Phytologia(July 1998)85(l):51-60. WOODY VEGETATION DIVERSITY OF LONGLEAF PINE COMMUNITIES IN CALCASIEU PARISH, LOUISIANA RayNeyland,HarryA.Meyer,&HeatherHarrington DepartmentofBiologicalandEnvironmentalSciences,McNeeseStateUniversity, LakeCharles,Louisiana 70609-2000 U.S.A. ABSTRACT LocatedinsouthwesternLouisiana, Calcasieu Parish oncesupported large areas of undisturbed longleaf pinelands. These pinelands were clear cut around 1900and have beenconverted largely toforest, range, and cropland. AlthoughvirginlongleafpinelandsnolongerexistintheParish, afew remnant sitesretainmanyoftheirpresettlementplantspeciesand characteristics. Three distinctlydifferentlongleafpine communitieswere identified and sampled to determinewoody species diversity and importance. Diversity values of the threecommunitiesdiffered significantly from each other. The acid longleaf pineflatwoodscommunitywas lessdiversethaneitherthehillsideseepagebog community or the sodic longleaf pine flatwoods community. The low diversity value for the acid longleaf pine flatwoods community was due primarily to low species richness when compared to the other two communities. Although the hillsideseepage bogand the sodic longleaf pine flatwoods each had identical species richness values, the disparity in their diversity values is explained primarily by the greater evenness of species abundance at the sodic longleaf pine flatwoods community. The various mixtures of woody species sampled suggest a high degree of heterogeneity amongthesiteschosenforstudy. Forexample,oftheelevendifferentspecies found among the three communities, only Pimis palustris was present at all three. Piniispaliistriswas themostimportantinallbut thesodic longleafpine flatwoods community where it was second to P. elliottii. The importance value forall combinedcommunitieswas clearlydominated by P. palustris at 140.69. Pinus eUiottii had the second highest importance value for all combinedcommunitiesat62.83 with Myricacerifera third at31.58. Relative coverageforall communitiescombined was alsodominated by P. palustrisat 55.82% withP. elliottiiandM. ceriferaat31.73% and6.56% respectively. KEY WORDS: Calcasieu Parish, Louisiana, longleaf pine communities, woodland remnants 51 PHYTOLOGIA 52 July 1998 volume85(1):51-60 & The longleaf pineland region of the West Gulf Coastal Plain (Bridges Orzell 1989) extends across northern Calcasieu Parish (Figure 1). This region originally extendedfromsoutheasternTexastonorth-central Louisiana(Wahlenberg 1946; Little 1971). These pinelands were clearcut in the early 1900's (Roy & Midkiff 1988; Smith 1991) and now existprimarilyas forest, range and cropland (Roy & Midkiff 1988). Remnantlongleafpinecommunitiesofthe WestGulfCoastal Plain identified byOrzell(1987)andBridges(1988) arecurrentlythreatenedby grazing, silvaculture, andwildlifemanagement(Bridges&Orzell 1989). Theoncevirginlongleafpinelands as described by Schwarz(1907) and Wahlenberg (1946) are now rare in Louisiana (Smith 1991). However, a few comparatively undisturbed sites within Calcasieu Parish appear to retain many of their presettlement plant species and remain recognizablelongleafpinecommunities(sensu Bridges & Orzell 1989; Smith 1991, 1996). Theseraresitesaresmallandmaybeconsideredremnantsofaoncedominant ecosystem. The purpose of this study is to characterize the adult woody vegetation of three remnantlongleafpinecommunitiesin CalcasieuParish in termsofdensity, frequency coverage, and importance. These communities are referred to as hillside seepage bogs, sodic longleaf pine flatwoods (sodic flatwoods), and acid longleaf pine flatwoods(acidicflatwoods). ThisterminologyissensuSmith(1996). STUDYAREA Located in southwestern Louisiana, Calcasieu Parish is adjacent to Beauregard Parish to the north, Jefferson DavisParish totheeast, Cameron Parish to the south, andtheSlateofTexastothewest(Figure 1). Approximately46%oftheParish's land isdevotedtoeitheragricultureorrangeland;approximately23% is woodland; 11% is marsh;4%swamp;andtheremainingareaisurban (Roy & Midkiff 1988). Land use isdevotedprimarilytotimber,cattle,rice,andsoybeans. Peux)leum-relatedindusuies arepresentintheP^shandareconcentratedaround theareaof Westlake. Calcasieu Parishisnoteworthyfortheabrupttransition from thelongleafpinelands in the north totheprairieandcoastalmarshesinthesouth(Figure 1). The first study site is a hillside seepage bog situated about 8 km northeast of DeQuincy;(93** 23' 83" x 30** 28' 92");rangeand township (RlOW x T7S Sect. 1) (Figure 1). This is the only known representative of this community type in the Parish. The siteextends approximately600 m on a north-south-axis by 36 m on an east-westaxis and is bounded to the west by a semi-evergreen broadleaf acid seep forest {sensu Bridges & Orzell 1989), and to the east by a stand of Pinus eUioaii Engelm. Thisbogappearstobetheheadwatersofasmall unnamed stream. The soil type is "Glenmora silt loam" (Roy & Midkiff 1988) and is characterized by an extremelyacidicsilt loam surface layer and a strongly acid silty clay loam subsoil. Distinctiveherbaceous species include Rhynchospora oligatuha A. Gray, Sarracenia alata (Wood) Wood, and Utriculariacomuta Michx. Charred bases of P. palustris Mill, indicatethesitehadburnedwithinthepastseveralyears. Neylandetal.: WoodyvegetationinCalcasieuParish 53 CoastalMarsh BottomlandForests CoastalPrairie Swampland LongleafPine Figure 1. Area of longleaf pineland in relation to other regions within Calcasieu Parish, Louisiana. Insetshows thelocationofthe Parish within the State. Study site #1 = hillsideseepagebog;#2= sodicflatwoods;#3 =acidicflatwoods. PHYTOLOGIA 54 July 1998 volume85(1):51-60 Thesecondstudysiteisasodic flatwoods thatis situated nearafew othersimilar sites in thewestern part of Calcasieu's longleaf pineland region (Figure 1). These flatwoods are typicallysaturatedduring the winterand spring but may become very dry during summer droughts. The specific site studied is situated about 22 km southwest ofDeQuincy;(93' 34' 15" x 30** 18' 74");rangeand township (RllW x T8SSect.35)(Figure 1). Itisboundedtothesouth by the Houston Riverand to the westbyCreekRoad. Thecommunityis approximately2.5 km long on itseast-west axisand2.0 km longon its north-southaxis. Distinctiveherbaceous species include Spartina spartinae (Trin.) Hitch., Chaetopappa asleroides (Nutt.) DC., Evolvulus sericeus Sw., and Liatris punctata Hook. The understory woody vegetation is stunted. Thesoil typeis"Brimstone"andhas avery stronglyacidicsiltloam layer, a subsurfacethatismediumacidsiltloam,andasubsoilofalkalinesiltyclaythatis high insodium(Roy&Midkiff1988). Thethirdstudysiteis an acidicflatwoodsand is situated in thecentral partof the longleafpinelandregion,about10kmsoutheastofDeQuincy;(93** 20' 85" x 30** 22' 63");rangeand township (R9W x T7S Sect. 11) (Figure 1). This acidicflatwoods extends about 1.2 km on its north-south axis and about0.4 km on its east-westaxis and is bounded on the east by Hoibrook Park Rd. and to the west by cleared land. Thesoil type is "Caddo-Messersiltloam"and has avery strongly acidicsurface and subsurfacelayer(Roy&Midkiff1988). Thesubsoil isstronglyacidsiltyclayloamor siltloam. Theterrainisleveltogentlyrollingwithsmallelevatedareastermed"pimple mounds"byHolland^/a?. (1952). Under natural conditionsand frequentfires, these flatwoods support a sparse canopy of longleaf pine with few other tree species (Bridges& Orzell 1989). The plantingofpine plantationsand firesuppression have neariyeliminatedthesecommunitiesin the Parish. Distinctiveherbaceous species in this communityincludeStylisma aquatica (Walter) Raf. and Platantfieranivea (Nutt.) Luer. Toour knowledge, this is the only acidic flatwoods community in Calcasieu Parishthatsupportsarichherbaceouslayer. MATERIALSANDMETHODS Thethreerepresentativecommunitytypeschosenforthisstudysharethefollowing characteristics: 1) longleaf pine is a major component; 2) community maintenance requires frequent fires; 3) herbaceous ground cover is species rich. Each site was selected torepresentone ofthe three longleaf pine communities in Calcasieu Parish that shares these criteria. Although these sites are not pristine, we consider them & recognizable longleafpine communities(sensu Bridges Orzell 1989; Smith 1991, 1996). Eachsampledareameasured 100malongitslongaxisand36 m on its short axis. Datawerecollectedon treeandshrubspecies(vineswereexcluded) with adiameterat breastheight(DBH);(1.3 m)of 1 cmor greater. Randomly selected ten by ten meter quadrats(plots)wereusedforsamplingwithin thedefined study sites. Plotsize was basedon thatrecommendedfor forest trees by Cox (1996) and Brower etd. (1998). : Neylandetal. WoodyvegetationinCalcasieuParish 55 The numberofplots sampled persite was determinedby constructinga species area curvetoensureweencounteredthebulkofspecies present (Cox 1996). Specifically, aplotofcumulativenumberofspeciesencounteredversuscumulativenumberofplots sampled indicated fifteen plots was adequate to sample neariy all of the species present. Samplingforhillsideseepage bog, sodic fiatwoods and acid fiatwoods sites were performedon April8, May3, and May 20, 1998 respectively. Nomenclature followsKartesz(1994). Frequency,density,coverage(basalarea)andimportancevalueswerecalculatedto determinethe dominantwoody species (Cox 1996). Basal area was estimated from DBHvalues. Theimportancevalue is equal to the sum of relativefrequency, relative density,andrelativecoverage. Thesevalueswerecalculatedforeachof the threesites individually and for all sites combined. Species diversity was calculated using Simpson's DiversityIndex(Cox 1996)andwascalculatedfromdensitydata. Because we consider the few small "pimple mounds" to be "intrusions" into the acidic fiatwoods site, woody species were excluded from sampling. However, a complete inventoryofthewoody speciesinhabitingthese mounds was madeand is included in thefollowingsection. RESULTS Species area curves (Cox 1996) suggest that fifteen samples were sufficient to encounter the bulk ofspecies presentateach site. That is, the final plot number for eachsitewas positionedwellintotheasymptoticportionofthecurve. Woodyvegetationdiversityvaluesamongthecommunitiesdifferedsignificantly(p < 0.05) (Table 1). The low diversity value for the acidic fiatwoods site was due primarily to low species richness as compared to the other two communities. Although the hillside seepage bog and the sodic fiatwoods site each had identical species richness values (Table 1), the disparity in diversity values is explained primarilybygreaterevennessofspeciesabundanceatthesodicfiatwoods. Density, frequency, coverage, and importancevalues forall sites are summarized inTable2. Thevarious mixturesofwoody species sampled suggest a high degreeof heterogeneityamongthethreesites. Oftheelevendifferentspecies discovered among thesites, onlyFinnspaliistriswas presentinallthree. Myricacerifera L. was present in all three communities but no individuals metour size criterion for sampling in the acidic fiatwoods due to a recent burn. Additionally, seven species were present in only one of the three sites: P. elliottii. Ilex vomitoria Aiton, Quercus nmrilandica Muench, ViburnumdentatianL., /. coriacea(Pursh) Chapm., Uquidambarstyraciflua L., and Sapium sebiferum (L.) Roxb. (Table 2). Heterogeneity among the communities is also demonstrated by the importancevalues ofP. palustris. That is, although P. palustris was the most important species for all communities combined (Table3), it was onlv the second most importantspecies in the sodic fiatwoods site (Table2). PHYTOLOGIA 56 July 1998 volume85(l):51-60 Table 1. Simpson's Diversity Index, variance, and species richness of woody vegetationbystudysite. Site Neylande/a/.: WoodyvegetationinCalcasieuParish 57 Tables. Density,frequency,coverage, andimportancevaluesby species summarized forallsites. Speciesarearrangedinorderofimportance. Species PHYTOLOGIA 58 July 1998 volume85(l):51-60 criterion. Ofthesethreespeciesencountered,onlyM. ceriferaappearedtobe common onthesite. Thewoodyvine,Smilaxlaurifolia, wasalsopresent. A completeinventoryofthe woody species inhabiting the pimple mounds within the acidic flatwoods community includes Sapium sebiferum, Acer rubrum, Myrica cerifera. Sassafrasalbidum (Nutt.) Nees, Quercusfalcata Michx., Q. nigra L., Ilex opaca, andLiquidambarstyraciflua. DISCUSSION Although not pristine, the three remnant sites analyzed in this study resemble longleafpinecommunities before settlementand clearcutting. However, the current stateofthesepinelandsisundoubtedlydifferentfrom thetimeprior tosettlement. For example,themostimportantspecies found in thesodic flatwoods site, Piniiselliotlii, is native only in Louisiana parishes east of the Mississippi River (Rora of North AmericaCommittee 1993) and, therefore, would have been absent in preseltlement times. Althoughthiscommerciallyintroducedspecieshistoricallyhasbeenused forits naval stores, P. elliottii has become an increasingly important plantation pine for lumberandplywood(RoraofNorthAmericaCommittee 1993). The establishmentof P. elliottiihasalmostcertainlydisplacedothernativewoodyspecies. Stepshaverecentlybeentakentopreserve atleasta partof the sodic flatwoods in Calcasieu Parish. The Nature Conservancy of LxDuisiana has recently purchased approximately240 contiguous acres ofsodic flatwoods along Persimmon Gully that areadjacenttostudysitenumbertwo(RichardMartin,pers. comm., 1998). Theacidflatwoodsandhillsideseepagebog sites support afew individualsof the introduced Sapium sebiferum. This native of subtropical China, commonly called Chinesetallow,hasbeenplantedinseveralareasoftheworidasanornamentalandfor theproductionofvegetabletallow and stillingiaoil from its fruits (Kahn etd. 1973). This species was introduced to the Southeast by the U.S.D.A. Bureau of Plant Industryintheearly 1900's toestablish local soap industries (Jamieson & McKinney 1938). TheabilityofSapium toaggressively invadeand quickly alterhabitatis well documented(Cameron& Spencer 1989; Bruceetd. 1995; Neyland &. Meyer 1997). Atpresent, theincidenceofSapiumatthesetwositesis low. However, itremains an openquestiontowhatextentthis species will altertheecologyofthese communities. Additionally, species composition in the hillside seepage bog has been altered since settlementtimes by the introducedPinus elliottii. Specifically,all but the lowermost 36mofthehillsideseepagebognow supportsadensestandofP. elliottii. The future of these remnant longleaf pine communities in Calcasieu Parish is uncertain. Most remain threatened by urbanization, wildlife management practices, and silvaculture. Withoutactiveprotection, these rarecommunities may be destined forextinctioninCalcasieuParish. : Neylandetal. WoodyvegetationinCalcasieuParish 59 ACKNOWLEDGMENTS We thankLatimcweSmith from the National Heritage Program of the Lx>uisiana & DepartmentofWildlife Fisheriesforreviewingpreviousdraftsofthis paperand for providingusthelocationofthehillsideseepagebogstudysite. Wealso thank Lowell E. Urbatsch (Louisiana State University) and Mark Paulissen (McNeese State University)forreviewingthispaperpriortosubmission. UTERATURECITED Bridges, E.L. 1988. A preliminary survey for potential natural areas in the pine flatwoods region of southwestern Louisiana. Unpublished report for the LouisianaNaturalHeritageProgram, BatonRouge, Louisiana. & Bridges, E.L. S.L. Orzell. 1989. Longleaf pine communities of the West Gulf Coastal Plain. NaturalAreasJournal9:246-263. Brower, J.E. J.H. Zahr, & C.N. von Ende. 1998. FieldandLaboratoryMethods forGenera,lEcology, FourthEdition. Wm. C. Brown, Dubuque,Iowa. Bruce, K.A., G.N. Cameron, & P.A. Harcombe. 1995. Initiation of a new woodland typeon theTexas Coastal Prairie by the Chinese tallow tree (Sapium sebiferum(L.) Roxb.). Bull.TorreyBot. Club 122:215-225. Cameron, G.N, & R. Spencer. 1989. Rapid leafdecay and nutrient release in a Chinesetallowforest. Oecologia80:222-228. Cox, G.W. 1996. LaboratoryManualofGeneralEcology, Sixth Edition. 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