). T. ascendenshasa narrOHer range,fromVirginiatoLouisianaonthecoastalplain.Athirdspecies,T.mucronatwn,isfoundinnorthernMexico(Small,1931).Knownasthe"woodeternal",cypress has beenusedforoutsideconstructionsuchassiding,sashes,doors,blinds,steps,railings,porches,buckets,tanks,mouldingandshingles 1915).InFlorida,cypresshasbeenusedtomakecratestocarrysoftdrinksandoranges.Onecommonuseforpondcypressisfenceposts,althoughtheportionofthepost Hhich isalternately Het anddrydecaysfasterthantheportionswhichremain Het orareconstantlydry.TodayinsouthernFlorida,smallsawmillscutpondcypressintostakes,usedtosupporttomatoplants gro;m inthewinter.Taxodiwnismonoecious.ThefemaleconesappearinMarchandApril,andmatureinOctoberthroughDecember(Bonner,1974).Themaleconesarepresentfrom Decemberto andformCassle-likeclusters8to12emlong.Thefemaleconesare1.3to3.1emindiameterandcontain18to30seeds(Mattoon,1915).Twoovulesareassociatedwitheachscaleonthecone(Bonner,1974).Theconesusuallydisintegrateonthetree,butmayfallentiretothe Duetotheirsize,theseedsarenotdispersedby windbutby floHing(F01'lells, 1965).Theseedmustsoakfrom onetothreemonths 1916),nottobreakaphysiologicaldormancy,butsothat Hater canpermeatethethickseedcoat(Murphyand

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8Stanley,1975).Theseed Hill remainviable undenrater uptooneyear 1959),yet Hill notgerminate while underwater(Demaree,1932).Germination Hill occurondryground,butnotonwell-drainedsoils(Mattoon,1915).Percentgerminationvaries: (1959)reported 35%, Mattoon(1915)gavevaluesrangingfrom55to88%,Bonner (1974) measuredfrom 67to 93% (fullseedonly),and Hurphy andStanley(1975)reportedvaluesfrom 13 to 90% (fullseed).Thegerminatedseedlinghasfromfourtoeightcotyledons,averagingsix(Welch,1932).Water fortheseedlingseemtodeterminethedistributionofthespecies.Irreversiblewiltingoccursthreetofourhoursfollowinglossofsoilmoisture(DicksonandBroyer,1972). Growth onanaerobic,floodedsoilislessthan onaerobic,floodedsoil(DicksonandBroyer,1972).Theseedlingwillnotgrowifcompletelyinundated,yet;rillproducenewsproutsfollowingasix Heek inundation(Welch,1932).Sufficient growth duringthefirstdryseasonmustoccurfortheseedlingtoescapeinundation(Demaree,1932),asprolongedinundation will resultinmortality(Demaree,1932;Bull, 1949;Betts, 1960).Bonner (1974) reportedthatnoanimalsarebelievedtoeattheseedorresinousconematerial,but at GorkscreH SwampSanctuaryhavebeenobservedeatingtheseed.Cypressseed Has a componentofthedietofthe nOH extinctCarolina (Sprunt, 19.5'+). Rabbitsandnutriamayeatthe nSH shoots,branches,barkandrootsofseedlings(Fowells,1965).Felledtrees,25to 40 cmindiameter,willproducehealthysproutsfromthestump.Treesupto200yearsoldmaysprout when cut do,m, butthemoreadvancedtheageofthetree,thelessvigorousthe (DetHiler,1916).Treesthataregirdledandfelledusuallydonotsprout (Nattoon, 1915).

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9Thetaxonomicdistinctionbetween 1. distichumand 1. ascendensisquestionable.Sometaxonomists(Small, 1933; LongandLakela,1971)considerthemtobe twospecies.Otherauthors(WestandArnold, 1956; KurzandGodfrey,1962;Fowells,1965)considerpondcypresstobeavariety (1. distichumvar.nutans(Ait.) SHeet) ofbaldcypress.Differencesbet He enthetwo types includeleafshape,leafarrangementandorientationofyoungbranches. 1. distichumhaslinearleaves,flattenedalongthebranches,andyoungbrancheswhichspreadaway fromthemainaxisofthetrunk(LongandLakela,1971).T.ascendenshassubulateleaves,appressedtothestem,andyoungbrancheswhichascendorbecomeparalleltothemainaxisofthetrunk(LongandLakela,1971).Inareaswhereboth types ofcypressoccur,thecharacteristicsofleafshape,leafarrangementandbranchorientationmayintergrade,whichleadstodifficultyindistinguishingbetweenthem.ObjectivesSuccessionalcommunitiesresultingfromlumbering,burningandburning-folloHing-lumberingofTaxodium-dominatedstrandsarepresentwithinCorkscrew Sanctuary.Threequestionswereproposed:1).Isthereevidencethatrestorationofthepredisturbancecommunityisoccurring?2).Giventhatthecommunitywillagainresemblethepredisturbancesystem,cantheprocessofsecondarysuccessionbeaccelerated?3).Isthe vegetationlong-lived,constitutingadisclimax,oristhevegetationactivelychanging?Toanswerthefirstquestion,aninventoryofthevegetationofeachcommunity wasdone.Activelyregeneratingspecieswouldbepresentasseedlingsand/orsaplingsandseedsourcespresent may giveanindicationofwhichspeciesmaypotentiallydominatethecommunity.

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10SeedlingsandseedsofTaxodium wereplantedintothesuccessionalcommunitiestodetermineifpartsofthesuccessionalprocessmaybebypassed.Survivaloftheseedlingswouldindicatethatpropersiteconditionsexistfor andthatotherfactorssuchasseedsupply,seedviability,orgerminationrequirementsmaybelimitingregenerationofTaxodium.Vegetationmaps wereconstructedfromchronological sequences ofaerialphotographs.Fromthemaps,vegetationchangesweredeterminedtomeasureratesofchangefollowinglogging,burning,andlogging-and-burning.

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STUDYSITE CorkscrewSwampSanctuary,locatedinnorthernCollierCounty,Florida(Fig.1),isthelargestremnantofthecypressstrandecosystemswhichatonetimecollectivelymadeuppartsoftheBigCypressSwampasdescribedbyDavis(1943).TheSanctuarypresentlyconsistsofapproximately4200haownedandmaintainedbytheNationalAudubonSociety,primarilyforthepreservationofthecypressforestsandtheassociatedwoodstork,Mycteriaamericana,rookery.Beforepermanentdwellingswerebuiltin 1954, thesitewasusedasa camp,firstbyhuntersandlaterbygamewardensprotectingwaterbirdsfrom plumehunters.LoggingwasinitiatedtothesouthoftheSanctuarybytheLeeTidewaterCypressCompanyinthelate1940's.Loggingcontinueduntil 1954 whentheCorkscrewCypressRookeryAssociationwasformedtoprotectremaininghabitatsusedasrookeries.TheAssociationpurchased907hafor$170,000,andtheLeeTidewaterCypressCompanydonated259hatoformthesanctuary.TheNationalAudubonSocietyassumedsupervisionandmaintenanceoftheSanctuaryandacquired1295 morehectaresfromtheCollierEnterpriseoveratwelveyearperiod.Anadditional1748hectareswerepurchasedin1968asbufferareas.AfireenteredthesanctuaryfromthesoutheastonJune8,1962.Theeasternstrandofloggedbaldcypresswasburnedalongwithportionsoftheunloggedpondcypress.RainsextinguishedthefireonJune10,1962.Inthelate1960's,officialsoftheSanctuaryfearedthatdrainagefromanextensivecanalsystemassociatedwiththeGolden Gate11

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12MorfinCo.Palm Beach Co.Dade Co. BrowardCo.J --) U p Lake OkeechobeeII I>OkeechobeeCO'r--S_t_._L_U_Ci_e._c_o_...r Co. Hendry HighlondsCo. Carlier Co.o I r kilemztei"S Ij Io2040 rr----I De SclQCo.I r/ Co. GladesCo,t--\ t ; ,'<,3 r ..;.VMYEflSLea Co.\ fi'9{CORKSCR:::W\1fiA.DU::S SWAMP\ SANCTUARY \\.. IFigure1Locationof CorkscreSHamp Sanctuaryinsouthern l____ __F_l_o_r_i_cla.

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13development,located5kmtothesouth,couldlowerwaterlevelswithinthesanctuary.In1969construction was completedon adikedesignedtoimpoundwaterintheSanctuary.Twoexistingrailroad werejoinedacrossacentralmarshtocompletetheearthendike.Thedikeisapproximately2mhighand5m wideatthetop.MajorhabitatsandrecenthistoriesofthestudysiteswithintheSanctuaryareshowninFigure2.Experimentalsiteswereestablishedintheloggedareaofthewesternstrand,oneabove(northof)thedikeandonebelow(southof)thedike.Twositesweresetupinthelogged-andburnedareaoftheeasternstrand,oneestablishedabove(northof)thedikeandone below(southof)thedike.Onesitewasestablishedintheburnedpondcypressareaabovethedike.Figure 3 includesaphotographofthevegetation of eachofthefivestudysites.Asaninventorywas doneaspartoftheresearch,thevegetationofeachofthestudysitesisdescribedindetailintheResultssection.Taxodium dist"chum dominatesthebaldcypresszoneofthevirginstrandand I. ascendensdominatesthepondcypresszone.MarshhabitatsaredominatedbySagittariagraminea,Pontedariacordata,Cladiumjamaicensis,Panicum hemitomum,andSpartinabakerii.Pinuselliottiivar.densaandSeronoareDensprevailinthepinehabitat.Salixcarolinianacoverstheburnedarea.Acerrubrum,Ilex Perseapalustris,Myrsineguianensis,MyricaceriferaandSalix arepresentintheloggedcypressarea.

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14o.sPinePordcypress Kilometers 0. l'3Logged duringearly1950'5 E3 a'Hoedin1962 Seuth dixe, construction completed in1969STUDY SITES' LA -Logged,notburned,cbovefhe dike LBLogged,notburned, below the dike LBA -Loggedandburned,above thedike LB 8 -Loggad ond burned, belowfhe dike SA -No! logged,butburned, obav the dike Figure2.HabitattypesandrecenthistoryofstudysiteswithinCorkscrewSwampSanctuary.SiteBAwasformerlydominatedbypondcypress;theotherswereformerlydominatedbybaldcypress.

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Figure;.PhotographsofstudysitesatCorkscrewSwampSanctuarytakeninDecember, 1976. (A) Logged,above-the-dikestudysite.Noticefelledcypresstree,notremovedbecausehollow.(B) Logged,below-the-dikestudysite.Mixed hardwoods. (C) Logged-and-burned,below-the-dikestudysite.DensethicketofSalixcaroliniana.(D) Logged-and-burned,above-the-dikestudysite.ClumpsofSalixcarolinianagrowing onburned-outstumpsinterspersedamongdeeperwaterareas.(E) Burned,butnotlogged,above-the-dikestudysite.Borrowpitand vegetationareintheforeground.Thedarkvegetationisa bermcoveredwithMyricacerifera.ThegraybackgroundisastandofSalixcarolinianaand emergentsnagsareamixtureofliveand dead Taxodium.

PAGE 24

16 '-":.,

PAGE 25

NETHOD3 Thisstudyincludedadescriptionofthevegetationpresentatthefiveexperimentalsites,plantingof seedandseedlings,monitoringof ,ater levelsateachstudysite,andanexaminationof setsofaerial photogrc.pr.s. VegetationDescriptionTodetermineifrestorationofapredisturbancesystemisoccurring,a vegetationdescriptionofeachofthefiveexperimentalsites ,as done.Threecomponents were considered:trees,shrubs,andunderstoryplants.Treeswereconsideredtobeallplants Hith diametersatbreastheight (DBn =1.J7maboveground)greaterthanJ.8cm.Shrubsweredesignatedasall Hoody plantslessthanJ.8 CIL DBHandgreaterthan1mtall.Allvascularplantslessthan1mtallandall non-,roody plantsgreaterthan1mtallwereconsideredtobepartoftheunderstory.TreesWeremeasuredina 25 D x 25 mplot,andshrubswere measlITed ina10mx10mplotnestedwithinthetreeplot(Fig.4).AIm ,ade pathWaSclearedaroundthe25 mx 25 mplot,andeachplotwasdelineatedbystringstretchedfromcornerstakes. For eachspeciesineachofthethreecategories,ameasureof domin3.nce, density,and "as made.Thesemeasureswereconvertedintorelativevalues(%)and toyieldan import3.nce value(Curtis,1959)foreachspecies.MaximumvalueforasinglespeciesisJOO.17

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18 [Jseedling plot E2 bold Cypress seed plot E:Sseed plot50m TR.l,NSECTFOR SEED AND SEEDLINGPLOTSI -.;fJ-'+50 mTRANSECTFORVEGETATION INVENTORY --UC-Samplingareaior understory vegetalion ,I 25m10m f 25mTree inventory plot Shrub plot 1+--i 0m ----to; I L------,-'----.J'i o \'jELL SITE10 Plots among existing vegetation mzters-Tl o5 Cleared areosFigure4.Schematicmapofsamplingprocedureateachstudysite.Arrangementoftree,shrub,andunderstoryinventoryplots,baldcypressandpondcypressseedplots,seedlingtransplantplotsandwellsiteisshown.

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19Dominance Hasexpressedforthetreeandshrubcategoriesbymeasuringdiametersandcalculatingbasalareas.Treediameters(DBH)weremeasured 'lith asteeldiametertapetothenearest0.1em.ShrubdiametersHeremeasuredtothenearest0.013emataheightof30emusingverniercalipers.Relativedominance wascalculatedforeachspeciesas:basalareaofaspeciesx100totalbasalareainplotDensityofeachspecieswasobtainedbycountingthenumberofindividualsofeachspecieswithintheplot.Relativedensity Has calculatedas:numberofindividualsofaspeciesx100totalnumberofindividualsinplot oftreespecies Has measuredasthe oftimesthespeciesoccurredintwenty-five,2mx2mplots.Theplotswereselected by randomlyselectingcoordinatesOnagridsystemestablishedinthetreeplot. ofshrubspecies talliedasoccurrenceintwenty-five1mx1mplotsnestedHithinthe2mx2mtreeplots.Relative wascalculatedas: nQmber ofoccurrencesofaspeciesin25plotsx100totaloccurrenceofallspeciesin25plotsImportancevalueswerecalculateddifferentlyforunderstoryspecies.Thecompositionoftheunderstoryvariesseasonally,sosixharvestsweremade,attHo-monthintervals,startinginNovember,1975andendinginSeptenoer,1976.Ateachharvest,three1mx1mplots Here selectedat random alongthe50mnorth-southtransectateachstUdysite(Fig.4).Theplots Here clippedofallabovegroundunderstorybiomass.Theclip-pings Here driedat70Cfor48hoursanddryHeightHasrecorded,by

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20species,tothenearest0.1gram.Relativedominance wasexpressedas:dryweightofaspeciesx100totaldry Height Stemswerecountedasameasure of densityandrelativedensitycalculated:numberofstemsofaspecies x100Frequencywascalculatedas orabsenceinthethreeharvestquadrats,andconvertedtorelativefrequencyasfollows:frequencyof inthreeplots100totalfrequencyper datexRelativedominance,relativedensityandrelativefrequencyweresummedtoyieldanimportancevaluefor each speciesoneachharvestdate. Seedlings Totestthehypothesisthatsecondarysuccessioncanbeacceleratedbyintroducingthedominantpredisturbancespeciesintopostdisturbanceseres,cypressseedlingswere intothefiveexperimentalsites.ThecypressseedlingsweretransplantedduringJanuary,1976.Theseedlingswerecollectedfromtheperipheryoftheundisturbedcypressstrand,wherethetreesexhibitpondcypresscharacteristics.Thecri-teriaforselectionwasanycypresstreebetween0.5mand1mtall,havingabasaldiameterlessthan12mm.Theseedlingswereallfoundwithin0.5kmofeachother.Removal wasaccomplishedbybreakingthegroundaroundtheseedlingwithashovel,graspingfirmlyatthebaseofthestemandgentlyextracting. The rootswere washedofremainingsoilandplacedinplasticbagswith '8ter andbladderwort(Utriculariasp.).Plantingwas donewithinonehourofremovalbyinsertingaflat15cmx40cmshovelintotheground, a:,d rockingitbackandforthtocreate

PAGE 29

21aV-shapedhole.Thebarerootseedlingwasinsertedandtheholefilledbypackingwithsurrounding soH. Atotalof120seedlingsweretransplantedintothefivesites.Six1mx1mplotsweresetupateachsite,withfourseedlingsperplot,oneseedlingplantedneareachcorneroftheplot.Eachseedlingwas marked by anumber-letter-numbercode Hritten onanaluminumtagattachedtothestakenearesttheseedling.Threeplots Here established among theexistingvegetation,andthree Here centeredin5mx5mclearedareas(Fig.4).Allplots Here atrandomlyselecteddistancesalonga50mtransectparalleltothetransectestablishedinthevegetationsurvey(Fig.4).As atransplant-shockcontrol,threeplots Here establishedintheareafrom whichtheseedlingswere removed. Theheightofeachseedlingwasmeasuredtothenearest12mmwhentransplantedandagainonFebruary29,March22,April17,May14,June11,JUly17,andOctober 23, 1976.Theseedlings Here observedforpresenceorabsenceofleavesoneachdate.Theleafshapeandarrangementandbrancharrangement Here observed folloHing transplantingandonthefinalmeasurementdatetodeterminetheeffectsofsiteonleafmorphology.SeedsTodistinguishwhichstageofthelifecycleofcypressmaybelimitingsurvival,seedswerealsoplantedateachexperimentalsite.MurphyandStanley(1975)statedthatcypressseedsgerminate folloHing a 50to60daysoakingperiodduring Hhich thethickseedcoatbecomespermeable.Demaree (1932) listedrequirementsforcypresssurvivaltoincludegerminationondrygroundandsufficient groHth toescapeinundation.Seedplotswereestablishedadjacenttotheseedlingplotstodetermine Hhether theseconditionsexistinthestudysites.

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22OctoberthroughNovemberthefemaleconesareabundantontrees (f1attoon, 1916).LateinNovemberconeswereretrievedbyshakingthebranchesuntiltheconesfellorbyharvestingwithascreennet-likebagattachedtoa 3 mpole.Becauseofthequestionastoexactdifferencesbetweenthetwotypesofcypress(pondandbald),seedswerecollectedfromtreeswhichexhibitedpondcypressattributesandfromtreeswithbaldcypresscharacteristics.Theconeswereair-driedinthree1.5mx0.6mscreen-bottomedflatsforfiveweeks.OnJanuary4,1976bothbaldcypressandpondcypressseedsplusconedebrisweredividedintofiveequalportions.Eachportionwasputintoafiberglassscreenbagand SOlin shut.Onebagofpondcypressseedandonebagofbaldcypressseedwerenailedtoapostsothatanimalscouldnotremove themandputindeep-waterareasforsoakingateachexperimentalsite.Asthewaterwentbelow 5ound ateachsite,theseedplusconedebriswere weighedoutin250 gand125gsamplestoyieldapproximately250pondcypressseedsand100baldcypressseedsrespectively.Theseedswereplantedin1emdeepfurrows.DatesofplantingwereFeb. 13 (burned,above-thedike),February27(logged-and-burned,below-the-dike),March16(logged,below-the-dike)andMarch 23 (logged-and-burned,above-the-dikeandlogged,above-the-dike).Onthedayofplanting,200pondcypressseedsand200baldcypressseedswereseparatedfrom conedebris.Four50-seedreplicateswereplantedinorganicsoilinwoodenflatsandplacedundergreenhouseconditions.Daily germin2tion wastalliedasappearanceofcotyledonsabovetheground.Theresultingseedlingsweremonitoredfordifferencesinleafmorphology.Anothergroupofconeswascollectedon December29,1976todeterminepotentialgerminationunderlaboratoryconditions.Theconeswere

PAGE 31

23air-driedandtheseedseparatedfromdebris.Four100-seedlotsofeachtype(pondcypressandbaldcypress),weresoakedforfourhoursin 95% sulfuricacid(specificgravity asrecommended by Murphyandstanley(1975),thenrinsedovernightintapwater.Twentyseedswereplacedonmoistfilter paper ineachoffiveplasticpetridishesfor each 100seedsample.Thedisheswereplacedinanincubatorseton acycleofexposuretolightfor8hoursat30 Canddarknessfor16hoursat20 C.Germinationasevidencedbytheappearanceofcotyledons monitoreddaily. Hater Levels levelsweremeasuredweeklyatonewellineachofthefiveexperimentalsites.Thedistancesfromthetopofa5.1cmdiametersteelpipetothewaterlevelinsideandoutsidethepipeweremeasuredtothenearest6.25mm.Insidemeasurementswere made bydroppingafloat,attachedtoafishingreel,intotheHell.Theslack ,las removed fromthelineandthedistance Has measuredfrom a mark onthefloattothewelltop.ThetopsoftheHellsweresurveyedbyM.J.DueverandL.A.Riopelleandcalibratedtomeansealevelheights.TheHaterlevelmeasurementsatdifferentsitesHeremadeHithinonehourofeachothertominimizedifferencesduetodailyoscillationscausedbyevapotranspiration.AerialPhotographsVegetationmaps Here made fromthreesetsofaerialphotographs.Thefirstset(1:20,000)wastakeninJanuary,1953,beforethestudyarea loggedorburned.Thisset(FlightCodeDSM-4L,#'s21,22,23,55,56,57) Has purchasedfromtheUSDASoilConservationService,CartographicDivision,Hyattsville,Maryland.Thesecondgroup(1:20,000) was takeninNovemberandDecember,1963,nineyearsfollowingloggingandeight

PAGE 32

24monthsfollowingtheburn.These photographs (FlightCodeDSM-JDD,#'s178,179,180and DSM-4DD, #'sJO,Jl,J2)wereborrowedfromtheSoilConservationServiceOfficeinGaines-rille,Florida.The mostrecentphotographs(1:75,000)weretakeninDecember, 1972 by Mark HurdAerialSurveys,Inc.(FlightCode IRC-CUJ,#'s1594,1595,1102).Allphotoswere22.8cmx22.8cm(9"x9") blac:< andwhitecontactprints.

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RESULTSVegetationInventory The southern coastal willow,Salixcaroliniana,hadthehighest valueinboththetreeandshrubplotsattheburned,abovethe-dikesite(Table1). The highdensityofstemsandhigh frequency of occucrence ofSalixresultedinhighimportancevaluesintreeplot#1, therelativelylargerdiametersofTaxodiumand Sabal accountforthe INier importancevalueofSalixintreeplot#2.Myrtle, Nyrica cerifera, secondinimportance value intreeplot#1andthirdintreeplot#2.Thisisduetohigherrelative frequency, andahigherdensitythan Taxodilli", SabalorPersea.Taxodiuminthetreeplotswereeithersurvivorsfromthefireornaturallyregeneratedfromseed.Persea,CephalanthusandSabal Here encounteredinthetreeplotasscatteredindividuals.Saplings(treespeciestallerthan1 m andless 3.8emDBH)of Nyrica, Taxodium, Annona,andCephalanthusinthe sbxub plotsindicatethatregenerationofthesespeciesisoccurring.Compositionandimportancevalues Here similarattheplotsaboveand beloo thedikeinthelogged-and-burnedarea(Table2).Only twospecieswereencounteredinthetreeplotsaboveandbelowthedike,whereSalixformsalmostapurestrand. MyricaHas slightlymoreimportantintheplot belo" thedikethanintheplotabovethedike, probably becausethereismoredry g-.coundbelOl'I thedike.NoTaxodium was encounteredinthetreeplotsaboveand belo;" thedikeintheareathat was logged-and-25

PAGE 34

26Table1.Importancevaluesforoverstoryspeciesattheburned,above-the-dikestudysite.Relative+Relative+RelativeImportanceDensityDominanceFrequencyvalueTreePlot#1Salixcaroliniana9187 81 259Myricacerifera681428Perseapalustris11 1 3Taxodiumascendens2 3 4 9ShrubPlot#1Salixcaroliniana83 8871242Myricacerifera1391236Cephalanthusoccidentalis 3 2 4 9Baccharishalmifolia11 4 4Annonaglabra11 1 1TreePlot#2Salix 72 3882192 Hvrica cerifera15141241Taxodiumascendens1126 643Cephalanthusoccidentalis11 1 1 SabaJ,. palmetto1211 22ShrubPlot#2 9alix caroliniana939679268l1yricacerifera 3 2 813Taxodiumascendens1 1 810Baccharishalmifolia11 4 5 LUdwigia peruviana1 1 1 1Cephalanthusoccidentalis1 1 1 1

PAGE 35

27Table2.Importancevaluesforoverstoryspeciesatthelogged-and-burned,above-the-dikeandbelow-the-dikestudysites.Relative+Relative+Relative=ImportanceDensityDominanceFrequencyvalueTreePlotAbove-the-DikeSalixcarolin;ana Nyrica ceriferaShrubPlotAbove-the-DikeSalixcarolinianaCephalanthusoccidentalisBaccharishalmifolia Nyrica ceriferaLudwigiaperuvianaAnnonaglabraTreePlotBelow-the-DikeSalixcaroliniana Nyrica ceriferaShrubPlotBelow-the-DikeSalixcarolinianaBaccharishalmifoliaCephalanthusoccidentalis Nyrica ceriferaLUdwigiaperuviana97 ) 50 )) 9 4 ) 1964622014 ) 19917517741196484592 199171i)271194 661179 9429551966)18155 ) 2861420742)214 6

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28 burnei. Salix,Cephalanthus aniHyricaHere inbothshrubplotsaboveand belo;; thedikeinthelogged-and-burnedarea. ll-ard;;oods suchasdahoonholly,Ilexcassine,redmaple,Acer moTtle, Myricacerifera, bay,Perseapalustris,popash, caroliniana,andpond apple, Annonaglabra Here conspicuousinthe tree plotatthe studysite,above-the-dike(TableJ). inthetreeplotare fromloggingandsproutsfromstumps. CeDh2.1anthusHaS thedominantspeciesfoundintheshrubplot.Saplings Here presentofthefolloHing r-ard.Hoods: Myrsine,Myrsineguianensis, Myrica, Ilex,PerseaandAcer. No individualsofTaxodium Here encounteredin plot,butasapling noticedinthetreeplot. Hard-,fOod speciesalsodominatethetreeplotatthelogged-notburned,beloH-the-dikestudysite(Table4).ImportancevaluesofAcerand :=lex-;;ere thehighestin tl:e-treeplot, wheretenspecies l-lere encountered.Notreesof Taxodima;;ere observedinthetreeplot.Impor t,wce valuesofAcer,Annona,Fraxinusand !1y:rica Here higheratthe belo;;-the-dike plotthantheabove-the-dikeplot, Hhile Salixand Here moreimportantintheabove-the-dikeplotthaninthe belo;:-the-dike treeplot. CeplnlanthusHas thedominantspeciesinthe shIue plot,as Has thecasein the above-the-dikesite.SaplingsofMyrsine,Persea,AcerandIlex He:",e foundintheshrubplotatthe beloHthe-d.ike site.Importancevaluesaregivenforeachunderstoryspeciesoneach harvestdate foreachstudysite in Tables5-9.Thevaluesforeachspecies >;ere summedoverthesixharvestdatesand raQked ineachtablewiththelargestcumulativevalueatthetop,decreasingtOHardthebot tee,. fern,Blechnum se:",,ula tum, thedominantunderstory

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29Table 3. Importancevaluesforoverstoryspeciesatthelogged,above-the-dikestudysite.Relative+Relative+RelativeImportanceDensityDominanceFrequencyvalueTreePlotIlexcassine 31 192777Acerrubrurn12251148Salixcaroliniana17 13 1141Myricacerifera15 911 35 Taxodiumdistichum 3 244 31 Cephalanthusoccidentalis1241127Perseapalustris4 3 1522Fraxinuscaroliniana214 6Ficusaurea1 1 3 5Annona glabra 211 4ShrubPlot CephAlanthus occidentalis20294594Myrsineguianensis 36 201167Myricacerifera222014 56 Ilexcassine5 81427Iteavirginica510419Persea pa1ust-ris 411116Acer rub:twn3 4411Salixcaroliniana2 5 1 8Baccharishalmifolia 3 11 5

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30Table4.Importancevaluesforoverstoryspeciesatthelogged, beloi,-the-.iike studysite.Relative+Relative+Relative ImportanceDensityDominanceFrequencyvalueTreePlotAcerrubrwn28381581Ilexcassine19223374Myricacerifera1462242Perseapalustris127 726Annona glabraJ 16423Fraxinuscaroliniana104 721Cephalanthusoccidentalis7 2 4 13Salixcaroliniana J 24 9Ficusaurea124 7Myrsineguianensis3 11 5ShrubPlotCephalanthusoccidentalis538015148Baccharishalmifolia184426Myrsineguiancnsis7 3 1525Iteavirginica9 21122Perseapalustris4 3 411Acerrubrwn6 3 110 l1yrica cerifera11 79Ilexcassine1 2 4 7

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31speciesatallsites.SeedlingsofSalix, Hyrica andCephalanthus Here theonlytreespeciesintheclippedplotsattheburned,above-the-dikestudysite(Table5).No Taxotiium Has harvestedfromanyofthesamplequadratsatthissite.Seedlingsof HyricaHere foundintheNovemberand Harch harvestsatthelogged-and-burned,above-the-dikesite(Table6).SeedlingsofSalixand HyricaHere harvestedinJanuaryand Hay, respectively,atthelogged-and-burned,below-the-dikesite(Table7).THenty-twospecieswereharvested beloH thedike,andthirteen Here harvestedabove. r;o cypress Has foundinplotseitheraboveor beloH thedikeattheloggedand-burnedsites.SeedlingsofAcer Here abundantintheharvestsfromthelogged,beloH-the-dikesitefrom November,1975throughJuly,1976(Table8).Myrsineseedlings Here foundintheNovemberharvest.IlexseedlingswerefoundintheJanuaryharvestbelowthedike,andAcerseedlingswere componentsoftheMarch,MayandJulyharvestsatthelogged,abovethe-dikesite(Table9).SeedlingsofAnnona Here encounteredduringtheMarchand harvestsabovethedike.Myrsine "as sampledinMayandCephalanthusseedlings Here sampledintheMarchharvestsabovethedike.NoseedlingsofTaxodium wereclippedateitheroftheloggedsites.Highbiomass values occurredbetweenlatespringandlatefallatallthestudysites(Table10).Maximumbiomass >laS measuredduringtheJuly attheloggedsitesandtheburnedsite(Table10). Maximl biomassoccurredabovethedikeatthelogged-and-burnedsiteinMarch. Thelogged-and-burned,below-the-dikesiteyieldedhighbiomassharvestsinNovember,JUlyandSeptember,1976.

PAGE 40

Table5.Importancevaluesofspeciesineachharvestofunderstoryplantsattheburned,above-the-dikestudysite.SpeciesDate(month-day-year)Mean11-16-75 1-22-76 3-4-76 5-13-76 7-14-76 9-24-76Blechnumserrulatum0 122 35 861527478.2 Panicum hemitomon61.0 0 652067 35.5 Pontedariacordata86 31 56280 0 33.5 Diodiavirginiana212846 015 3323.8 palustris2119 45 8 050 23.8 Sarcostemmaclausa0 0 1+ 51 32 921.0 'i1ood'o'larclia virginica13 1503150018.2Pol yr;onumpuncta:tuffi 2011'+510 0 1516.8Andropogon 1+ 0 0 0 0 10 7.3 Mikaniabatafolia0 16 16 0 0 86.7Sagittariagraminea0 30 80 0 0 6.3LuclHiOjia repens0200 0 0 0 3.3 Baccharishalmifolia6 0 0 0 7 02.2Typhalatifolia 13 0 0 0 0 02.2Sagittarialancifolia 13 0 0 0 0 02.2Bacopa maritima0 0 12 0 0 02.0Astercarolinianus0 0 0 10 0 01.7 OSffiL:ndarepolia 0 0 0 0 9 01.5Cyperus 0 9 0 0 0 01.5Cephalanthusoccidentalis0 0 0 0 0 8 1.3EUE,torium compositifolium0 0 0 7 0 01.2Juncu3polycephaluc6 0 0 0 0 01.0 Nyrica cerifera6 0 0 0 0 01.0Salixcaroliniana0 0 0 0 6 01.0 cylindrica00 0 0 5 00.8 VJN

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Table6.Importancevaluesofspeciesineachharvestofunderstoryplantsatthelogged-and-burned,above-the-dikestudy sHe. SpeciesDate (month-clay-year)Mean11-16-75 1-22-76 3-4-765-1 3-76 7-14-76 9-24-76Blechnumserrulatum1458017900 1369.5 POlygonum punctatum0127202893955.9Mikaniabatafolia17482312013110 53.2Boehmeriacylindrica J4 0 1027 J4 3723.7Peltandravirginica1600007314.8Astercarolinianus02412220 911,2Cyperus E. 0200280 1811.0Sarcostemmaclausa0 01045 009.2Myricacerifera400150009.2Woodwardiavirginica5100 0008.5Osmunclaregalia0 000 42 88.3Typhalatifolia0 011300 98.3Azollacaroliniana00 15 0002.5 \.,.) \.,.)

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, !!I Table7.Importancevaluesofspeciesineachharvestofunderstoryplantsatthelogged-and-burned,below-the-dikestudysite. ;I SpeciesDate(month-day-year)MeanI11-16-75 1-22-76 3-4-76 5-13-76 7-14-769-24-76Blechnumserrulatum1032898718311482.8Baccharishalmifolia2373 J6 15571135.8 lancifolia21761725 J4 1230.8Pontedariacordataa53a6829a25.0Ncphrolepis exalta 48a519 7a19.2Spirodela aaa a a10717.8Polygonum punctatum72097221413.2Cyperus ll.. aaa52343 11.8 cylindrica77172115a 11,2 LUdwigiarepensa18101225a10.8Mikaniabatafolia7 8 97 8118.3Thelyptrisnormalis18a a9a187.5Astercarolinianus7a1815aa6.7Osmundaregalia23aa6aa4.8Andropogon ll.. 14a10a a a4.0Peltandravirginica158aa aa3.8Hydrocotleumbellataa7a14aa3Woodwardiavirginica7a13a aa2.2Typhalatifoliaaa12a aa2.0 ]'lyrica ceriferaaaa10aa 1,7 Salixcarolinianaa7aa aa 1,2 quinquefoliaaaa7aa 1,2'i-

PAGE 43

Table8.Importancevaluesofspeciesineachharvestofunderstoryplantsatthelogged,below-the-dikestudysite.SpeciesDate(month-day-year)Mean11-16-75 1-22-76 3-4-765-1 3-76 7-14-76 9-24-76Blechnumserrulatum5972 0 75306149.5Osmundaregalia44 8 107789640,5Nephrolepisregalia0317923102 039.2Acer rubrum52267129 270 34.2 Sagittarialancifolia005312 455227,0 Woodwardiavirginica 3616311523020.2Boehmeriacylindrica243824119 1219.7Myrsineguianensis00 141+302513,7Polyrjonum punctatum0290 8 0 168,8Peltandra virf5inica 220 0 0220 7.3 Thelyptrisnormalis238 0 00 137.3Iteavirginica0210 0 0 95.0Myricacerifera00 9 17 0 04.3Diodiavirginiana0 10 0 0 10 0 3,3 Melothriapendula00 0 0 18 03,0Pontedariacordata0 17 0 0 0 02,8 BaccDAris halmifolia0 15 0 0 0 02Astercarolinianus0 0 00 0 152Mikaniabatafolia14 0 0 0 002Persea 13 0 0 00 02.2Smilaxlaurifolia12 0 0 0 0 02.0Panicumsp.00 9 00 0 1,5 Ilexcassine06 0 00 0 1,0

PAGE 44

TO-bIc 9. Importancovaluos ofspociesinoach rorvcst ofundorstoryplantsat thelo[!;[!;ed, abovo-tho-dikestudysito. C' Do-to(month-dO-y-year)Moan0peCles 11-16-75 1-22-76 34 -76 5-13-76 7-14-769-24-76Blechnumserrulatum111186 53 57527889.5Spirode1apolyrhizae00 0 00 170 28.3 vioodwa-rdia virginica460 355413 0 24.7Boohmeriaoylindrica4022501/+18a24.0Thelyptrisnormalisa0 0 0 752116.0Sagittarialancifolia021500240 15.8 Nophrolopi::;oxalta 1710 00550 13.7 Crinumamericanum00065aa10.8Osmunda regalia 290 15 11+0 0 9.7 Hikania batafolia0 102119 7 09.5Panicumsp.3212a a8a8.7Astercarolinianus8 13 209 0a8.3Acer rubrum00 14 15 15 07.3Annonaglabraa0 17 17 005Melothriapendula8000240 5 Pistiastratioides000 00305.0Myrsineguianensisaaa26a0 4.3 Cephalanthusoccidentalisa026aaa 4.3 Polygonum punctatum0 10 0 0 7 02.8Campyloneuroncostatum00010001.7Habenariasp.8aa aa0 1.3\...)'"

PAGE 45

StudySiteTable 10.Abovegroundunderstorybiomassduringthestudy period. Eachvalueisameanofthree1m 2 samples ( :: onestd.deviation).Valuesareovendryg/m2 Date (month-day-year) 11-16-751-22-763-4-76 5-13-76 7-14-76 9-24-76 Burned,above-the-dikeLogged-and-burned,above-the-dikeLogged-and-burned,below-the-dikeLogged,below-the-dikeLogged,above-the-dike 51::!:. 36.061 ::!:. 59.7111 ::65.324:: 28.551 :: 46.5 35::!:.16.7 24+14.339 13.0 27::!:. 4.161 25.9 24. 8.1 104. 30.7 320' 96.0 329. 470.480 ':::'109.46:: 3.935+38.251+77.497+ 34.2 ---38+27.380+75.7132+98.1--46:: 22.9 44:: 33.9 126.:::. 64.291 :!: 32.1 104 :!: 127.0103 ::!:. 21. 0 89 :!: 36.841 :: 24.8 '=:j

PAGE 46

WaterLevelsThegroundattheburned,above-the-dikewell(Fig.5) the high estinelevationofallstudysites,at5.10mabovemeansealevel(MSL). Thesurfaceofthewellwasdryfor180days,fromFebruary16toAugust14.Thewaterlevelsinsidethepiperangedfrom a lowof4.03maboveMSL(-1.07m belowthesoilsurface)onMay12toahighof5.43maboveMSL(+0.33mabovethesoilsurface)onOctober2,1976.Theelevationoftheseedandseedlingplotsrangedfrom5.10mto5.13maboveMSL.Thegroundsurfaceatthelogged-and-burned,above-the-dikewell was dryfor107days,from !1arch 11toJune27(Fig.6).Lowestwater measuredonMay12at4.14maboveMSL(-0.78m belowthesoilsurface).Waterlevelsreachod5.35m(+0.43mabovethesoilsurface)on November2,1975andOctober2,1976.Theelevationsoftheseedlingplotsrangedfrom4.61to5.17maboveMSL.The water levelsinsidethepipeatthelogged-and-burned,belowthe-dikesitewereconsistentlyhigherthantheleveloutsidethepipe(Fig.7),indicatingthatthedike was loweringsurfacewatertothesouth.However,thesitebelowthedike was dryfor108days,approximatelyequaltothedryperiodabovethedike.Thus,thedikeresultsinhigher water onitsupstreamside,butdoesnotseemtoalterthedurationofthetimeperiodwhenthesoilisflooded.Thelowestwaterlevel was measuredonMay18at4.38maboveMSL (-0.54 m belowthesoilsurface)Maximumwaterlevelwas5.19maboveMSL(+0.27mabovethesoilsurface)beforewaterretainingboardsintheculvertsthroughthedikewere removedinJune,1976.Levelsreached5.38maboveMSL(+0.46mabovethesoilsurface)followingremovalofthewaterretainingboards.Theelevationsoftheseedlingplotsbelowthedikedidnotvaryasmuchasthoseofthe

PAGE 47

:;'"'",UI osAE-amongexisting vegetationJ J A MFJD Nr'1 1"" < :b -Io Z oI ","'UO'-...".'>. Levelof soil surfaceat well...... II '1"" I'CI,C2,C3 fll orzGJ-'0 ro -1(I)4.0 I'!fI 1t \ -J \II!6.0rI I OJQ) E 55[-1k:J-"W-1 I (,) ((l
PAGE 48

o"1(f)MfYl orZ Gl tJro -IL13'"roM r M::I oz F.:3Cc. C3C!C-in clrored areasE-among existing vegetation outside -) 4.55.0 l.evel of 50:1 surfaceat well",""'-:::: II [I 6,0rIII5.5" f:r: (!)UJ:r:-":;:zW;;,;:lei> o :n<'t':.1iiIE-lLuGj_I<:J:W'J) 4.0I' ,I I!I!r!II j-J DJF MAMJJAs o I-1975I1976-------------Figure6.Waterlevelsatthelogged-and-burned,above-the-dikestudysite. .t: o

PAGE 49

", r ", < l>:::! 0 z 0 E3,Ct (flrn EI rn. E2,C3 IC2_ Z(;)"1l5-ju)3o. ..'" Eamong existing vegetation Cin cleared arcas.:!!Q)E ...l 5.5 w ;... w...l
PAGE 50

42plotsabovethedike,rangingfrom 1[.92 mto5.04maboveMSL.Atthelogged,butnot-burned,above-the-dikestudysite(Fig.8)the 10HestHater level Has measuredon May 15at4.66maboveMSL(-0.33mbelowthesoilsurface).WaterlevelspeakedonOctober2at5.53mabove MSL(+O.54m abovethesoilsurface).Elevationsoftheseedlingplotsrangedfrom4.80to5.01maboveMSL,resultingin dry periodsranging fro", 38to103days.WaterlevelsbeloH-the-dikeattheloggedstudysitewerelowerthanlevelsabovethedike,evenafterremovalofthewaterretainingboards.Inside-the-wellmeasurementswerehigherthanoutside-the-wellmeasurements beloH thedikeintheloggedarea(Fig.9).Nowater visibleonthegroundoutsidethewellfor114days,fromApril1toJuly17.Low Hater measuredat4.47m(MSL)(-0.40m beloH thesoilsurface)on 18.Highwater measuredatthe Hell onOctober2at5.25maboveMSL(+0.38mabovethesoilsurface).Elevationsoftheseedlingplotsdifferedby0.18m(4.83to5.01illabove MSL) andweredryforperiodsoffrom100to155days.SeedlingsCompensation madeforthevariationininitialsizeofthetransplantedseedlingsbyexpressingheightincreasesaspercents.Percentincreases Herestmuned foreachplot,thendividedbythenumberofsurvivingseedlingstoyielda meanpercentheightincreaseperplot.Assumptionsofananalysisofvarianceincludehomogeneityofvariancesandthatthedata folloH a normaldistribution.Achi-squaregoodnessoffittest(SnedacorandCochran,1967)wasusedtodeterminethatthepercentheightincreasesfolloweda normaldistribution.Bartlett'stestforhomogeneityofvariances(SokalandRohlf,1969)

PAGE 51

3 ,"'"'"!'1rM o z o .,U1!'1rntJ r zGl"1J ro uJ E2,E3[I tr"C3 -.C2o s AE among existing vegetction JJM ;\ Level of soil at wellMFJDN I I I 4,01. ,,..,,,, -'" 're", ,4.5 50F"C'In I 'C I6.0 r 15.5 ...JlcJ >W ...J oQJf-:I:,nGj :.c '".c'CjE1--1975 I 1976IFigure8.Waterlevelsatthelogged,but-nat-burned,above-the-dikestudysite. -t:\.,)

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:3,n-noV>"5 -I(f) C1 r n1 < po;j o z[J E-amongeXistingvegetationC-incleared areaso (f) C1 I .n, Level ofSOIlsurface at wei!)Lz==:::=j E2 i2 } [1,CI,C2 .C3 Z..G'I 6.0r, I '"Q'00E 5.5 ..Jt.cJ > !.U..J0 CD
PAGE 53

indicatedthatthevarianceswere homogeneous. A analysisofvariance (SokRl andRohlf,1969)detectednosignificantdifferenceingrowthamongstudysites,orbetweengrowthofseedlingsinclearedareasandamongexistingvegetation.Theseedlingsgrowinginclearedareasexhibitedagreateraverage grolrth atallsitesthanseedlingsgrowinginunclearedareas(Fig.10).A one-wayanalysisofvariance(SnedacorandCochran,1967) usedtocomparetreatmentsateachsite.Astatisticaldifference(p=O.OS)wasrevealedbetweentreatmentsatthelogged-and-burnedsitebelowthedike,butnodifferencesweredetectedattheothersitesduetolargevariances.Bytheendofthestudy,therewasgreateraveragemortalityofseedlingsgrowingamongexistingvegetationatallsitesexceptthelogged-and-burned,above-tho-dikesite(Fig.11).Becauseofvariability, hOHever, a two-Hay analysisofvarianceyieldednosignificantdifferenceinmortalityeitherbetHeenclearedandunclearedareas,noramongsites.The numberofseedlingswithleavesincreasedatallofthesitesbyMarchastheseedlingsleafedout(Fig.12).Theseedlingstransplantedintotheclearedareassproutedleavesearlierthantheseedlingsgrowingamongexistingvegetation.The numberofliveseedlingsremainedroughlyconstantfromAprilthroughJuly,whenwaterlevelsrose.BetweenJUlyandOctoberseedlingsdiedfollowinginundationattheabove-the-dikesitesintheloggedandlogged-and-burnedareas.Overallseedlingsurvivalwas66%attheexperimentalsites,yetonly 58% survivalwastalliedatthecontroltransplantsiteestablishedintheareafrom whichtheseedlingswere removed. Theseedlingsdiedatthetransplant-shockcontrolsitebecauseoflossofrootcontactwithsoilmoisture,notinundation,as lO'derHater levelsandlongerdryseasonwereobservedatthis

PAGE 54

10 Burned,ebove-dike Logged -ondburnedI oi:rove.dii<8 Logged-end-LeggedI burn3d I abovebelow-dikeLeaced-" below!'Ji:'{e Figure10.Heightincreasesoftransplantedseedlingsatthestudysites.Valuesaremeans;barsrepresent+onestandarddeviation.

PAGE 55

D oAmongexistingvegetation Clearedareas12 :> 6 cr:::l
PAGE 56

48 ClearedareasoAmong existing vegetationLogged,abovethe dike Lagged,belowthe dikeLogged and turned, abovethe dike Logged andburned, below thedike 12 en 1w> z(; Burned,abovethedika M AMJJAsa Figure 12.Numberoftransplantedseedlingswithleavesfrom 24Februaryto 23 October1976ateachstudysite.

PAGE 57

site.Leafshape,leafarrangement,andbranchorientationweredeterminedontheseedlingsatthetimeoftransplanting(January)andattheendofthestudy(October).Theleafshapedidnotchangefollowingtransplanting,howevertheothercharactersdidchange.AttheendofthestUdy,theleaveswereflattenedand fromthestemontheseedlingstransplantedamongexistingvegetation.Theendsofthestemswereflat,notascending.Theseedlingstransplantedintotheclearedareasexhibitedpondcypresscharacteristicsofappressedleavesleavesalongthestemandascendingbranches.SeedsSeedGerminationintheFieldAtallsitesseedgerminationwaslow.averaging2.1%ofallseedplanted. Naximum germination >:a,s 4.8%forpondcypressseed SOlTel intheclearedareaatthelogged,above-the-dikesite.Nodifferenceinpercentgerminationwasdetected betHeen pondcypressandbaldcypressseed.Attheburned,above-the-dikestUdysite,seedshadbegungerminatingintheclearedareasbytheeleventhdayfolloHingsowing(Table11)andamongexistingvegetationafter17days.The numberofgerminatedseedlings greatestintheclearedplotson March2,andintheplotsamongexistingvegetationon Barch 16.ByMay14thenumberofgermi nated seedlingsinallplotsbegantodecrease,probablyduetorecedingwaterlevels.Oneseedling sUL'rived throughbothdryandwetseasonsandwasstillaliveinOctober. Mean germinationforbaldcypressseedsownattheburned,above-the-dikesitewas 2.8%,while1.6%ofthepondcypressseedgerminated.Meangerminationofallseed 801m atthesiteHas 2.2%.

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IiIIIIITable11.Numberoflivingseedlingsattheburned,above-the-dikestudysite.I f Feb. Feb. March March March MarchApril May JuneJulyOctober Treatment Seed Source !1t.1!L...1..229162217 14 ..1.L.JL .23Existing pondcypress 1 250* 0 134 532 11 -l vegetation2 250* 0 0000000 0 --2 3250* 0 0031111 1 0baldcypress1 100* 0013111 11 0 2 100* 0 035 1110 0 --2 3100* 0 23 3221 1 10Clearedpondcypress 1 250-* 1 5 5 55 4 00 0 --2 of2 250* 037 55 1100 -.!2 existing3250* 1 4 4 2 1 0 0 0 0 0vegetationbaldcypress1 100* 0 11 23000 0 0 2 100* 0 12 2 1000003100* 0 1 1 2 0000 0 0 *numberofseedsplanted jnundated\.n o

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51Theseedplacedatthelogged,above-the-dikesiteforsoaking ",as lost,sotheseedsoakingatthelogged-and-burned,above-the-dikesitewasdividedandsownatbothsites.Thisiswhythenumberofseedsownatthese two sites(Tables12and14)isone-halftheamount sownatother sites.GerRination washighestatthelogged-and-burned,above-the-dikesite 12)on 14,butbyJuneII,allbuttwoofthe14germinatedseedlingshaddiedfollowinglossofsoilmoisture.Theremainingseedlings >:ere inundatedbyJuly17anddeadbyOctober2J.Averagegerminationatthesitewas 1.5% for seedand 1.7% for baldcypressseed.Meangerminationatthesitewas1.6%.Peak gerRinationwas observedintheclearedareasattheloggedand-burned,below-the-dikesite(TableIJ)one monthbeforemaximum germinati::m intheplotslocatedamongexistingvegetation.MortalityintheclearedplotswashighestinAprilandMay, when levelswerelowest. Hean germinationatthesitewas1.9%.Thepondcypressgerminationaveraged2.1%;thatofbaldcypressaveraged1.7%.Meangerminationatthelogged,above-the-dikestudysite(Table14)was 2.2%,,dth pondcypressseedaveraging2.8%,and baldcypress seedaveraging 1.7%. was muchlowerbelowthedike,averaginglessthan 1.0%. Pondcypressseedgerminationaveraged1.1% belowthedike;thatofbaldcypressaveraged0.8%.Germination higherintheclearedareas,bothaboveandbelowthedike,roughlyone monthbeforepeak intheunclearedareas.MortalityoccurredbothaboveandbelowthedikebeforeinundationinJune. Mean percent germination washigherintheclearedplotsatthelogged sites forbothpondcypressandbaldcypressseed,althoughthe

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Table12.Numberoflivingseedlingsatthelogged-and-burned.above-the-dikestudysite.MarchAprilMayJuneJulyOctoberTreatmentSeedSourcePlot221714 -1L..1L2'3 Existingpondcypress1 12Y 2 110 0vegetation2 125* 0 1 0 0 0 3 125* 0 3 0 0 0baldcypress1 5 0 0 1 110 2 5 0 0 0 0 00 3 5 0 0 3 0 0 0Clearedofpondcypress1 125* 1 0 00 -..2 existing2 125* 0 1 0 0 0vegetation 3 125* 0 3 0 0 0baldcypress150 -x -0 000__0 2 5 0 0 0 0 0 0 3 5 0 0 1 0 0 0 *numberofseedsplantedinundated f
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Table13.Numberoflivingseedlingsatthelogged-and-burned,below-the-dikestudysite.MarchMarchMarch MarchApril t1ay JuneJulyOctoberTreatmentSeedSourcePlot29162217 1411 -1L __ 23 Existingpondcypress1 250* 0 0 0 00000vegetation2 250* 0 3 497 7 753 250* 0 21 00 000baldcypress1 100* 00 0 0 0000 2 100* 0 1 3 22221 3 100* 0 20 00 0 0 0Clearedpondcypress1 250'0 4115 0000of2 250-* 285 00 0 0 0existing3 250* 0000 20 0 0vegetationbaldcypress1 100* 0 3 110000 2 100* 0 10 0 0000 3 100-* 0020 000 0 *numberofseedsplanted_inundated

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Table14.Numberoflivingseedlingsatthelogged,above-the-dikestudysite.MarchAprilMayJuneJulyOctoberTreatmentSeed SourcePlot2217 1411 -1L2'3 Existingpondcypress1125*0 2 0 0 0vegetation2 125-* 0 0 0 0 0 3 125*1 0 0 0 0baldcypress1 50* 0 1 0 0 0 2 0 0 0 0 0 3_50* 0 0 0 0 0Clearedofpondcypress1125*90 0 0 0existing2 125* 0 0 0 0 0vegetation 3 90 0 0 0baldcypress1 50* 2 1 1 0 0 2 50* 0 0 0 0 0 3 _50*2 0 0 0 0*numberofseedsplanted_inundated ':B-

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Table15.Numberoflivingseedlingsatthelogged,below-the-dikestudysite.March MarchAprilMayJuneJulyOctoberTreatmentSeedSourcePlot16 22 17 11+..1L -1.L2'3Existinc; pondcyprcsG 1 000 0 00vegetation2 250* 0 1 2 0 0 0 3250-* 0 0 00 0 0baldcypress1 100* 0000 0 0 2 0 110 0 0 3 100* 0 00 0 0 0Clearedofpondcypress1 250* 0 3 3 000existing2 250* 0 10 1 0 0 0vegetation 3 250* 01000 0baldcypress1 100* 0 1+ 0000 2 100* 000 0 0 0 3 100* 00 0000 *numberofseedsplanted_inundated di

PAGE 64

observeddifferencesarenotstatisticallysignificant(Fig.13).Atothersites,nOdifferencesingerminationofpondcypressseedandbaldcypressseed Here observed.Nosignificantdifferenceinseedgerminationwasobservedeitheramongsitesorbetweentreatments.SeedGerminationintheGreenhouseGerminationofcypressseedwas muchhigherinthegreenhousethaninthefield.Meangerminationofpondcypressandbaldcypressseedsoakedatthestudysites,thensowningreenhouseflats,isshowninFigure14.Theseedsoakedfor41daysattheburned,above-the-dikesite,for 58 daysatthelogged-and-burned,below-the-dikesite,for72daysatthelogged,below-thedikesite,andfor78daysatthelogged-and-burned,above-the-dikesite.Theseedatthelogged,above-the-dikesite ,2S lost.Nostatisticaldifference Has detectedbetHeenpondcypressandbaldcypressseedgerminationintheflatsusinga two-Hay ofvariance,norwasanydifferencedetectedwhichcouldbeattributedtosoakingtreatment.SeedGerminationintheLaboratoryGerminationofacidscarifiedseedinthelaboratorywashigherthangerminationofseedsowninthefieldorgreenhouse.MeangerminationpercentageswerecalculatedfromfourlOa-seedreplicates(Fig.15).Germinationcommenced onthefifthdayfollowingtreatment.Anaverageof 5.5% ofthepondcypressseedand 11.3% ofthebaldcypressseedhadgerminatedbythetenthday.Afterthirtydays,anaverageof lO.5b ofthepondcypressseedand14.8%ofthebaldcypressseedhadgerminated.Thegerminationofpondcypressseedrangedfrom 6to16%,whilegerminationofthebaldcypressseedrangedfrom11to18%.Nosignificantdifferenceinthirty-daygerminationpercentageswasdetectedbetween

PAGE 65

sr4 o z 3o r-<::z::;:n::2'wt9l5J PONDCYPRESSoBALDCYPRESS nature!! clearednatural01 corednatura!natural cleorod natural cleoredarcosareosarcaS oreasarcasorcasorcasOrcasorcosarcasBurned, Logged andBurned,LoggedandBurned, Logged, abovethe di!(e above the dike balowthedike above thedike belowthedike 13. Meangerminationofpondcypressahdbaldcypressseedssownateachstudysite.

PAGE 66

58 rsJ pONDCYPRESS c=J BALDCYPRESS4 2161214 10 z 0 t-oz? 8 D:: W rG 6Burned) Loggedand bumed)Logged, Logged end burnad) Gbove fhebelow tha dike below tha dike above the dike Figure14.Germinationofpondcypressandbaldcypressseed so,m ingreenhouseflatsfollowingsoakingatstudysites.Valuesaremeans;barindicates+onestandarderror.

PAGE 67

592030252015DAYS105 "0 15100 '" '" '""0'"'0'""0-'" '"'"'"'" 1067 -=oJ!C.-0 z 0 !C.-z z 0 533 .....re<-LUZI.!)::;: BALDCYPRESSSEED Ct: w C) PONDCYPRESSSEEDo57100 33 30252015 "0'"'".0"0'"-g 10 z o !;r 5 zDAYS Figure15.Cumulativegenninationofpondcypressandbaldcypressseedinincubatorfollowingacidscarification.Percentagesarebasedonalltreatedseedandonallfull,ungenninatedseed.Valuesaremeans,+onestandarderror.

PAGE 68

60pondcypressandbaldcypressseedusingat-test(Mendenhall,1974).Afterthirtydays,theremainingungerminatedseedswerecutopenandexaminedtodeterminethenumberoffullseed(asevidencedbyfleshyendosperm).Germinationbasedonfullseedwas 97% forbaldcypress,whereasonly 72% ofthefullpondcypressseedhadgerminated.AerialPhotographsThevegetationmapmade fromphotographstakenin1953(Fig.16)showsanextensiveTaxodium-dominatedforestpriortologging.Thecypressclosertothecentralmarshhadlargercro,rnsthanthoseclosertothewetprairies.Also,therewere morecypressperunitareanearthecentralmarshthantherewerenearthewetprairies.Willow(Salix)wasfoundontheborderbetweenthecypressstrandandthecentralmarsh.Prominentfeaturesonthe1953photographsincludeda mixed hardwoodstandwithinthecypresseastofthecentralmarsh,plUStherecentlycutoverpineland.Themapfrom1963photographs(Fig.17)indicatesthatmixedhardwoods werepresentontheformercypress-dominatedstrandapproximatelynineyearsfollowinglumbering.Sixmonthsaftertheburntherewereextensiveamountsofbaregroundintheloggedareas.Thetramroadsandscarsfromskiddin6weredistinguishableonthephotographsandremnant,uncutcypress Here visibleonthefringesofthecentralmarsh. WilloH stillborderedthecentralmarsh,butwasnotasextensiveasithadbeenin1953.Theremainingcypressclosetothetramroadshadsmalldiametersandwerecloselyspaced.AgricUlturalfieldswereprominentontheeasternborderofthemap. ThemapinFigure18(fromphotographstakenin1972)indicates th.at unburnedmixed hardHoods stillpersisteighteenyearsafterlogging.The

PAGE 70

62,',.... ... .. .... .,,.-,.'..'. ...m WATER [] WILLOW tJ BARE GROUNDDPleJE o0.5 tKILOMETERS oCYPRESSL [JJMIXED HARDWOODS[]\'lETPRAIRI"::._-------

PAGE 72

oCYPRESS [[[]lMIXEDf-'ARD'ivOODS o0.5 ItKILOMETERS IN OPINE MARSH oWET PR.l\:RIE WATER WILLOW El BARE GROUND 64

PAGE 74

66 WATER W WILLOW EI BARE GROUND +.;' : :.."-"-D PINE IS:]r,lARSH0" ,, ;; ,',' oCYPRESS ITJI]] MIXED 0WET PRAIRIE :-, () ,o0.5 )' KI LOI,lETERS N.i ; 'I'j,-."" ,T.J';" 'r-::rh: : .. ...

PAGE 75

67logged-and-burnedarea,which wasbaregroundin1963wascolonizedbywillow.Anareaofunlogged,smalldiametercypress(northoftwinlakesinlowermiddleright)which was burnedalsobecamedominatedbywillow.Cypressstilldominatesnorthandwestoftheloggedarea,andisfoundonthefringesofthemarshintheloggedarea.Noticeableistheadditionoftheleveeacrossthecentralmarshandthecreationoffournewlakes:borrowpitsforthesoilusedinthedikeconstruction.

PAGE 76

DISCUSSIONCypresstreesarescarceintheformercypress-dominatedcommunities "hich"ere logged,burned,andlogged-and-burnedat Corkscre"S"amp Sanctuary.Verylimitedregenerationofcypressisoccu=inginthestrandthat HaS loggedandthestrandthat "as burned,butthereisnocypressregenerationintheareathat "as bothloggedandburned.Requirementsforseedgerminationandseedlingsurvivalmayhavebeenalteredbythedisturbancesofloggingandburning.Becauselittleornocypressregenerationisoccurringintheloggedandburned sHamp, itmaynotreturntoitsoriginalfloristiccomposition.Iftherequirementsforcypressregenerationare management schemesmaybedevised "hich couldresultinthereestablishmentofcypressasa componentofthesedisturbedsystems.CypressRegenerationVerylittleregenerationofcypressintheloggedareasandburnedareasof Corkscre" hasbeenfromvegetativepropogation.Recoveryoftheforestbyvegetativemeansafterloggingwaseliminatedbecausethetrees "ere girdledpriortofelling,apractice "hich reducescoppicing 1915).Thefirekilledordamagedtherootsystemsofthetreesintheburnedareas,alsoeliminatingregenerationbycoppice.Therefore,thenaturalregenerationwhichhasoccurred,andcanbeexpectedtooccur, r2S beenfromseed.Naturalregenerationfromseedisdependentupon avarietyoffactors "hich affecttheseedavailability,germinationandseedling68

PAGE 77

69survival(Smith,1962).SeedAvailabilityAnadequatesupplyofseedforabundantregenerationisnotavailabletothedisturbedsites.Thescatteredremainingcypresstreesproducealimitedamountofseedandthe seed whichisproducedinsitumaybeinfertileorconsumedbyanimals.Thedisturbedareasareisolatedfromareaswhere muchseedisproducedduetotheimmobilityofcypressseed.Someseedisproducedbyacypresstreeeveryyear,butgoodproductionoccursonlyaboutonceeverythreeyears(Mattoon,1915).Thetreesattheburnedstudyareaandloggedstudyareawerenotobservedtoproducea goodseedcropduringtheyearofstudy,andnoregenerationwasnoticedduringthisyear.Thepresenceofcypresssaplingsatthesesites, h01'ieVer, indicatesthatsufficientseedisproducedforlimitedregenerationduring years.Theseedsthatareproducedmaybelessfertilethanseedproducedinundisturbedareas.Taxodiumismonoecious,butcrosspollinationbywindaccountsformostfertilization (Heier etaL,1970).Fertilizationisnotaslikelytooccurinthedisturbedarea,wherelesspollenisproduced,asinadenseuncutorunburnedstandoftrees.Also,squirrelsconsumecypressseedinCorkscrewSwamp.Predationonseedinthedisturbedareasdecreasesanalreadylimitedsupplyofseedavailableforregeneration.Viablecypressseeds,whichareproducedintheundisturbedportionsofthestrand,donotmoveinthedisturbedareas.Cypressseedsmoveonlywhilefloatingonwater(Fowells,1965).Theslow flOH ratesofwaterinCorkscreHSwamplimitthedistancetheseedcantraveL

PAGE 78

70 suchasvegetation,felledlogsortheabandonedtramroadsalsoimpede movementoftheseed.Theseedsdonotfloatmorethanacoupleofdaysandconsequentlydonotmovemuch beyondtheshadowoftheparenttree. Germination Thewaterrequirementsforgerminationofcypressseedaremetatthestudysites,butlowseedviabilitylimitsnaturalregeneration.Cypressseedmusthaveanabundantmoisturesupplyforonetothreemonthsafterseedfall 1915),whichoccursfromOctoberto Decenber (Fowells,1965).Standingwater observedatthestudysitesfrom onetothreemonthsafterseedfallandwaterlevelswere lo;rerthangroundlevelbeginninginFebruaryatthesitewiththeshortesthydroperiod.othersitesremainedfloodeduntilMarch.Atallstudysites,theseedswereinundatedlongenoughtoimbibeasufficient arr,ount of forgermination.The numberofseedwhichgerminateddidnotseemtobeaffectedbythelengthoftimetheseed inundated.ThegerminationoftheseedfromCorkscrewSwamp was low,averaging10to 15% ofallincubatedseed.Only15%ofallseedcollectedwasfull,indicatinglowviability.Nodifferenceinviability notedbetweenpondcypressseedandbaldcypressseed.ThepercentagesoffullseedwhichgerminatedduringthethirtydaytestperiodwerecomparabletothosemeasuredbyBonner(1974)andMurphyandStanley(1975).LowviabilityofthecypressseedatCorkscrewaccountsforlowpercentgerminationofseedsplantedin flatsandinthefield,andisalimitingfactorincypressregeneration.SurvivorshipDuringtheFirstYear Asurvivorshipcurve (Fig. 19) Has constructedforthestUdyperiod

PAGE 79

(IiCC o?:> c::;CJen 1000500100-5010 5 SEED SOWN (normalized 101000 individuals) t::JIAEAN GERMINATION OF INCUBATED SEED &MEAN GERMINATIONOFSEEDINGREENHOUSE GMEAN GERMI,\jAT!ONOFSEEDINTHE FIELD7140, J !80120160200 TIMEFROM SOWING(DAYS) 240 230Figure19.Meansurvivorshipofcypressseedsownatstudysites.

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72toestimatea meansurvivalrateofcypressseedsowninthe field. The mean numberofgerminatedseedlingssurvivingtoagivenday was calculatedbaseduponallseedsown.Althoughthecurveisforlessthanoneyear,itprobablycoversthemostcrucialtimeofseedlingdevelopment.Thecurveisa TypeIII(Deevey,1947),typicalofaspecieswithhighinfantmortality.ItissimilartothesurvivorshipcurveofAcersaccharumseedlings(HettandLoucks,1971)inthatthemortalityrateisnotconstantduringthefirstyearofexistence.Avarietyoffactorsinfluencethesurvivalofcypressduringtheyearfollowingseedfall.Immediatesurvivalisdeterminedbythenumberofseedwhich gerrranate. Someofthenewlygerminatedseedlingsdieasaresultofdroughtconditionscreatedbyrecedingwaterlevels.Theseedlinglnust growtall duringthedryseasontoescapeinundationaswaterlevelsrise.Abundantnaturalregenerationoccurs onyearswhentheproperconditionsforsurvivalcoincideswitha goodseedcrop(BUll,1949;Rathborne,1951;Stubbs,1972).Survivalthroughdryseason.Initialmortalityofthenewlygerminatedseedlingoccursduringthedryseasonaswaterlevelsrecede.Demaree(1932)andLangdon (1958) bothindicatethatsufficientgrowthtoescapeinundationiscrucialforcypressseedlingsurvival.AtCorkscrewSwamp,lackofsoilmoistureisagreatercauseoffirstyearmortalitythaninundation. A totalof148seedlings fromseedsownatthestUdysites.Ofthattotal,123(83%)diedduringthedryseason,15(10%)sufferedinundationmortalityand10(7%)survived(Fig.20).Theoldertransplantedseedlingssurvivedthedryseasonbetterthanthenewlygerminatedseedlings.Thetransplantedseedlingsprobablymaintainedrootcontact }ath thesubsurfacewater,andasa

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10080 Cl..J 60 aenL1.. 0 .....z 40wu c:w CL20o(130)(22)(120)73Pond cypress BoldcypressPondcypress BaJdC'jpressPcndcjpres.s ClearedAreasAmong Transplanted r&l TRANSPLANT MORTALITY MORTALITY DURING DRY SEASON MORTALITYDURINGWET SEASON oSURVIVEDTHROUGHSTUDYPERIOD() NUMBER OFSEEDLINGS 20.Fateoftransplantedseedlingsandseedlingsgerminatedfromseed S01in atstudysites.

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74only17%oftheseedlingsdiedduringthedryseason.Rootcontactwithwaterisimperative,becauseirreversiblewiltingoccursincypressseedlings3-4hoursfollowinglossofsoilmoisture(DicksonandBroyer,1972).Theseedlingsaremorelikelytobeeatenbyanimals,suchasrabbits(Fowells,1965),duringthedryseasonbecausetheseedlingsaremoreaccessable.Inundationmortality.Demaree(1932)andBull(1947)reportthatTaxodiumcannotsurviveprolongedsubmergence.Welch(1932)observedresproutingfollowingasix-weekinundation.AllseedlingsthatdiedatCorkscrewSwampwere submergedforlongerthansixweeks. Thesurvivors(bothplantedandtransplantedseedlings)were onhigher,driersitesandthereforeabletoescapeinundation.CompetitiveAbilitvofCypressSeedlingsTheexistingvegetationateachstudysiteappearstoinhibit,butnotprohibit,cypressregeneration.Growthandsurvivalofthetransplantedseedlingstendedtobehigherinclearedareasthanamongexistingvegetation(Fig.21),butthedifferenceswerenotstatisticallysignificant.Anysignificantdifferenceingrowthwhichreflectstoleranceorintoleranceofcompetitionmaytakeyearstodetectbecauseoftheslowgrowthratesofcypress.Postetal.(1975)measuredadifferenceingrowthratesofcypressseedlingsaftertwoyears(butnotafteroneyear)followingtransplantingintocypressdomesreceivingsecondarilytreated sevrage anddomesreceivinggroundwaterinputsnearGainesville,Florida.Thedifferencein grovrth betweenclearedareasandunclearedareas(Fig.21)ismaskedbyhighvariation.Over alongerstudyperiod,thedifferencemighthave becomesignificant.The

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75o80100o50 40L30 60 w(I):r:20 :;: 40 <.'>c::W::> :r: (I) Z Z
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76developingseedlingsmightgrowbetterinclearedareas,indicatingthattheexistingvegetationinhibitsregrowthofcypress.Thepresenceofcypressseedlingsatsiteswherethereisanoverstoryofcypressindicates,however,thatregenerationisnotentirelyexcludedbyoverstoryvegetation.Theexistingvegetationmayactasashelterearlyinthelifeofaseedling,butoncetheseedlingistallenoughtoescapeinundationtheeffectsseemtobedetrimental.Theseedlingswhichgerminatedunderexistingvegetationsurvivedbetterthanseedlingswhichgerminatedinclearedareas(Fig.20).Thevegetationprobablydecreasesthe trans pirationaldemand ontheseedlingbyincreasingthehumidityanddecreasingthesUnlight.Moretransplantedseedlingssurvivedinclearedplotsthaninplotsamongexistingvegetation(Fig.21).Percentsurvivalbasedonthehighestnumberofleafed-outseedlingsforeachtreatmentat each sitewascalculatedtoseparatetransplantdeathfromdeathduetocompetition.The meanpercentsurvivalofallseedlingstransplantedintoclearedareas,excludingdeathsduetotransplantshock,was 86%,whereasonly 6J.J% ofseedlingstransplantedamongexistingvegetationsurvived.The two meansarenotstatisticallydifferent,yetsuggestthatsurvivalmayhavebeenhigherinclearedareas.Taxodiumexhibitsmorphologicalplasticityinresponsetomicroclimaticvariables.Theseedlingstransplantedintotheclearedareashadleavesappressedtothebranchandbranchesthatwereascending.Thesecharacteristicsareprobablyaresponseto "'ater stresscreatedbyincreasedevapotranspiration.Theleavesoftheseedlingsplantedamongexistingvegetationwereflattened fromthebranchandthebranches

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77wereflattenedaway fromthemainaxis.Theseedlingsintheclearedareasleafedoutbeforeseedlingsplantedamongexistingvegetation.Seedlingsgrowninthegreenhouseleafedoutpriortoseedlingsinthefield,indicatingthattemperature,butnotphotoperiodorwaterrelations,maybeanimportantpartofthetriggeringmechanism.THOofthethreeobservedcharacteristics,leafandbranchorientation,variedwithmicroclimate,whereasleafshape remained constant.Seedlingswhich grewundersimilarconditionsinthegreenhouse,butgerminatedfrompondcypressandbaldcypressseed,exhibiteddifferencesinmorphology.Thus,myobservationsofseedlingsproducedfromseedsharvestedfrom knownparenttrees,plusmyobservationsofcypresstreesofvariousagesandon avarietyofsitesinthefield,convincemethatpondcypressandbaldcypressaredifferent.Untildefinitivegenetic,cytological,andtaxonomicstudiesareconducted,itisprobablyadvisabletocontinuetoregardthemastwodistinctspecies.SuccessionalStatusofCypressForestsAgeneralizedschemeofsuccessioninsouthFloridaswampsisproposedinFigure22;itincludesfourcommunitieswithcypresspresentandtwowithoutcypresspresent.Thelargestremnantofthecypress-mixedhardwood communityinsouthF'loridaisinCorkscrewSwampSanctuary.Thiscommunityischaracterizedbylarge-diametercypresstreesforminganoverstoryaboveanunderstoryofmixedhardwoods.ThisvegetationtypewasprobablythehistoricallyubiquitousswampforestinsouthFlorida,SUbjecttochangebynaturalforcessuchashurricanes,frosts,andfire.ChangesduetoFireAsurfacefireinthecypress-mixedhardwoodforestusuallykills

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78 cypressregenerationnofire cypress regenerationmixedhard Hoods +remnant cypress lightfiressevereburnlogging HilloH firenofiremixedhardHoodscypress+mixed hardwoodssurfacefirenofireseverefiremonospecificsevereburnorlogging cypress forestssurfacefire -----r-----j nofire HilloH +remnantcypress Fi@lre 22.GeneralizedsuccessionschemeinsouthFlorida sHamps.

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79thehardwoods.Cypressismoretolerantofasurfaceburnthanmost hardwoods(Cypert,1961;EwelandMitsch,1975),sosurfaceburnstendtoweedoutthehardwoods,leavingmonospecificstandsofcypress.Ifthecypressisunburnedforalongperiodoftime,thehardwoodsinvadeandreestablishacypressmixed hardwoodforest.Aseverefireinthecypressmixed hardwoodtypewillnot killthehardwoods,asinasurfaceburn,butwillalsokillsomeorallofthecypress.Cypert(1961)andAlexanderandCrook(1973)statethat ofthepeatsubstratebyseverefiresdestroystherootsystemsofalltrees,eliminatingrecoverybycoppice.Removaloftheorganicsoilalsoalterssoilpropertiesofwaterretentionandnutrientavailability.Theeliminationoftreesandvegetativeregrowthleavesthesiteopenforinvasionbyotherspecies. forestsdenudedbyasevereburninsouthFloridaareoftencolonizedbywillow,Salixcaroliniana.Willowhasreadilydispersedseedandrapidlyinvadesseasonallyfloodedsitesafterdisturbancessuchasburning(Robertson, 1953) orlogging(Allen,1962)havealteredthesubstrate,providedthatsomesoilorganicmatterremains(Loveless,1959).Willowstandsaremaintainedby freQuently recurringfires(Craighead,1971).Ifremnantcypressarepresentandifsurfaceburnsrecur,thewillow-dominatedcommunitiesonceagainbecomecypressdominated.Iffireisexcludedfromthewillow-and-remnant-cypresscommunity,theninvasionofhardwoodscoupledwithcypressregenerationwillcontinueuntila mixedforestofcypressandhardwoodspersist.Thus,thenaturalpatternsexpressedinFigure22canbesummarizedasfollows:fireinsouthFlorida s"Bmp forestshaslongbeenoneofthedeterminantsofvegetationchange.Priortoman'sintervention,

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80thewetlandforestswereprobablyoneofthreegeneraltypesdependinguponthefrequencyandseverityofburning. S,Bmps which werenotburnedforlongperiodsoftimeweremostlycypress mixed hardwoods.Surfacefireskilledthehardwoods,leavingpurecypressforests.Severefireswoulddestroyallofthevegetation,allowinginvasionbywillow.Thewillowforestswere,andare,maintainedby frequent fire,butchangeintoacypressmixed hardwoodforestiffireisexcluded.Changes duetoLoggingorLogging-and-BurningSuccessionincypressmixed hardwood swampsfollowinglumberingresultsincommunitiesdominatedby mixed hardwoods. The cypressareremovedbylogging,leavinga fewsmall-diametercypress.Thehardwoodsarereleasedfromoverstorycompetitionandcapturethesite.Asurfacefireinthemixed hardwood,remnant-cypressforest rzy enhancecypressregenerationbyreducingcompetitionfromthe hardooods. Thecypresscanthensuccessfullyregenerateifproperconditionsforgerminationandseedlingsurvivalarepresent.Astandofmixedhardwoods,withremnantcypress,shouldpersistiffireisexcluded.Redmaple,Acerrubrum,anddahoonholly,Ilexcassine,arethedominanttreesattheloggedsiteswithinCorkscrewSanctuaryandbothspeciesareregeneratingactively.Red maplemayliveupto150years(Fowells,1965),sothistree.maycontinuetodominateforalongtimeiffireisexcluded.Annona,Cephalanthus,Myrsine,Perseaand arealsoactivelyregenerating.Remnantcypressshouldremaininthe conoounity forlongperiodsoftime,withorwithoutregeneration,becauseofthelongevityofcypress.IfsufficientnumbersofcypressarepresentinthemixedhardWOOdsafterlogging,thecypresswillreproduceandthecommunitywill return

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81tothepre-loggingcypressmixed hardwoodstatus.Ifinsufficientseedarepresentand/orcompetitionfromthehardwoodseliminatescypressregeneration,theforest ,nIl become astandofmixed hArdwoods withnocypress.Cypert(1961)andAlexander and Crook(1973)agreethatloggedswampsaredamagedmuchmorebyfirethanunloggedswamps. The hardwoodsareallkilledbytheburn,whichismoreintensebecauseofthethick undergro1rth, andtheresultantbareareaisopentoinvasionbywillow.Thewillow ismaintainedbyfrequentfiresandunchangedwaterlevels(Craighead,1971).Withtheeliminationoffire,hardwoodsinvadethelogged-and-burned,willow-dominatedstand.SeedlingsofAnnona,AcerandPerseawerefoundwithinthelogged-and-burnedstUdyarea.Alexander and Crook(1973)reportedsimilarobservations,indicatingsuccessiontowarda mixed hardwoodforestwhichisdevoidofcypress.Cypressisdifficult,butnotimpossible,toreestablishonceremovedbylumbering.Cypress-freemixed hardwoodforests,however,arealikelylong-termsequeltologgingfollowedbyasevereburn.ManagementRestorationoftheTaxodiumdominated inthedisturbedareasofCorkscrewSwampSanctuarymaybeonedesirablegoalofsanctuarymanagement;actionsareavailablebywhichtheproblemsassociatedwithcypressregenerationmaybeOvercome.Cypressseemtobereproducingnaturallyinareaswhereseedsourcesremain,butloggingandburningremovedmostoftheseedsourcesandcypressseedsdonotmovefromtheundisturbedareasintothe disttITbed areas.Lowseedviability,highseedlingmortalityandslow ratesofseedlingsseemtolimit inareaswhereseedispresent.Ifcypressisreestablished

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82inthedisturbedareas,theresultantcommunitymaybe moreresistanttochangefollowingburning.Also,itwouldeventuallyreplacesomeoftherookeryhabitatlostbylogging.Twomethodscouldbe employedinrestoringtheTaxodium:directseedingorseedlingplanting. So'Ning seedwould belessexpensive and laborintensive,buttheexpectedsurvivorshipwouldbeverylow.InthisstUdylessthan1seedoutofevery1000plantedsurvivedthefirstyear,indicatingthatapplicationratesofseed would havetobeveryhighforabundantregeneration.Purchasedseedmaybehigher in viabilitythanseedcollectedon-site,butdrought and inundationmortalitywouldlimitthenumberof survivors. Theothermethodofreforestationwouldbetotransplantyear-oldseedlingsinthedisturbedsites,a method whichhasworkedsuccessfully in Louisiana(Bull, 1'147; Rathborne, 1951). Tomaintainthegeneticstock,seedcouldbeharvestedinthearea,scarifiedwithacidtoincreasethegerminationrate, and sowninpots.One-year-oldseedlings,whichwereraisedfromtheseedgerminatedingreenhouseflats,averaged1m in height:tallenoughtoescapeinundationifplantedinthedisturbedareas.Transplantingofpottedseedlingsintothefieldshouldprobablybedone fromJanuarythroughMarch, whentheseedlingsarestillleaflessandwaterlevelsarelow enoughsothatmuchoftheswampisreadilyaccessible.Bull (1947) recommended a 5 mx5mplantingdistance.NaturaldensitiesoftreesincypressstrandsatCorkscrewSwamprangefrom6m 2pertreeinthepondcypressregionto100 m 2pertreeinthebaldcypressregions(Dueveretal.,1975).Assumingfirstyearmortalitytobe 33% ofalltransplantedseedlings(thisstUdy),the planting Hould result in adensityof38m 2pertree one yearfollowing

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83planting.Ifthedisturbedarea(175ha)isplantedatthisdensity( 25 n2 pertreeor400treesperha),then70,000seedlingswouldhavetobeplanted.Plantingofseedlingsatlowerdensitieswoulddecreasetheamountoflabor butwouldincreasetheregrowthtime.Seedlingsplantedatalowerdensitywould,intime,serveasseedsourcesforadditionalcypressreforestation. The waterimpounded bythedikeseemstobeaffectingthevegetationin the cutoverandburnedareasofCorkscrewSanctuary.Theimpactseemstoresultfromhigher v2ter levelsabovethedikeratherthanincreaseddurationofthehydroperiod.Becausethewaterisdeeperabovethedikeduringthewetseason,lesssubstrate--inclUdingsoil, hUIlllilocks, andstumps--isexposedthanbelowthedike.Cypressseeds exposedsubstratesforgermination,sotheincreaseddepthofwaterabovethedikemayberetardingthealreadylimitedamountof natural regenerationoccurringthere.Removaloftheboardsblockingthe c'llverts whichtraversethedikemightresultinlower v2ter levelsand increasetheamountofsubstrateavailableforrecolonizationby c;Yl'ress. The regenerationofcypressonlandswhich wereoncedominatedbyTaxodiuminCorkscrewSwampSanctuaryisconstrainedbybiologicalfactors, seedavailabilityandviability,narrowgerminationandseedlingestablishment inherentdifferencesbetweenpondcypressand andsitecapturebysuccessionalvegetation.Withintheseconstraints,however,therearetoolsavailabletotheSanctuarymanagement.Theyincludethepotentialforartificialregeneration,waterlevelmanipulation,andtheuseoffire.Usedtogetherinthepropercombin atioLs, thesetoolsshouldmakepossiblethereestablishmentofcypressforestsonmuchofitsformerrangewithinCorkscrewSwampSanctuary.

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LITERATURECITEDAllen.P.H. 1962.Blackwillowdominatesbaldcypress-tupeloswampeightyearsaftercutting.U.S.D.A.ForestService.SoutheasternForestExperimentalStationResearchNote177.2p.Alexander.T.R.andA.Crook.1973.VegetationchangesinsouthFlorida.AppendixG.InSouthFloridaEnvironmentalStudy.National Park Service.827p.Applequist.M.B.1959.Longevityofsubmergedtupeloandbaldcypressseed.LouisianaStateUniversityForestryNote 27. 2p.Beaven. G.F.1939.Pocomoke Astudyofa swampinMaryland.Bull.TorreyBot.Club66:367-389.Betts.H.S.Serial.1960.6p.Baldcypress.U.S.ForestService,AmericanWoodsBonner,F.T. 1974. Unitedstates.Taxodium.p.796-798. In SeedsofWoodyPlantsintheU.S.D.A.ForestService.AgricultureHandbook No. 450.Bull,H. 1949. CypressplantinginsouthernLouisiana.SouthernLumberman179:227-230.Carter.M.R., L.A.Burns.T.R.Cavinder.K.R. Dugger.P.L.Fore.D.B.Hicks.H.L.Revells,andT.W.Schmidt.1973.EcosystemAnalysisoftheBigCypressSwampandEstuaries.ReportU.S.EnvironmentalProtectionAgency.RegionIV. 374p.Craighead.F.C.1971.TheTreesofSouthFlorida,Vol.I:TheNaturalEnvironmentsandtheirSuccessions.Univ.ofMiamiPress,CoralGables.Fla.212p.Craighead,F.C. 1974. HammocksofsouthFlorida.pagesP.J.Gleason,(ed.).EnvironmentsofsouthFlorida:present.Memoir2:MiamiGeologicalSociety.53-60.InpastandCurtis.J.T.1959.TheVegetationofWisconsin:Anordinationofplantcommunities.Univ.ofWisconsinPress,Madison.657p.Cypert.E. 1961. TheeffectsoffireintheOkeefenokeeSwampin 1954 and1955. Amer. Nidl. Nat.66:485-503.Cypert.E.1973.PlantsuccessiononburnedareasinOkeefenokee S,mmp followingthefiresof 1954 and1955.Proc.Ann.TallTimbersFireEcol.Conf.12:199-218. 8l+

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Davis,J.H.1943.TheNaturalFeaturesofSouthernFlorida,especially tb.e Everglades.FloridaGeologicalBull.No.25. 311p.DeBell,D.S.,J.Stubbs,andD.D. Hook.1968.Standdevelopmentafteraselectioncuttingina rBrdwood bottomland.SouthernLumberman 217:126-128.Deevey,E.S. 1947. Lifetablesfornaturalpopulationsofanimals.Quart.Rev.BioI.22: 283-314. Demaree,D.1932.SubmergingexperimentswithTaxodium.Ecology13: 2.53-262. Detwiler,S.B.1916.The characteristics.baldcypress(Taxodiumdistichum),Amer.Forestry22: 577-581. identification R.E.andT.C.Broyer.1972.Effectsofaeration,watersupply nitrogensourceon grolrth anddevelopmentoftupeloandbaldcypress.Ecology53:626-635.Duever,M.J.,J.E.Carlson,andL.A.Riopelle.1975.EcosystemanalysesatCorkscrewSwampSanctuary,p.627-725. Irr H.T.Odum,(ed.), WetlandsforWater RecyclingandConservation. 2nd AnnualReport,CenterforWetlands,Dniv.ofFla.,Gainesville.Egler,F.E. 1954. composition,aVegetatioActaVegetationscienceconceptsI:factorinold-fieldvegeta,tionGeobotanica 4:412-417. initialfloristicdevelopment. EHel, K.C. and H.J.Nitsch. 1975. Theeffectoffireonthespecies composition oftreesin tHO cypressdomes,p.215-222.InH.T.Odum,(ed.), CyPress WetlandsforWater Management,RecyclingandConservation.2nd AnnualReport,CenterforWetlands,Univ.ofFla.,Gainesville. FOHells, H.A.1965.SilvicsofForestTreesoftheDnitedStates.U.S.D.A.ForestService.AgricultureHandbook 271. 762p.Gooch,F.S.1953. nursery clLltureofcypress.SouthernLumberman186:37-38.Harper,R.M.1927.NaturalResourcesofSouthFlorida.18thAnnualReportStateGeologicalSurvey,Tallahassee,Fla.p.25-206.Hett,J.M. and O.L. Loucks,1971.SugarMaple(AcersaccharumMarsh.)seedlingmortality.J.Ecol.59:507-520. Johrcson, R.L. 1972. Oak-Gum-CypressintheMidsouth,p.98-103.In SilvicultlLre Systemsof Najor ForestsTypesintheU.S. U.S.D.A. AgricultureHancibook No. 445. Kurz, H. andR.K.Godfrey.1962. TheTreesofNorthernFlorida.Dniv.ofFla.Press,Gainesville,Fla.311p.

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Langdon, O.G. 1958. Silvicalcharacteristicsofbaldcypress.ForestService,Southeasternstationpaper 94. 7p.Long,R.W.andO.Lakela.1971. AFloraofTropicalFlorida,ofSeedPlantsandFernsofSouthernPeninsularFlOrida.MiamiPress,CoralGables,Fla.962p.86 U.S.D.A. A Manual Univ.ofLoveless,C.M.1959. AstudyofthevegetationintheFloridaEverglades.Ecology40:1-9.Mattoon,W.R.1915. TheSouthernCypress.U.S.D.A.BulletinNo.272.74p.Mattoon,W.R.1916. Waterrequirementsandgrowthofyoungcypress.Soc.Am.For.Proc.11:192-197. Mendenhall,W.1975.Introductiontoprobabilityandstatistics.4thed.WadsworthPublishingCo.,Belmont,California.460p.Mueller-Dombois,D.andH.Ellenberg.1974. Ecology. John Wiley andSons,NewYork.AimsandMethodsofVegetation 547 p.Murphy,J.B.andR.G.Stanley.1975.Increasedgerminationratesofbaldcypressandpondcypressseedfollowingtreatmentsaffectingtheseedcoat.Physiol.Plant35: 135-139. Oosting,H.J. 1956. TheStudyofPlantCommunities. 2nded.W.H.Freeman andCo.,SanFranciscoandLondon. 440p.Pen found ,W.T.1952.Southernswampsandmarshes.Bot.Rev.18: 41; 446.Post,D.M., C.A.Hollis,J.B.MurphyandG.Deghi. 1975.Forestgrowthseedlingsandfertility,p. 48;-554. InH.T.Odum,(ed.),Cypress WetlandsforWater Management,RecyclingandConservation.2nd AnnualReport,CenterforWetlands, Univ.ofFla.,Gainesville.Prestridge,J.A. 1947. TheBigCypressSwampwherethemodernandprimitivemeet.SouthernLumberman175:126-130.Rathborne,J.C.1951. Cypressreforestation.SouthernLumberman183: 239-240.Robertson,W.B. 195;. A SurveyoftheEffectofFireinEvergladesNationalPark.U.S. Dept.ofInteriorReport,NationalParkService.169p.Small,J.K.1920. CypressandpopulationinFlorida.Jour.N.Y.Bot.Garden 21:81-86.Small,J.K.type.1931. Thecypress,southernremnantofanorthernfossilJour.N.Y.Bot.Garden 32: 125-135.

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87Small,J.K.1933. oftheSoutheasternFlora.Univ.ofNorthCarolinaPress,Chapel ,Hill. Smith,D.H. 1962. ThePracticeofSilviculture.7thed. John WileyandSons,NewYork, LondonandSydney. 578p.Snedacor, G.H. andH.G.Cochran. 1967.StatisticalMethods.6thed.The IO'B StateUniversity Ames, Iowa. 593 p.Sokal,R.R.andF.J.Rohlf.1969.Biometry. H.H. FreemanandSons,SanFrancisco.776p.Sprunt,A.Jr. 1954. FloridaBirdLife.CowardMcCann,Inc.,NewYorkandtheNationalAudubonSociety.527p.Sternitzke,H.S. 1972.Baldcypress:endangeredorexpandingspecies?Econ.Bot.26: 130-134. Stubbs,J.1972.Atlantic Oak-Gun-Cypress, p.89-93.InSilvicultureSystemsofMajorForestTypesintheUnitedStates.U.S.D.A.AgriculturalHandbookNo. 445.Heier, T.E.C.R.Stocking,andM.G.Barbour.1970. WileyandSons,Inc.,NewYork, London, Sydney.Botany.708p.John Welch, H.H. 1932.AnecologicalstUdyofbaldcypressinIndiana.Proc.Ind.Acad.Sci.41:207-213. West, E.andL.E.Arnold. 1956. TheNativeTreesofFlorida.Univ.ofFloridaPress,Gainesville,Fla.218p.

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BIOGRAPHICALSKETGHLanceH.Gunderson wasbornDecember7,1952inMiami,Florida.HeattendedpublicschoolsinLee County,FloridauntilgraduationfromFortMyersSeniorHighSchoolinJune,1970.InSeptember,1970heenteredtheUniversityofFlorida.HereceivedaBachelorofSciencedegree,majoringinBotany,inJune,1975.Upongraduationhe was acceptedintotheGraduateSchoolintheDepartmentofBotany.88

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IcertifythatI havereadthisstudyandthatinmyoplnl0nitconformstoacceptablestandardsofscholarlypresentationandisfullyadequate,inscopeandquality,asathesisforthedegreeofMasterofScience.IcertifythatI havereadthisstudyandthatinmy0plnl0nitconformstoacceptablestandardsofscholarlypresentationandisfullyadequate,inscopeandquality,asathesisforthedegreeofMasterofScience. 'if' ArielE. LugoAssociateProfessorofBotanyThisthesiswassubmittedtotheGraduateFacultyoftheDepartmentofBotanyintheCollegeofArtsandSciencesandtotheGraduateCouncil,andwasacceptedaspartialfulfillmentoftherequirementsforthedegreeofMasterofScience.August 1977 Dean,GraduateSchool

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,''.., .." .......'...,,'.'.'..00.5 0 PINE WATER---.. + KILOMETERS MARSH 19 WILLOW NDCYPRESSI!IIllMIXED HARDWOODS o WET PRAIRIE BAREGROUND

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,."""."""',.:::".".",""""""""..,"".'"."".:",""."",." .".".: ..00.5 0 PINE WATER",-.', f KI.LOMETERS MARSH e WILLOWNoCYPRESS ffiIlI MIXEDHARDWOODS IZ:I WETPRAIRIE EI BAREGROUND

PAGE 100

00.5 [] PINE WATER---"." f KILOMETERS MARSH Il WILLOWNoCYPRESS lIII] MIXEDHARDWOODS o WETPRAIRIE 8 BAREGROUND


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