You are on page 1of 156

Ministry of Housing and the Environment

Proceedingsofthe1stResearchSymposiumon BiodiversityinTrinidadandTobago HeldinCommemorationoftheInternationalYear ofBiodiversity2010 PortofSpain Trinidad

Editors
AndrewLawrence HowardP.Nelson DepartmentofLifeSciences TheUniversityoftheWestIndies St.Augustine DepartmentofLifeSciences TheUniversityoftheWestIndies St.Augustine

Editors
AndrewLawrence DepartmentofLifeSciences TheUniversityoftheWestIndies St.Augustine Trinidad,WestIndies email:Andrew.Lawrence@sta.uwi.edu HowardP.Nelson DepartmentofLifeSciences TheUniversityoftheWestIndies St.Augustine Trinidad,WestIndies email:howien@hotmail.com

CoverArtwork:2011EleanorS.Devenish 2011 The University of the West Indies and The Ministry of Housing and the Environment, Government of the Republic of Trinidad and Tobago. All rights reserved.

Publishedby TheDepartmentofLifeSciences
TheUniversityoftheWestIndies St.Augustine Trinidad WestIndies Email:aabio.edulink@sta.uwi.edu Website:http://sta.uwi.edu/fsa/lifesciences/edulink/

ISBN:9789766202750

Copies of these Proceedings may be purchased from The Department of Life Sciences. An electronic version of these proceedings, searchable by author title and abstract, is posted on The Department of Life Sciences biodiversity programme website. Conference presentation proposals are screened by a committee composed of academic staff from the University of the West Indies, the Ministry of Housing and the Environment and invited reviewers. Accepted presenters are given guidelines for submitting their papers/posters for the proceedings.Papersmayberejectedbytheeditor(s)iftheyarenotsubmittedin ausableformatorbythedeadline.Researchpapersmayalsoberejectedifthe authordeclinesaneditorsrequesttomeetqualitystandards.Editorialdecisions aremadebytheeditor(s).

TableofContents

Foreword....................................................................................................vi Preface......................................................................................................viii ResearchPapers..........................................................................................1 HowmanyspeciesareinTrinidadandTobago? ChristopherK.Starr.....................................................................................2 PopulationDensityoftheCook'streeBoa(Corallus ruschenbergerii)intheCaroniSwamp,Trinidad. KeleTaylor,HowardP.Nelson,andAndrewLawrence..............................8 Demographyandgeneralecologyofanintroduced primatethetuftedcapuchin(Cebusapella)in Chaguaramas,Trinidad. DarshanNarang,HowardP.Nelson,andAndrewLawrence....................19 SpatialandTemporalDiversityinGroundLevelFruit FeedingButterflies. ImranKhan,ChristopherK.Starr,HowardP.Nelson, andAndrewLawrence...............................................................................30 Acomparisonofbeachmorphologyandphysical characteristicsofTurtleBeach,TobagoandGrande Riviere,Trinidadanditsimplicationsforturtlenesting. SheetalJankieandAndrewLawrence.......................................................42 SpatialDistributionofintertidalbenthicmacrofaunainthree sandybeachesinTrinidad. LanyaFanovich,HowardP.NelsonandAndrewLawrence.......................55 GeneticDiversityandStructureoftheNeotropical MonodominantSpeciesMoraexcelsa(Benth.)inFive NaturallyFragmentedPopulations. NigelAustin,MichaelOathamandPathmanathanUmaharan.................72 TheconservationstatusofMetastelmafreemani. GayatrilakshmiRaghavaSinghandMichaelOatham................................88

ii

TableofContents(cont)
FireintheAripoSavannasEnvironmentallySensitiveArea: CausesandConsequences. AditiBisramsinghandMichaelOatham....................................................99 BiodiversityandbiogeographyoflichensinTrinidadand theimplicationsforforesthealthandbiosensitivity. AndreaScobie.........................................................................................110 ElectricalEnhancementofCoralGrowth:APilotStudy L.S.Beddoe,T.J.Goreau,J.B.R.Agard,M.George,and D.A.T.Phillip............................................................................................116 PopulationdensityoftheagoutiDasyproctaleporinaatthe CentralRangeWildlifeSanctuary,Trinidad. HowardP.Nelson,IndiraOmahMaharaj,NadraNathaiGyan, andAntonyRamnarine............................................................................123 Abstracts.................................................................................................130 NaturalHistoryandConservationoftheTrinidadPipingGuan KerrieNaranjit.........................................................................................131 BiofoulingonrecreationalvesselsinTrinidadandTobago JudithF.Gobin,AlanaJuteandAnuradhaSingh.....................................132 PatternsofbiodiversityinTrinidadianspiders. JoanneSewlal..........................................................................................133 Aninitialinvestigationintothethirdrecordedmassbleaching eventinTobago. JahsonAlehmu........................................................................................134 SpatialDistributionandextentofmangrovesinTrinidad. RahannaJumanandDeaneshRamsewak...............................................135 Developingpublicawarenessandeducationtoolstopromote anunderstandingandappreciationofbiodiversityinthe coastalandmarineenvironment. LoriLeeLum............................................................................................136

iii

TableofContents(cont)
Aninnovativeapproachformonitoringabioticfactors influencingmangroveforestbiodiversityinanestuarine ecosystem. M.Atwell,M.Wuddivira,J.Gobin,andD.Robinson................137 Monitoringandmanagementofmarineinvasivealien speciesinTrinidadandTobago. RosemarieKishore,FrancisWeekesandKhamaPhilip.............138 TheextentoftheseaturtlefisheryinTobago,WestIndies. MichelleCazabonMannette.....................................................139 Mitigatingathreatofinvasivealienspeciesintheinsular CaribbeanATrinidadandTobagoPerspective. VeldaFergusonDewsbury........................................................140 Lifeanddeathinthesavannasastudyoftherare terrestrialorchidCyrtopodiumparviflorum. HowardP.Nelson,SharonLaurent,CarlysleMcMillan andEleanorDevenishNelson..................................................141

iv

Foreword
In 1992, after the Earth Summit in Rio, the Convention on Biological Diversity was established and opened for signature. It was at that occasion the Government of the Republic of Trinidad and Tobago took the opportunity to become a signatory to the Convention, which was subsequently ratified in 1996. By so doing, Trinidad and Tobago had signalled to the international community that this country was aligned with international efforts aimed at reducing biodiversity loss, and the equitable sharing of the benefits arising from the wise use of biological resources. The United Nations has declared 2010 as the International Year of Biodiversity, with the theme Biodiversity is life, biodiversity is our life. ThisthemeisveryrelevanttoTrinidadandTobago.Oneoftheobjectives of the International Year of Biodiversity is the mainstreaming of biodiversity issues in public policy, so as to sensitize and raise public awarenessofecologicalissues,andtoensurethatthoseissuesaretaken on board in the implementation of environmental agendas. In my GovernmentsPolicyFrameworkforSustainableDevelopment,ourfocus is,ManagingourEnvironment,becauseourunderstandingisthat,there canbenosustainabledevelopmentwithoutrespectfortheenvironment. Thepolicyframeworkfortheenvironmentthereforeseekstostrengthen andpromoteeffortsatnatureconservation,inparticulartheconservation ofbiodiversityregimes. Being a tropical Caribbean island in close proximity to the Equatorial region, our country has been greatly endowed with a rich and unique biodiversity,whichhasinlargemeasurecontributedtooureconomicand social development. This development has been facilitated by the available knowledge of our natural resources, together with evidence baseddecisionmakingwithrespecttotheuseofthoseresources.Itisin thiscontextthatthecollaborationbetweentheLifeSciencesDepartment oftheUniversityoftheWestIndiesandtheMinistryofHousingandthe Environment for the conduct of the Research and Poster and Paper Symposium in commemoration of the International Year of Biodiversity has generated value to Trinidad and Tobago, and by extension, the internationalcommunity. BiodiversityresearchconductedbytheUniversityoftheWestIndieshas always been a valuable service, in terms of quantification of species for

management, and the identification of causes and development of solutionsforproblemsoccurringinourecosystems.However,asanation with an emerging economy, there is need to emphasize that more and continuousresearchisnecessaryforustokeepabreastofthecountrys developmentalneeds. Tertiarylevelresearch,asseeninthecollaborativeworkbetweentheLife Sciences Department of the University of the West Indies and the Ministry of Housing and the Environment, is therefore an ideal opportunity which allows post graduate students of the University to creatively communicate their research work in a format that would be accessibletoallsectorsofthepublic,whileatthesametime,providing valuableexposuretothesalientbiodiversityissueswhichmattertoallof us. Tertiary level research also identifies gaps and shortcomings that policy makers and planners need to address within the policy and planningframework.Forthesereasonstherefore,andinmycapacityas the Minister of Housing and the Environment, I am committed to promoting and supporting biological research among post graduate studentsattheUniversityoftheWestIndies, ThehostingoftheResearchPosterandPaperSymposiumwasintended to contribute to this effort and reinforces the linkage between my MinistryandtheLifeSciencesDepartmentoftheUniversityoftheWest Indies.Together,weshouldlookforwardtotheSymposiumbecominga fixtureonthecalendarsofbothinstitutions,andexpectanincreaseinthe numberofextractstobepresentedinthesuccessiveyears.Astheevent grows,wecancometoexpectagreatermomentumandhigherlevelsof interestandparticipationfromthevariouspublics,suchthatbiodiversity protectionandconservationbecomeanintegralpartofourexistenceand ethos. Dr.TheHonourableRoodalMoonilal MinisterofHousingandtheEnvironment May,2011

vi

Preface
The 2010 International Year of Biodiversity (IYB 2010) has provided the world with a unique opportunity to take a critical look at the state of biodiversity resources at a global level. The emerging picture is not encouraging,astherecentreflectionsattheConventiononBiodiversitys ConferenceofthePartiessuggest,wehavenotdonesufficientgloballyto stem thetideof biodiversity loss. This inability to slowdown or reverse therateofbiodiversitydegradationhastremendousimplicationsforthe quality of human life in the future. Nowhere is this link between biodiversitys goods and services and the quality of human life more relevant than on Neotropical islands like Trinidad and Tobago, where issuesasdiverseasrurallivelihoods,potablewatersupply,foodsecurity and resiliency to climate change and natural disasters are inextricably linkedwiththestateofthecountryslivingresources. One key element of any attempt to stem the loss of biodiversity in Trinidad and Tobago mustbe the development of stronger mechanisms for information exchange between those individuals and institutions involved in biodiversity research, and those agencies involved in policy making and management of these resources. This need for better communicationbetweenthesegroupswasidentifiedasacriticalissuein thecountrysNationalBiodiversityStrategyandActionPlan,andremains an important challenge to improving biodiversity management in the country. Only through directed research on biodiversity can the gaps in our knowledge on the living elements of Trinidad and Tobago be filled and so allow for development of scientifically justifiable, costeffective and culturally relevant interventions to manage the countrys living resources. The IYOB 2010 Research and Poster Symposium has provided a unique opportunity to develop a bridge between the research community and the policy and management community in Trinidad and Tobago. The papersandposterspresentedattheSymposiumreflectworkonadiverse range of taxonomic groups, ecosystems and management questions currently being investigated by the research institutions operating in Trinidad and Tobago. These proceedings contains 11 abstracts and 12 papers based on the posters and papers presented at the International Year of Biodiversity Symposium held on 25th November 2010, at the CrownePlazaHotel,PortofSpain,Trinidad.Acceptedabstractandposter presenters/authors who participated in the Symposium were invited to

vii

submit full papers for inclusion in these proceedings. Those full papers presented here represent those presenters/authors who agreed to submitfullpapersandhavemettheeditorialstandardsetbytheeditors. WewouldliketoclosebyrecognizingourcolleagueDr.DavidI.Persaud, whose vision and support for a stronger link between the work of the national and regional research institutions and their policy and management agency counterparts in Trinidad and Tobago provided us withthemeanstomaketheSymposiumareality. ProfessorAndrewLawrenceandDr.HowardP.Nelson St.Augustine, March,2011

viii

ResearchPapers

HowmanyspeciesareinTrinidadandTobago? ChristopherK.Starr
DepartmentofLifeSciences,TheUniversityoftheWestIndies,St.Augustine,Trinidad,West Indies.Email:christopher.starr@sta.uwi.edu

Abstract KnowledgeofthenumberofspeciesoforganismsnativetoTrinidadand Tobago varies widely among taxa. At one extreme, represented by the microorganisms,solittleisknownthatnomeaningfulestimatesareyet feasible.Attheoppositeextreme,representedbylandandfreshwater vertebrates,specieslevelinventoriesarevirtuallycomplete.Insectsmake upabouthalfofthedescribedspeciesofallorganismsonEarth,andthey certainlymakeupthebulkofanimalspecies.Aboutonemillionspecies havebeendescribed,andestimatesofthetruetotalconvergearound2.5 to6million.ApplyingtothisrangetheTrinidadandTobagofractionsof the world fauna in nine wellstudied groups, the true total for these islandsisestimatedtobebetween67.5and312thousandspecies. Keywords Invertebrate,speciesrichness,Tobago,Trinidad,vertebrate,diversity Introduction Itcanreasonablybeassumedthatmostreadersofthispaperareinfull sympathy with the view that humanity is not the owner but rather the caretaker of the Earth's biotic resources. In addition, it can be argued thathumanitycandoabetterjobofcaringforthemifitisknownwhat they are. The worldwide enterprise of inventorying the world's species, then, is a key foundation of any conservation effort. In this regard, nationalboundariesareaconvenientifhighlyartificialwayofdividing theplanetintomoremanageableunits,sothatinventoriesareusuallyon a countrybycountry basis. The purpose here is to review the state of knowledge of the specieslevel biotic diversity of one such country, TrinidadandTobago. In the standard classification, the phyla of living organisms are arranged into five kingdoms (e.g. Margulis 1998). Three of these Monera, bacteria and other prokaryotes; Protista, protozoans and most algae;andFungiaretreatedcollectivelyas"microorganisms",although this does not imply that together they form a natural, monophyletic

group. The other two kingdoms are the plants (Plantae) and animals (Animalia). Microorganisms The number of species of microorganisms in Trinidad and Tobago remains unknown. About 157,000 species have been described worldwide(HawksworthandKalinArroyo1995),butthisisplainlyagross underestimate of the true total. It is quite certain that the described speciesineachofthethreekingdomsrepresentonlyasmallfractionof those in existence. For Trinidad and Tobago, in particular, even less is known. Over the course of the next generation, it can be hoped that a breakthrough in the assessment of microbial biodiversity will be achieved. Plants ThesituationofthePlantKingdomisverynearlytheoppositeofthatof microorganisms. The vascular plants, and in particular the seed plants, makeupthegreatbulkofdescribedandofallexistingspecies.Theseare relatively well catalogued worldwide, as well as in Trinidad and Tobago (BakshComeauetal.,inpress).Thedescribedspeciesofvascularplants herenumberabout2465:270ferns,25fernallies,1431dicotyledonous flowering plants, and 739 monocotyledonous flowering plants. These figuresarebelievedtobeclosetothetruetotals. Animals For convenience, attention will be restricted to land and freshwater species. The waters around Trinidad and Tobago harbour a richness of animals, but there is no biologically meaningful way of drawing boundaries. Vertebrates Thevertebratesappearevenmorecompletelyknownatthespecieslevel than the vascular plants (Table 1). Further close examination will undoubtedly modify these numbers slightly, but they are unlikely to significantlydisturbthepresenttotalofabout600species. Insects Insects appear to be by far the most speciose group of organisms on Earth, with slightly over one million described species worldwide. Nonetheless, this is evidently just a fraction of the true total. Furthermore, confidence in the reliability of present numbers varies

widely among orders (Nielsen and Mound 1997: Table 2). The various attemptstoestimatethenumberofexistingspecies(Table2)applyvery differentmethods,butallrelyonacomparablemixtureofknownfacts, ratiosandexplicitassumptions.

Table 1 Numbers of land and freshwater vertebrate species in Trinidad and Tobago. VertebrateGroup Species References
_____________________________________________________________

Fish Amphibians Reptiles Birds Mammals Total

42 32 106 321 98 599

PhillipandRamnarine(2001) Murphy(1997) Boos(2001),Murphy(1997) ffrench(1991) Boos(unpubl.)

Toillustratethisgeneralapproach,therelativelysimplemethod ofHodkinsonandCasson(1991)canbesummarized.Theseauthorsare specialists in Hemiptera, an insect order with about 71,000 described species worldwide. In a set of large samples collected by insecticidal fogging in Indonesia, they found 1690 species of Hemiptera. Of these, 62.5%werepreviouslyundescribed.The71,000describedspecies,then, were estimated to be just 37.5% of the true total. Hemiptera make up 7.5% of described insect species. Assuming this ratio also holds for undescribedinsectspecies,wereachanestimateof71,000X1/0.375X 1/0.075=2.5millionspeciesofinsectsintheworld. There is no strong reason to favour any one of the estimates given in Table 2. Nonetheless, it is remarkable that with one exception theyfallwithintherangeof2.5to6millionspecies.Giventhevarietyof methods, it is a reasonable working hypothesis that the true total falls withinthismanageablerange. Noattempthasyetbeenmadetoestimatethenumberofinsect species occurring in Trinidad and Tobago. Even at this small scale, no directcensus approach will yield a meaningful answer. However, it is possible to reach a reasonable rough estimate by reference to known numbersofspeciesinthefewwellstudiedgroupsina)theworld,andb) Trinidad and Tobago, together with an estimate of the total number of speciesintheworld.AsseeninTable3,TrinidadandTobago'sshareof the world fauna in nine wellstudied groups ranges from 2.2% to 7.8%, withallbutthreefiguresfallingbetween2.7%and5.2%.

Table 2 Estimates of the number of insect species in the world. _______________________________________________________ Estimate(millions) Reference _______________________________________________________ 2.5 HodkinsonandCasson1991 2.7 Gaston1992 4 May1994 4.8 degaard2000 4.86 Novotnyetal.2002 5.7 Hammond1992 6 Groombridge1992:2425 30 Erwin1982 _______________________________________________________

Iftheworkinghypothesisisadoptedthattheoverallfractionforinsects as a whole is within this range we arrive at the following estimates of theTrinidadandTobagofauna: Minimum2.7%of2.5million,or67,500. Median4.5%of4.8million,or230,400. Maximum5.2%of6million,or312,000. This is almost a fivefold range, certainly far from a precise estimate. However, it is a fair beginning and a significant improvement over the lack of any previous estimate. Furthermore, it serves to emphasize that insects comprise the great majority of multicellular species under our nationaljurisdictiononland.
Table 3 Numbers of known insect species in the world and in Trinidad and Tobago. Taxon Knownspecies FractionReferences World TandT inTandT Odonata 5500 119 2.2% 14 Isoptera 2100 56 2.7% 56 Heteroptera:Gerromorpha 2400 78 3.3% 78 andNepomorpha 64 5 7.8% 910 Heteroptera:Dysdercus Lepidoptera:Papilionoidea 13,688 387 2.8% 1112 Lepidoptera:Hesperiidae 3592 272 7.6% 11,13 Diptera:Culicidae 3209 160 5.0% 14,15 Hymenoptera:socialVespidae 800 38 4.8% 1619 Hymenoptera: DorylinaeandEcitoninae 248 13 5.2% 2021

References: 1.Bridges(1993).2.Geiskes(1932).3.Geiskes(1946).4.Michalski(1988). 5.Constantino(2010).6.Scheffrahnetal.(2003).7.SlaterandO'Donnell(1995).8.Nieser and AlkinsKoo (1991). 9. Freeman (1947). 10. Doesburg (1968). 11. Shields (1989). 12. Barcant(1970).13.Cock(1982).14.Ward(1992).15.Tikasingh(unpubl.).16.Carpenter (1996).17.CarpenterandKojima(1996).18.MatsuuraandYamane(1984).19.Starrand Hook(2003).20.Bolton(1995).21.Watkins(1992).

References
BakshComeau,Y.S.,D.W.Hawthorne,S.A.Harris,S.S.MaharajandD.L.Filer.InPress.The VascularFloraofTrinidadandTobago:AChecklistandConservationStatus. Barcant,M.1970.ButterfliesofTrinidadandTobago.London:Collins314pp. Boos,H.E.A.Unpubl.ChecklistofmammalsofTrinidadandTobago. Boos, H.E.A. 2001. The Snakes of Trinidad and Tobago. College Station: Texas A and M Univ.Press270pp. Bolton,B.1995.ANewGeneralCatalogueoftheAntsoftheWorld.Cambridge:Harvard Univ.Press504pp. Bridges,C.A.1993.CatalogueoftheFamilyGroup,GenusGroupandSpeciesGroupNames oftheOdonataoftheWorld.2nded.Urbana:Publ.byauthor. Carpenter, J.M. 1996. Phylogeny and biogeography of Polistes. Pp. 1857 in: S. Turillazzi and M.J. WestEberhard (eds.), Natural History and Evolution of PaperWasps. Oxford: OxfordUniv.Press. Carpenter,J.M.andJ.Kojima1996.ChecklistofthespeciesinthesubfamilyStenogastrinae (Hymenoptera:Vespidae).J.NewYorkent.Soc.104:2136. Cock,M.J.W.1982.Theskipperbutterflies(Hesperiidae)ofTrinidad.PartII.Asystematiclist of the Trinidad and Tobago Hesperiidae. Occ. Pap. Dept. Zool. Univ. West Indies, St. Augustine(5):47pp. Constantino, R. 2010. Online catalog of the termites of the New World. http://www.unb.br/ib/zoo/docente/constant/catal/cat.htm. Doesburg, P.H. van 1968. A revision of the New World species of Dysdercus Gurin Mneville(Heteroptera,Phyrrhocoridae).Leiden:E.J.Brill215pp. Erwin, T.L. 1982. Tropical forests: their richness in Coleoptera and other arthropods. Coleopt.Bull.36:7475. ffrench,R.1991.AGuidetotheBirdsofTrinidadandTobago.2nded.Ithaca:CornellUniv. Press426pp. Freeman,P.1947.ArevisionofthegenusDysdercusBiosduval(Hemiptera,Pyrrhocoridae), excludingtheAmericanspecies.Trans.r.ent.Soc.London98:373424. Gaston,K.J.1992.Regionalnumbersofinsectandplantspecies.FunctionalEcol.6:24347. Geiskes, D.C. 1932. The dragonflyfauna of Trinidad in the British West Indies (Odonata). Zool.Meded.14:23262,15:96128. Geiskes, D.C. 1946. Observations on the Odonata of Tobago, B.W.I. Trans. r. ent. Soc. London97:21335. Groombridge, B. (ed.) 1992. Global Biodiversity: Status of the Earth's Living Resources. London:ChapmanandHall585pp. Hammond, P. 1992. Species inventory. Pp. 1739 in: B. Groombridge (ed.), Global Biodiversity:StatusoftheEarth'sLivingResources.London:ChapmanandHall. Hawksworth,D.L.andM.T.KalinArroyo1995.Magnitudeanddistributionofbiodiversity. Pp. 10791 in: V.H. Heywood (ed.), Global Biodiversity Assessment. Cambridge: CambridgeUniv.Press. Hodkinson, I.D. and D. Casson 1991. A lesser predilection for bugs: Hemiptera (Insecta) diversityintropicalrainforests.Biol.J.linn.Soc.43:10109.

Margulis,L.1998.FiveKingdoms:AnIllustratedGuidetothePhylaofLifeonEarth.3rded. NewYork:W.H.Freeman520pp. Matsuura,M.andSk.Yamane1984.BiologyoftheVespineWasps.Berlin:Springer323 pp. May, R.M. 1994. Past efforts and future prospects towards understanding how many species there are. Pp. 7184 in: O.T. Solbrig, H.M. van Emden and P.G.W.J. van Oordt (eds.),BiodiversityandGlobalChange.Wallingford,Oxon:CABInternational. Michalski, J. 1988. A catalogue and guide to the dragonflies of Trinidad (order Odonata). Occ.Pap.Zool.Dep'tUWI,StAugustine(6):1146. Murphy, J.C. 1997. Amphibians and Reptiles of Trinidad and Tobago. Malabar, Florida: Krieger245pp. Nielsen,E.S.andL.A.Mound1997.Globaldiversityofinsects:Theproblemsofestimating numbers. Pp. 21322 in: P.H. Raven (ed.), Nature and Human Society: The Quest for a SustainableWorld.Washington:NationalResearchCouncil. Nieser,N.andAlkinsKoo,M.1991.ThewaterbugsofTrinidadandTobago.Occ.Pap.Zool. Dep'tUWI,StAugustine(9):1127. Novotny,V.,Y.Basset,S.E.Miller,G.D.Weiblen,B.Bremer,L.CizekandP.Drozd2002.Low hostspecificityofherbivorousinsectsinatropicalforest.Nature416:84144. degaard,F.2000.Howmanyspeciesofarthropods?Erwin'sestimaterevised.Biol.J.linn. Soc.71:58397. Phillip,D.A.T.andI.W.Ramnarine2001.AGuidetotheFreshwaterFishesofTrinidadand Tobago.StAugustine:Dep'tofLifeSciences,Univ.oftheWestIndies79pp. Scheffrahn, R.H., J. Krecek, B. Maharajh, J.A. Chase, J.R. Mangold and C.K. Starr 2003. Termite fauna (Isoptera) of Trinidad and Tobago, West Indies. Occ. Pap. Dep't Life Sci. Univ.WestIndies(12):3338. Shields,O.1989.Worldnumbersofbutterflies.J.Lepid.Soc.43:17883. Slater,J.A.andJ.E.O'Donnell1995.ACatalogueoftheLygaeidaeoftheWorld(19601994). NewYork:NewYorkEntomologicalSoc.410pp.(Modifiedinlinewithpers.comm.from J.E.O'Donnell.) Starr,C.K.andA.W.Hook2003.TheaculeateHymenopteraofTrinidad,WestIndies.Occ. Pap. Dep't Life Sci. Univ. West Indies (12):131. Updated version: http://www.ckstarr.net/aculeates.pdf. E.Tikasingh.Pers.comm. Ward,R.A.1992.Thirdsupplementto"Acatalogofthemosquitoesoftheworld"(Diptera: Culicidae).MosquitoSyst.24:177230. Watkins, J.F. 1992. Ecitoninae: Distribution of New World army ants by genus: species: country:state.Unpubl.list.

PopulationDensityoftheCook'streeBoa(Corallus ruschenbergerii)intheCaroniSwamp,Trinidad.

KeleTaylor HowardP.Nelson2,andAndrewLawrence2
1,2 2

DepartmentofLifeSciences,TheUniversityoftheWestIndies,St.Augustine,Trinidad, WestIndies.Email:keletaylor@hotmail.com 1 CorrespondingAuthor

Abstract Themonitoringofanimalpopulationsisnecessarytoprovidethebasisfor understanding population changes over time, as well as to conserve, protect and manage a species. Corallus ruschenbergerii, commonly known as the Cooks Tree Boa, is found in the Caroni Swamp, Trinidad. The aim of this study was to estimate the population density of the Cooks tree boa in the Caroni Swamp, Trinidad, using line transect distancesurveys.Thesurveywasconductedduringthewetseasonperiod for Trinidad, i.e. the months of September and October 2010 along six transects.Atotalof115.97kmoftransectsweresurveyedviaboatwithin the swamp channels and drainage outlets, as well as on foot along the banksofthemangrove.Distance6.0wasusedtoestimatedensityofthe Cookstreeboa,basedontransectobservations.Atotalof32Cookstree boas were observed during this study. The best fitting model for the surveydata,basedontheAkaikeinformationcriterion(AIC)score,wasa uniformkeyfunctionwithsimplepolynomialexpansion.Cookstreeboa densityestimateswereestimatedat11.022treeboasperkm2. Keywords Cooks tree boa, Caroni Swamp, Trinidad, population density estimate, linetransectsurveys,distancesampling. Introduction Urban expansion, pollution and deforestation have resulted in the dramaticdecreaseintheabundanceofmanysnakesglobally(Mullinand Seigel 2009). In addition, snakes may be considered among the most persecuted organisms, with public attitudes towards them ranging from fascination, awe and worship, to fear and loathing (Mullin and Seigel 2009).Thislackofempathytowardsthesevertebratesissurprising,given that less than 25% of these species are venomous and most perform importanttopdownregulationofmammalianpestspecies.

Given these challenges, thedevelopmentof conservation strategies that maintain the role of snakes as indicators of ecosystem health, and important predators, is imperative (Mullin and Seigel 2009). Whilst numerous studies on the ecology and natural history of many snakes have been conducted, a complete understanding of their ecology, and particularly population biology, remains lacking (Dorcas and Wilson 2009). Such gaps in knowledge limit the ability of managers to develop effective conservation and management strategies. Accurate population estimatesforsnakescanbedifficulttoobtainduetoproblemsassociated withlowdetectabilityandbiasedsamplingmethods(DorcasandWilson 2009). The population status of the Cooks tree boa in the Caroni Swamp, andontheentireisland,isunknown,astherehavebeenfewsystematic snake surveys done in Trinidad. The Cooks tree boa (Corallus ruschenbergerii)belongstothesnakefamilyBoidae,whichcontainssome of the largest living snakes in the world. Its range includes Trinidad and Tobago,NicaraguatonorthernColombia,andnorthernVenezuela.Local names include common tree boa, cascabel dormillon, yellowmarbled tree boa, mangrove cascabel and mangrove dormillon. The Cooks tree boa can be found in the Caroni Swamp, which is located on the west coastofTrinidad.Cookstreeboahasalsobeenrecordedfromthenorth coastofTrinidad,inforestedareasbetweenPariaandPetiteTacaribBays (Boos 2001). It is a mainly nocturnal and arboreal snake, with colours ranging from khaki green or brown, and yellow beneath the chin, neck and fore body. During the day, individuals can be seen coiled in a tight ballintrees,oftenoverhangingwaterandtheirpreyincludesfrogs,bats, birds and iguanas (Wehekind 1955; Hendserson and Boos 2001). Although the Cooks tree boa is nonvenomous, they can be extremely aggressiveanddefensivewhendisturbed. Reliable census techniques and accurate assessments of tree boa densities are fundamental to wildlife research and future monitoring of populations. A density estimate of the Cooks tree boa is crucial to the understanding of the temporal dynamics of its population and for evaluating the effectiveness of any management strategies used to conservethespecies.Itwouldbeimportanttodeterminethepopulation of existing Cooks tree boas in the Caroni Swamp as the area has been undergoingchanges,whichcandirectlyaffectthesnakepopulation.The Caroni Swamp is experiencing pressures such as reclamation for roads, housing and industrial development, industrial and chemical pollution, poaching, unmanaged and uncontrolled tourism and drainage modification(ForestryDepartment,FoodandAgricultureOrganizationof

theUnitedNations2005).Inaddition,thereisgeneralpublicindifference to the protection and conservation of Cooks tree boa in the Caroni Swamp, and the ecological system as a whole. Snakes have lifehistory characteristicsthatmakethemvulnerabletopopulationdeclines,suchas long life spans, late sexual maturity and low reproductive rates (Mullin and Seigel 2009). This, coupled with the fact that many snake species occupy the highest levels in their respective trophic webs, suggest that anydeclineinsnakepopulationsarelikelytohaveimpactsontheirprey populations,andforthewetlandecosystemasawhole(MullinandSeigel 2009). WithoutquantitativeassessmentoftheCookstreeboapopulation, the conservation and management of this species cannot be implemented in the long term. As such, this paper describes the application of a distance sampling survey to estimate the population densityoftheCookstreeboaintheCaroniSwamp. MaterialsandMethods StudyArea This study was conducted in the Caroni Swamp, Trinidad, which has an areaofapproximately6000ha,makingitthelargestwetlandonthewest coast of the island. The swamp is situated between PortofSpain and Chaguanas, where the Caroni River meets the Gulf of Paria. The Caroni River,themainwatercourseenteringtheswamp,runsalongthenorthof theswamp.OthernaturalwatercoursesoftheswampincludetheBlue River,CatfishRiver,PhaggRiver,GuayamareRiverandMadameEspagnol River. There are many canals and channels within the swamp, some of whicharenaturalandothersdredged. TheCaroniSwampispredominantlyestuarineandischaracterizedby dense mangrove vegetation, reaching some 23 m in height. The swamp holds the largest single stand of mangrove on Trinidads west coast, which includes approximately 60 % of Trinidads mangroves. This ecosystem includes Rhizophora mangle, Rhizophora harrisonii, Rhizophora racemosa, Avicennia germinans, Avicennia schaueriana, LagunculariaracemosaandConocarpuserectus.Themostwidespreadis Rhizophoramangle,followedbyR.harrisoniiandR.racemosawhichare also very common (Forestry Department, Food and Agriculture Organization of the United Nations 2005). The climate at this site is tropical humid, with two distinct, wet and dry, seasons. The Caroni Swampwasrecognizednationallyasanenvironmentallysensitivesystem, andtodaymostoftheCaroniSwampisaprohibitedareawithentryby

10

permit only. The site was also designateda Ramsar siteof international importancein2005. DistanceSamplingSurveys Population density data for many terrestrial species can often be estimated using line transect surveys (Hyrenbach et al. 2001). This technique uses counts of individuals and is widely used in animal populationecologyfordensityestimation(Plumptre2000;Marquesetal. 2007). Such line transect surveys can be relatively effective for the estimation of animal densities over large areas or where budgets are limited(Neweyetal.2003). ThisstudyusedlinetransectsurveysforCookstreeboaintheCaroni Swamp, as a means to determine population density of this predator. Distancesamplingiswidelyusedinwildlifeecology(Burnhametal.1980) and its advantages include flexibility in data collection and processing, andprovisionofhighlypreciseestimatesofanimaldensities(Harrisand Burnham2002).Inthisdistancesamplingstudy,itwasassumedthatall Cooks tree boas were not detected during the surveys due to visual obstructionsandobservererror. Distancesamplingusessampledataonspeciesdetectionprobability to estimate a detection function that describes how detection of the specieschangeswithincreasingdistancefromthetransect(Somershoeet al.2006).Theareaaroundthetransectcanbederivedfromthisfunction, anddensityisthencomputedasthenumberofindividualsencountered dividedbytheeffectiveareasampled. Distance sampling makes three critical assumptions (Newey et al. 2003, Buckland, et al. 2001): First, all individuals on the transect are detectedbytheobserver.Thisassumptionaffectstheconstructionofthe detection function, and importantly, it relies on the 100% detection of individualsonorveryclosetothesampleline.Asecondkeyassumption isthatindividualsdonotmovebeforedetection.Distancesrecordedafter any movement by individuals under observation will introduce errors in densityestimation;andthird,isthatdistancemeasurementstothetarget animalareaccurate.Again,anybiasinthemeasurementswillintroducea biasintheestimateofindividualdensities. The distance surveys undertaken here were conducted through the months of September and October 2010, during the morning period to midday period (0700 1300) and evening period (1400 1900). Surveys were conducted on six transects, along the channels, drainage outlets and mangrove embankments throughout the swamp. Transect lines rangedfrom18kmto22kminlength(Figure1).Atransectwidthof50

11

m was estimated on either side of the transect axis. All surveys were conductedwithanaverageoffiveobservers.Surveyswereundertakenin aflatbottomboatonopenwater,withobserversonbothsidesandfront of the boat, and on foot in single file while traversing transects along mangrovebanks.WhenCookstreeboaswereobserved,theirpositions were recorded via handheld GPS receivers. Sighting distance in meters was measured by eye from the transect to the Cooks tree boa, and compass bearings were taken to determine the sighting angle between the Cookstree boa and the transect, by the same observer throughout thewholestudy.Theradialdistancesandsightingangleswerethenlater converted to perpendicular distances (Buckland et al. 2001). Environmental data on viewing conditions such as weather and glare werealsorecorded. Dataanalysis All field data were analysed using the conventional distance sampling (CDS) engine in the software Distance 6.0 (Thomas et al. 2004). This programusesanarrayofmodelsforestimationofdetectionfunctionand populationdensity.Threedetectionfunctions(halfnormal,uniformand hazard rate), and their series expansions (simple polynomial, cosine or hermitepolynomial),wereusedinthisanalysis.Exploratoryanalysisand modelselectionfollowedthoseguidelinesrecommendedbyBucklandet al.(2001).Exploratoryanalysisincludedplottingofahistogramtoallow examination of observationfrequency and distance relationships. Models were compared using the AIC scores, and the model with the lowestAICvalueselected(BurnhamandAnderson1998). Results Atotalof32Cookstreeboaswereobservedfromatotalof115.97km line transect surveys in the Caroni Swamp. Buckland et al. (2001) recommends data truncation to eliminate outliers and improve model fitting. However, there were no observations remarkably different in measurementandthusnotruncationwasmadeintheanalysisconducted in Distance 6.0. The probability of detecting the Cooks tree boas decreasedasthedistancefromthetransectlineincreased.Basedonthis datastructure,thehalfnormalandhazardratemodelswereconsidered because they are known to be able to manage data with rapid falls in detectionrateswithdistance(Bucklandetal.2001).

12


Figure1Studyareaandlocationofsurveyroutes(dottedlines)andallCookstree boaobservedwithintheCaroniSwamp(greydots)

Using the AIC values, the halfnormal simple polynomial model provided the best fit to the data. The density of the Cooks tree boa estimatedfromdistancesamplinganalysisusingthelowestAICvaluewas 11.022 tree boas per square kilometre. The histogram of perpendicular sightingdistanceshadanarrowshoulderthatsuggestedmanyofthetree boas were detected on the transect line, and that detection rates decreasedrapidlywithperpendiculardistance(Figure2).
Figure 2 Histogram of perpendicular distances and the halfnormal simple polynomial density estimator model for Cooks tree boa at Caroni swamp, Trinidad.

13

Discussion Distance sampling methods are often constrained by the reality of field conditions(Bucklandetal.2001).Ideallylinetransectsshouldbeplaced randomly, and subsequent transects evenly spaced across the swamp regardless of the distribution of habitat type(s) (Kuhl et al. 2008). The major problems of distance sampling the Cooks tree boa population in thisarea,wastheuseofnonstraighttransectsalongthechannels,canals andembankmentswithintheswamp.Thisdoesnotconstitutearandom sample and possibly gave a biased estimate of the population density (Bucklandetal.2001;HibyandKrishna2001). Surveysonfootwereconductedalongrouteswhereitwaspossible towalk.However,therewerefewareaswherethiswaspossible,asmost of the landscape within the study area was covered by water and mangroveroots.Theboatsurveyspermittedalargersamplesizetobe obtainedandalargerareatobecovered,aswellasallowingtwoviewsof thetransectswhilegoinginoppositedirections.Inaddition,althoughthe transect lines were not straight, the estimate may have been more precise because more of the sample site was covered (Plumptre 2000). Also, even though it was difficult to establish straight transect lines throughout all of the study area, transects covering more of the study areamadetheserepresentativeasfaraswaspossible,oftheentirearea. Moreover, since most of the surveys were conducted from a small slowmovingboat,Cookstreeboadetectionwaslikelytovarywiththe sizeoftheboatandheightofobserverabovewaterlevelandboatspeed. Boat size, especially, has implications regarding navigation, as a smaller flat bottom boat canpass through narrower channels as comparedtoa larger boat. Speed is also an important factor in detection; especially if observershavetosightCookstreeboasindensevegetation. Other environmental factors such as weather and sun glare affect detection of Cooks tree boas. Surveys were conducted during the morning period when glare was not a problem, and for those surveys conducted in the evening period when there was some glare present, observers were able to navigate transects oneway to count individuals ononesideoftheboatwithoutglare;andthenobserve/sightindividuals ontheothersideoftheboatwhenreturningintheoppositedirection. Theaccuracyofthiskindofsurveyislimitedduetotheconstraints imposedbytheassumptionsofdistancesampling.Thisisespeciallytrue ofobservationsonorclosetotheaxisofthetransect(Bibbyetal.2000). Forsomespecies,g(0)maybe<1andisaffectedbyplatformtypes(size of boat and those surveys conducted on foot). Furthermore, the

14

assumptionofperfectdetectionalongatransectisoftenveryunrealistic exceptinthecaseofnarrowstrips. Mostobservationsweremadenofurtherawaythan12metersfrom the observer,and this suggests that as distance away from the transect increased, detection probability decreased. The distance histogram calculatedfortheobservationsinthisstudysuggestedthatonlytreeboas close to transect lines were detected with certainty. However, in cases wheredistancehistogramsdeclinesteeply,detectionprobabilitiescanbe difficulttoestimate,andoftenresultinpoordensityestimates(Buckland etal.2001). Rapid declines in detection may be due to tree boa behaviour or inaccurate sighting distance and angle measurement. Cooks tree boas aretypicallystationaryduringthedayandsomeettheassumptionthat individualsshouldnotmoveinrelationtoobservers.Thetreeboasinthis studywereinactive,evenwhenunderverycloseobservation,andwere undisturbedbyobserveractionsand/ornoise. In this study, it was difficult to assess the assumption of measurement accuracy, especially as measurements were made by eye andnotwithequipmentsuchaslaserrangefinders.Untrainedobservers canbepooratjudgingdistancesbyeye(Alldredge,etal.2007).Assuch, observer training and testing is very important in obtaining correct distancemeasurements.Observersinthisstudyweretrainedtoestimate distancesbyeye.However,itisnotedthatcorrectmeasurementsinthe fieldwithouttheuseoflaserrangefinderscanbeverydifficulttoobtain, especially under conditions of dense vegetation and whilst on a moving platform,asperformedintheswamp. In general, careful survey design, and proper stratification and samplingofthestudyareaisnecessary,toavoidbiasesinsurveydata.In addition, it has been suggested that a sample size of 6080 is the minimum threshold for reliable population size and density estimates (Buckland et al., 2001), and an accurate population estimate may be unlikely with smaller sample numbers, such as those obtained in this survey. There have been surveys of other island boa species that have estimated population densities. A study of two endemic boas in the Caribbean (the Mona boa Epicrates monensis monensis, and the Virgin Islands boa Epicrates m. granti) conducted over 9 years from 1984, suggested that density values for these species can vary between 1.5 202boasperha.Thesespeciesarelistedasthreatenedandendangered, respectively,undertheU.S.EndangeredSpeciesAct.Botharenocturnal species and difficult to count in the field. Due to largescale habitat

15

destruction and the introduction of exotic predators on these islands (e.g.,rats,cats)extrapressurehasbeenexertedontheseboasovermost of their range (Tolson and Garcia 2003). Population data for these two species was crucial to the development of their recovery plans. The current Cooks tree boa survey was conducted over a short period and factors such as boa population dynamics were not assessed. However, distancesamplingcanbeadvantageouswhensurveyinglargeareaswith difficult terrain and once the key assumptions are met, can produce an accurateestimateofboadensity. Conclusions Thenaturalhistoryofsnakesoftenmakesthecollectionofreliabledata on their populations challenging (Lind et al. 2005). This study demonstrates that distance sampling can provide a means to estimate density of the Cooks tree boas in the Caroni Swamp, once the key assumptions are met. It also shows that distance sampling is advantageous in providing estimates on the Cooks tree boa when surveying large areas, where obtaining larger samples in the field is difficult,andwherebudgetortimeforresearchislimited. Other important considerations include the use of other data to obtainamoreaccuratepopulationdensity.Duetothesecretivenatureof most snakes and their intricate activity cycles, accounting for snakes annual activity is critical in obtaining accurate population estimates (Camachoetal.2005). The lack of a standardized techniques for monitoring population statusremainsamajorobstacleinthemanagementofsnakepopulations globally (Dorcas and Wilson 2009). However, a more complete understanding of snake ecology at the individual, population and landscape levels will be important in the development of effective conservation programs for these animals. An appreciation of the Cooks tree boa should also be encouraged to change the negative view of snakes by people, thereby making a difference in their protection and longterm survival. More longterm research is required into the population distribution and trends of the Cooks tree boa, if it is to be conserved at the Caroni Swamp. In nature, monitoring a populations changewithtimeismoreimportantforawildpopulationthananysingle estimateofpopulationdensity. Acknowledgements The staff at the Caroni Visitors Centre at the Caroni Swamp are recognised, for providing access to their office and information on the

16

CaroniSwamp.SpecialthanksMadooToursandC.Madoo,D.Madooand S. Madoo who provided transport, navigation of the Caroni Swamp and adviceonspeciesbehaviour.ThisresearchwasfundedbytheUniversity of the West Indies. The authors also wish to recognize the invaluable support of the many volunteers in the field, and for those constructive commentsgivenbyfamilyandfriends.

References
Alldredge, M. W., T. R. Simons and K. H. Pollock. 2007. A field evaluation of distance measurement error in auditory avian point count surveys. J Wildlife Management 71 (8):27592766. Bibby, C. J., N. D. Burgess, D. A. Hill and S. H. 2000. Bird census techniques, 2nd ed. AcademicPress,London. st Boos,H.E.A.2001.ThesnakesofTrinidadandTobago.1 ed.W.I.Moody,Jr.,naturalhistory series;no.31.TexasAandMUniversityPress. Buckland, S. T., D. R. Anderson, K. P. Burnham, D. L. Thomas and J. L. Laake. 2001. Introduction to distance sampling: estimating abundance of biological populations. OxfordUniversityPress,NewYork. Burnham. K. P. and D. R. Anderson. 1998. Multiple model selection and inference: an informationtheoreticapproach.Springer,London. Camacho, C., J. Feinberg and T. Green. 2005. Use of the Program DISTANCE to Assess PopulationSizeoftheEasternHognoseSnake(Heterodonplatirhinos)attheBrookhaven NationalLaboratory,Upton,NewYork,11973. Dorcas, M.E., and J.D. Wilson. 2009. Innovative methods for studies of snake ecology and conservation. In Chp 1 Snakes: Ecology and Conservation/edited by Stephen J. Mullin andRichardA.Seigel.CornellUniversity,NewYork. ForestryDepartment,FoodandAgricultureOrganizationoftheUnitedNations.2005.Global forestresourcesassessment2005:ThematicstudyonmangrovesinTrinidadandTobago Country profile. Forest resources development service forestry department. Forest resourcesdivisionFAO,Rome(Italy). Harris,R. B.,and K.P.Burnham. 2002.Onestimatingwildlifedensitiesfromlinetransect data.ActaZoolSinica48:812818. Hiby, L., and M. B. Krishna. 2001. Line transect sampling from a curving path. Biometrics 57:727731. Hyrenbach, K. D., C. L. Baduini and JR. Hunt. 2001. Line transect estimates of shorttailed shearwater Puffinus tenuirostris mortality in the southeastern Bering Sea, 19971999. MarOrnithol29:1118. Kissling, M. L., and E. O. Garton.2006. Estimating detection probability and density from pointcount surveys: a combination of distance and doubleobserver sampling. The Auk 123(3):735752. Kuhl, H., F. Maisels, M. Acrenaz and E. A. Williamson. 2008. Best practice guidelines for surveysandmonitoringofgreatapepopulations.OccasionalpaperoftheIUCNSpecies survivalcommissionno.36. Lind,A.J.,H.H.WelshandD.A.Tallmon.2005.Gartersnakepopulationdynamicsfroma 16year study: considerations for ecological monitoring. USA Ecological Applications, 15(1). Marques,T.A.,L.Thomas,S.G.FancyandS.T.Buckland.2007.Improvingestimatesofbird densityusingmultiplecovariatedistancesampling.TheAuk124(4):12291243.

17

Mullin, S. J., and R. J. Siegel. 2009. Snakes: Ecology and Conservation. Cornell University, NewYork. Newey,S.B.,M.Enthoven,andS.Thirgood.2003.Candistancesamplinganddungplotsbe usedtoassessthedensityofmountainharesLepustimidus?WildlifeBiol9:185192. Plumptre, AJ. 2000. Monitoring mammal populations with line transect techniques in Africanforests.JApplEcol37:356368. Somershoe, S. G., D. J. Twedt and B. Reid. 2006. Combining breeding bird survey and distance sampling to estimate density of migrant and breeding birds. The Condor 108:691699. Tolson,P.JandM.A.Garcia.2003.Mona/VirginIslandsBoa:AU.S/PuertoRicopartnership seekstorecoverendangeredboa.ToledoZoologicalGardens,P.O.Box4010,Toledo,OH 43609. Wehekind, L. 1955. Notes on the foods of Trinidad snakes. British Journal of Herpetology 2:913.

18

Demographyandgeneralecologyofanintroducedprimatethe tuftedcapuchin(Cebusapella)inChaguaramas,Trinidad DarshanNarang1,2,HowardP.Nelson2,andAndrewLawrence2

DepartmentofLifeSciences,TheUniversityoftheWestIndies,St.Augustine,Trinidad, WestIndies. 1 CorrespondingAuthorEmail:dsn95@yahoo.com

Abstract Twoendemicnonhumanprimatesubspeciesareknownfromtheisland of Trinidad: the Red howler monkey (Alouatta seniculus insulanus) and the Whitefronted capuchin (Cebus albifrons trinitatis). A third primate, the tufted capuchin (Cebus apella), appears to have been introduced to theChaguaramaspeninsuladuringtheUnitedStatesmilitaryoccupation of that site during 19411977. These tufted capuchins now occur sympatrically with the two endemic Trinidadian monkeys at Chaguaramas, while virtually nothing is known of the ecology and demography of this introduced Cebid. This study investigated the population density of the tufted capuchins within the Chaguaramas peninsula. Line transect distance surveys were conducted within the Chaguaramas peninsula from May to November, 2010. The cumulative distancesampledduringthestudywas34.6km,withasamplingeffortof 200hours.Conventionaldistancesampling(CDS)inDistance6.0software wasusedtoestimatedensity.Capuchindensitywithinthestudyareawas 17.7 troops per km2 with an overall average troop size of 5.2 0.6 individualspertroop(n=30)andarangeof215individualspertroop. Keywords Tufted capuchin, Cebus apella, population density, introduced species, Trinidad,distancesampling. Introduction Humans have introduced animals throughout the world for various reasons: aesthetics, food, hunting and sport, commercial enterprises, controllingpests,scientificresearch,andthroughaccidentalintroduction, escapeesandpetkeeping(Long2003).TheCaribbeanislandshavehada longhistoryofEuropeanandAmericancolonizationandasaresulthave experienced frequent species introduction. These exotics have included commensalrodents,themongoose(Herpestesauropunctatus),arangeof domestic species, as well as peccaries and monkeys (four species). In

19

total, the Caribbean has experienced at least 37 mammalian introductions,28ofwhichhavebecomeestablished(Long2003). Trinidad is home to two endemic nonhuman primate subspecies (Agoramoorthy and Hsu 1995): the red howler (Alouatta seniculus insulanus)ofthefamilyAtelidaeandtheTrinidadwhitefrontedcapuchin (Cebus albifrons trinitatis) of the family Cebidae. It has been suggested thatC.albifronswasintroducedtotheislandduringpreColumbiantimes (Long 2003). However, no comparative genetic studies with mainland South American populations have been conducted. One competing theory is that C. albifrons may have also occurred naturally on Trinidad and that this population was isolated when Trinidad became an island approximately 10,000 years ago, during the last ice age (Vuilleumier 1972). RecentresearchonprimatesinTrinidadhasfocusedonthreewildlife sanctuaries in Trinity Hills, Central Range and Bush Bush Island (Bacon andffrench1972,Neville1976;AgoramoorthyandHsu1995;Rylandset al. 1997; Phillips 1998, Phillips et al. 1998, Phillips and Abercrombie 2003).WhitefrontedcapuchinsinTrinidadarelimitedtoonlytwoofthe threewildlifesanctuaries:TrinityHillsandBushBushIsland(Phillipsand Abercrombie 2003). Whitefronted capuchins need larger ranges (>80 ha.) than Red howlers and thus are not as likely to persist in viable populations outside of large tracts of forest (Phillips and Abercrombie 2003). The major threats to primates in Trinidad include hunting, habitat destruction, and harvest of important plant species within and near protected areas (Phillips and Abercrombie 2003). Many important habitat areas for primates in Trinidad already enjoy de jure protection with 13 legal sanctuaries, comprising about 3.4% of the total land area, that are currently managed by the Forestry Division (Phillips and Abercrombie 2003). The Environmental Management Authority (EMA) has also designated some areas of the country as environmentally sensitive, including the Nariva Swamp an important primate habitat (EMA 2006). Some monkeys may also persist in unprotected areas, and protectionofthesepopulationsmaybeparticularlydifficult(Phillipsand Abercrombie 2003). The Conservation of Wildlife Act (67:01) 1953, providesprotectiontoallprimatesinthecountryregardlessofwhether theyoccurinStateorprivateforest. Athirdspeciesofprimate,thetuftedcapuchin(Cebusapella)of thefamilyCebidae,wasintroducedontheChaguaramaspeninsuladuring theUnitedStatesmilitaryoccupationfrom19411977(John1998).These monkeys were kept at a small zoo in the Scotland Bay area of the

20

peninsula.Thesetuftedcapuchinsnowoccursympatricallywiththetwo endemicTrinidadianmonkeyswithintheChaguaramaspeninsula.There has been no research on these freeranging capuchins within Chaguaramas. This study investigates the population density and distribution of the tufted capuchins within the Chaguaramas peninsula. The general ecology of these capuchins is also describedas it relates to populationdynamics. Methods Studyarea This study was conducted in the Chaguaramas National Park (CNP), locatedonthenorthwesternpeninsulaofTrinidad.Thispeninsulaisfive and a half miles long from East to West, and extends three and a half milesfromNorthtoSouth(Figure1),withanareaofapproximately40.5 km2 (CDA 2010). The national park was created and is managed almost exclusivelybytheChaguaramasDevelopmentAuthority(CDA)underthe CDAAct(35:02)1972.
Figure1:LocationoftransectswithintheChaguaramaspeninsulaand thedistributionoftuftedcapuchins

The CNP is characterized by several hills reaching a maximum height of 546 m at Morne Catherine. There are numerous seasonal streams, and the predominant vegetation is tropical dry forest (Beard 1946)withpatchesofmixedprimaryandsecondary,wetanddryforest, as well as various forms of agriculture. This study was primarily undertaken within the Tucker Valley and Scotland Bay areas of the peninsula.

21

The Tucker Valley consists of a mosaic of land uses including large farms, a golf course, a rifle and archery range, as well as various military institutions. It includes and stands of exotic bamboo (Bambusa vulgaris), which have become dominant in several areas of this valley. The Scotland Bay area shows higher plant diversity and the area is strongly deciduous. However, in adjacent areas Brazilian Rubber (Hevea brasiliensis)andB.vulgarisbecomedominant. The climate within the peninsula is seasonal with a dry season from January to May and a wet season from June to December (TTMS 2010). Surveys Linetransectdistancesamplingtechniquesarearangerelatedmethods for estimating the abundance of wildlife populations (Buckland et al. 1993, 2001, 2004; Thomas et al. 2002). Line transect distance surveys were conducted between 5:30 am to 11:30 am, during the wet season, between the months of May to November, 2010. These line transects mostlyfollowedexistingforesttrailswithintheScotlandBayareaanda trailfromtheGolfCoursetoMacqueripeBaywithintheTuckerValley. Observations were made while walking slowly (12 km/hr.) along trails, and recording encounters within 100 m of either side of these trails.Uponencounteringagroupofmonkeys,1520minuteswasspent on recordingnumber of individuals, canopy height,height of monkeyin canopy, angle of sighting, distance of animal from observer, habitat description, sex and age category of individuals, activity on sighting and direction of general movement. For each group of monkeys, perpendiculardistancewasestimatedfromthecentreofthetrooptothe axisofthetransectline.Detectionfunctionsforundetectedtroopswere thenestimatedfromtheseperpendiculardistances(Bucklandetal.2001, 2004). Analyticmethods The observational data was analyzed using the conventional distance sampling(CDS)inDistance6.0.(Thomasetal.2009).Distance6.0models the detection probability as a function of the animals perpendicular distancefromthetransect.Inthismodeldetectionfunction,clustersize andencounterrateareestimatedseparately,andtheresultsarepooled toderivedensity(Thomasetal.2009). Distance sampling makes three key assumptions: Firstly, animals on thetransectaredetectedwithcertainty.Inthisstudy,multipleobservers wereused,withdatapooledacrossthem.Thesecondassumptionisthat

22

theobjectsdonotmove.Thethirdassumptionisthatallmeasurements areexact.Untrainedobserversareoftenpooratjudgingdistancesbyeye orear(Alldredgeetal.2007).Forthisstudy,theobserversweretrained toestimatedistancespriortothesurveystoincreaseaccuracy.Itshould be noted that recent advances now permit some relaxation in these assumptions(Bucklandetal.2001). In addition to exact distances, the cluster sizes of the tufted capuchins were assumed to be accurately recorded, where these observationsoccurredclosetothetransect.Itwasalsoassumedthatthe twocapuchinspeciesatthesitewerenotmisidentified. CDS uses a parametric function that can be paired with one or more adjustment terms. The Distance software used in this study provides 4 functions (uniform, halfnormal, hazardrate and negative exponential) and these can be paired with series adjustment functions can include cosine, or hermite or simple polynomial identifies. Here, varianceofadensityestimatewascalculatedusingthedeltamethod,and comprises three components encounter rate, detection function and meanclustersize(Bucklandetal.2001).Theselectionofanappropriate model was done using a twostep process. The first step involved exploratorydataanalysis,whichalloweddetectionofanyproblemssuch as overdispersion in the data. The second phase of the analysis was model selection, which included determination of an appropriate truncationdistancefortheobservationdata(Bucklandetal.1997,2001). Ninemodelcombinations,eachconsistingofakeyfunctionand an adjustment term, were used to describe the observed data, and Akaikes Information Criterion (AIC) was used to select the most appropriatemodel(BurnhamandAnderson2002). Results Thirtyfour days were spent in the field with a sampling effort of 200 manhours. A total of seven transects, covering a total distance of 34.6 km and representing an area of 6.92 km2, were surveyed in the Chaguaramas peninsula (Table 1). One hundred and fiftysix individual animal observations were made during 30 troop observations. Considerable variability in count frequencies were detected in the data (Figure 2), and the shape of the histogram suggested overdispersion in the data. To compensate for this issue, a cutoff point of 50 m was selectedsinceonly7%oftheindividualobservationsoccurredinthe50 100mzone(Figure2).Theselectedmodelwasahazardratekeyfunction withacosineadjustmentthatresultedinthelowestAICvalue(222.15).

23

Table1TransectlengthsandobservedtroopsatChaguaramasstudysites

The estimated density of the tufted capuchins within the sampling area was 17.7 troops per km2 with the estimate of expected value of troopsize5.20.6individualspertroop(n=30).Theoverallestimateof densitywas92.3animals/km2. Therangeofanimalspertroopwasbetween215animalsand0.15 troops were observed per hour of observation. Most troops were observedalongtheMacqueripetrailandonlyafewtroopswereseenin theScotlandBayareaandalongElderRoad(Figure1).
Figure2DetectiondistancesofC.paellawithintheChaguaramaspeninsula.

Discussion Demographics The population of the tufted capuchin in the Chaguaramas peninsula appears similar to that of other locations where this species occurs. In

24

comparison,theencounterratefortuftedcapuchinsonMargaritaIsland hasbeenreportedatbetween0.02and0.23troops/hoursofobservation (MarquezandSanz1991),while0.15troops/hourswereobservedinthis study.TheaveragegroupsizeonMargaritaIslandwas4.5individualsper group, with 5.2 individuals per group in the Chaguaramas study population (Marquez and Sanz 1991). Similarly, group sizes in Chaguaramas were identical to that reported elsewhere (Rylands et al. 2008).ThetuftedcapuchinsdensityinChaguaramaswasalsoverysimilar to that of the red howlers density on Bush Bush Island, Nariva, of 83 individualsin20troops/km2(AgoramoorthyandHsu1995). Habitat Cebus is found in almost every kind of Neotropical forest. This generalizationisalsotrueofthetuftedcapuchin,whichisknownfroma broad range of forest habitats and regarded as ecologically plastic in its habitat requirements (Mittermeier and van Roosmalen 1981). These Cebids typically use the lower to midcanopy and understory of tropical forests (Fragaszy et al. 2004). In this Chaguaramas study, the tufted capuchin was found throughout the vertical forest profile, as well as foragingontheground. Diet The tufted capuchin is omnivorous and able to use a variety of food sources,manyofwhicharenotusedbyotherNeotropicalspecies.Other authorshavereportedthattheirdietcontainsfruits,pith,nectar,leaves, nuts, insects, birds, small mammals, lizards etc. (Terborgh 1983; Spironello 2001). In Chaguaramas, the tufted capuchins were observed feeding on a range of food sources including various palm fruits, exotic fruitsfromintroducedtrees,andvariousinsectlarvae.Thesecapuchins spentasignificantamountoftimeforaginginthebambooandfeedingon termites and ants. On one occasion, capuchins were observed eating a frog. The general daytime feeding habits of Cebus apella is categorized broadlyintotwotypes:eitherforagingasscatteredindividualsinagiven area; or in groups moving slowly or quickly in a given direction (Izawa 1979,1980).Thesetypesoffeedingactivitiesoccurredmainlyalongthe trailtotheMacqueripeBeach,wherethecapuchinswouldbemovingin an easterly or westerly direction while slowly feeding on mainly animal proteinamongthebamboo.Thecapuchinsfedopportunistically,thusthe specific movement type during feeding was not always obvious. The capuchins were always scattered among the vegetation and there may

25

havebeenanoverlapamongthefeedingareasofdifferenttroopsatthe sametime. Interspecificinteraction Tufted capuchins typically live sympatrically with other primates, where theyoccurnaturally(Fragaszyetal.2004).Notably,duringthisstudy,no interactions were observed between any of the primate species in Chaguaramas. During this study, only one troop of the Trinidad white fronted capuchin (Cebus albifrons trinitatis) was observed east of the Tucker Valley Road, while no tufted capuchins were observed there. In addition, no Trinidad whitefronted capuchins were observed in any transect west of the Tucker Valley Road, where the Tufted capuchins wereobserved. ComparativestudiesofCebusalbifronsandCebusapellainColombia suggest strong differences in their behaviour (Defler 1979). Cebus albifrons troops tended to be large and multimale, which were very aggressive towards neighbouring conspecific troops. These primates defended an exclusive territory with little overlap by conspecifics, and used longdistance calls and fighting to maintain their territories (Defler 1979).Incomparison,troopsofCebusapellaweresmall,didnotdefenda territory, displayed few agonistic interactions and had overlapping territories(Defler1982).ThetuftedcapuchinsinChaguaramasexhibited similarintraspecificbehaviourtothatobservedinColombia.However,no interactionsbetweenthetwocapuchinspecieswereobservedduringthis study. Redhowlermonkeyswereobservedthroughoutthestudyarea,but in relatively low numbers. However, no interactions were observed betweenthesehowlersandthetuftedcapuchinsinthisstudy. Tooluse Wild tufted capuchins can use tools to obtain food. Thus, fruits may be brokenopenontreesurfaces,orwithbatonlikebranches(Boinskietal. 2000). During this study, tool use was observed, including the use of batons to obtain insects from bamboo and use of bamboo leaves to obtainwaterfromhollowsofdeadbamboostalks.Thisisverysimilarto the tool use observed in wild Trinidad whitefronted capuchins, which usedleavesascupstoretrievewaterfromtreecavities(Phillips1998). Conclusion Thisisthefirstpublishedstudyoftheestablishmentofanalienprimate ontheislandofTrinidad.Invasivealienspeciesarecurrentlyoneofthe

26

most significant threats to biodiversity in the Caribbean, and have significant implications for the Caribbeans economy, the environment and for human health. Further research is therefore, required to understandthehistoryandecologyofthisintroducedprimatespecies,its potential invasiveness and its impact on the two endemic primate subspeciesinTrinidad. Acknowledgments ThankstotheWildlifeSection,ForestryDivision,fortheirsupportinthe field, to L. Guy for providing necessary arrangements for the access to Scotland Bay; to the Chaguaramas Development Authority for providing theresearchpermit;toR.HonoroftheTrinidadandTobagoRegiment for escorting us throughout the Scotland Bay surveys; to the National HerbariumofTrinidadandTobagoforprovidingsupportinthefieldand intheidentificationofplants;andtoallvolunteers. References
Agoramoorthy,G.andHsu,M.J.1995.PopulationstatusandconservationofRedHowling Monkeys and Whitefronted Capuchin monkeys in Trinidad. Folia Primatology 64:158 162. Alldredge, M.W., Simons, T.R. and Pollock, K.H. 2007. A field evaluation of distance measurement error in auditory avian point count surveys. Journal of Wildlife Management71:27592766. Bacon, P. R. and R. P. Ffrench. editors. 1972. The wildlife sanctuaries of Trinidad and Tobago.PreparedbytheWildlifeConservationCommittee,MinistryofAgriculture,Lands andFisheries,TrinidadandTobago. Beard J.S. 1946. The natural vegetation of Trinidad and Tobago. Oxford at the Clarendon Press. Boinski,S.,Quatrone,R.P.andSwartz,H.2000.Substrateandtoolusebybrowncapuchins in Suriname: ecological contexts and cognitive bases. American Anthropology 102(4): 74161. Buckland,S.T.1992.Fittingdensityfunctionsusingpolynomials.AppliedStatistics41:63. Buckland, S.T., Anderson, D.R., Burnham, K.P. and Laake, J.L. 1993. Distance Sampling: EstimatingAbundanceofBiologicalPopulations.ChapmanandHall,London. Buckland, S.T., Anderson, D.R., Burnham, K.P., Laake, J.L., Borchers, D.L. and Thomas, L. editors.2004.AdvancedDistanceSampling.OxfordUniversityPress,London. Buckland, S.T., Anderson, D.R., Burnham, K.P., Laake, J.L., Borchers, D.L. and Thomas, L. 2001.IntroductiontoDistanceSampling.OxfordUniversityPress,London. Buckland,S.T.,Burnham,K.P.andAugustin,N.H.1997.Modelselection:anintegralpartof inference.Biometrics53:603618. Buckland,S.T.,Summers,R.W.,Borchers,D.L.andThomas,L.2006.Pointtransectsampling withtrapsorlures.JournalofAppliedEcology43:377384. Burnham, K. P., and D. R. Anderson. 2002. Model Selection and Multimodel Inference: A PracticalInformationTheoreticApproach.2ndeditionSpringerVerlag,NewYork. CDA (Chaguaramas Development Authority). 2010. http://chagdev.com/ Accessed May, 2010.

27

Defler,T.R.1979.OntheecologyandbehaviorofCebusalbifronsinnorthernColombia,1: Ecology.Primates20:475490. EMA (Environmental Management Authority). 2006. The Environmentally Sensitive Areas (NarivaSwampManagedResourceProtectedArea)Notice.LegalNoticeNo.334. Fragaszy,D.M.,Visalberghi,E.andFedigan,L.M.2004.Thecompletecapuchin:thebiology ofthegenusCebus.Cambridge:CambridgeUniversityPress. Izawa,K.1979.Foodsandfeedingbehaviourofwildblackcappedcapuchin(Cebusapella). Primates20:5776. Izawa,K.1980.Socialbehaviourofthewildblackcappedcapuchin(Cebusapella).Primates 21(4):443467. John,B.1998.ChangingPatternsofLanduseinChaguaramas.UnpublishedUndergraduate Thesis. University of the West Indies. St. Augustine. http://www.triniview.com/CarenageChaguaramas/Chaguaramas.html Accessed August 2010. Long,J.L.2003.IntroducedMammalsoftheWorld:TheirHistory,DistributionandInfluence. CSIROPublishing,Australia. MarquezL,SanzV.1991.EvaluacindelapresenciadeCebusapellamargaritae(Hollister, 1914) en la Isla de Margarita. Trabajo Especial de Grado, Universidad Central de Venezuela,Caracas. Mittermeier, R.A and van Roosmalen, M.G.M. 1981. Preliminary observations on habitat utilizationanddietineightSurinamemonkeys.FoliaPrimatology36:139. Neville, M. 1976. The population and conservation of howler monkeys in Venezuela and Trinidad.In:NeotropicalPrimates:FieldStudiesandConservation,R.W.Thorington,Jr. andP.G.Heltne.editors.pp.101109.NationalAcademyofSciences,Washington,DC. Phillips, K. 1998. Tool use in wild capuchin monkeys (Cebus albifrons trinitatis). American JournalofPrimatology46:259261. Phillips, K. A. and Abercrombie C. L. 2003. Distribution and Conservation Status of the PrimatesofTrinidad.PrimateConservation19:1922. Phillips,K.A.,Elvey,C.R.andAbercrombie,C.L.1998.ApplyingGPStothestudyofprimate ecology:Ausefultool?AmericanJournalofPrimatology46:167172. Rylands,A.B.,Mittermeier,R.A.andRodrguezLuna,E.1997.ConservationofNeotropical primates:Threatenedspeciesandananalysisofprimatediversitybycountryandregion. FoliaPrimatologica68(35):134160. Rylands,A.B.,Boubli,J.P.,Mittermeier,R.A.andWallace,R.B.2008.Cebusapella.In:IUCN 2010.IUCNRedListofThreatenedSpecies.Version2010.4.http://www.iucnredlist.org/ AccessedJuly2010. Spironello,W.R.2001.Thebrowncapuchinmonkey(Cebusapella):ecologyandhomerange requirements in central Amazonia. In: Bierregaard, R.O, Gascon, C., Lovejoy, T.E. and Mesquita, C.G. editors. Lessons from Amazonia: the ecology and conservation of a fragmentedforest.NewHaven:YaleUniversityPress.p27183. Terborgh,J.andJanson,C.H.1983.TheecologyofprimatesinsoutheasternPeru.National GeographicalSocietyResearchReport15:65562. Thomas,L.,Laake,J.L.,Rexstad,E.,Strindberg,S.,Marques,F.F.C.,Buckland,S.T.,Borchers, D.L., Anderson, D.R., Burnham, K.P., Burt, M.L., Hedley, S.L., Pollard, J.H., Bishop, J.R.B. and Marques, T.A. 2009. Distance 6.0. Release 2. Research Unit for Wildlife Population Assessment, University of St. Andrews, UK. http://www.ruwpa.stand.ac.uk/distance/ AccessedJuly2010. Thomas,L.,S.T.Buckland,K.P.Burnham,D.R.Anderson,J.L.Laake,D.L.Borchers,andS. Strindberg. 2002. Distance Sampling. Pages 544552 in ElShaarawi, A.H and Piegorsch, W.W.editors.EncyclopediaofEnvironmetrics.JohnWileyandSons,Chichester,UK.

28

TTMS (Trinidad and Tobago Meteorological Service). 2010. Government of Trinidad and Tobago.Accessedathttp://www.metoffice.gov.tt/climate/article.aspx?id=4820Accessed June2010. Vuilleumier, B. S. 1972. Pleistocene changes in the fauna and flora of South America. Science173:771780.

29

SpatialandTemporalDiversityinGroundLevelFruitFeeding Butterflies ImranKhan1,2,ChristopherK.Starr2,HowardP.Nelson2,and AndrewLawrence2


2

DepartmentofLifeSciences,TheUniversityoftheWestIndies,St.Augustine,Trinidad, WestIndies.Email:immyboy323@yahoo.com 1 CorrespondingAuthor

Abstract Butterfly diversity has been proposed as an indicator of habitat disturbance.Rapidassessmentoffruitfeedingbutterfliesisoftenusedto predict disturbance impacts and so aid in the development approval process in Trinidad. However, such an approach makes several assumptions about the relationship between butterfly diversity and habitat disturbance. This study reports on an investigation of these relationships. Groundlevel fruit feeding butterflies were trapped within forest,agricultural,andcocoahabitatsinGrandeRiviere.Sevensampling stations were established in each habitat type, and trapping was replicatedsixtimeseachmonthfromMaytoSeptember2010.Shannons Diversity (H) was calculated for each plot to determine whether HF>HA>HC. H was also calculated monthly across all plots to examine temporalchangesindiversity.Hforthethreeplotswasfoundtobe2.48, 1.56,and2.00,respectively.HforthemonthsofMaytoSeptemberwas found to be 1.72, 1.98, 1.05, 1.99, and 2.15, respectively. Comparison withanotherstudyinTrinidadsuggeststhattheguildofgroundlevelfruit feeding butterflies may be used as biological indicators of disturbance, but not for rapid assessments. Areas dominated by C. minor are less disturbed, and a dominance of E. penelope and E. hermes appears associatedwithdisturbance. Keywords Caligominor,disturbance,Euptchiahermes,Euptchiapenelope,indicator, Shannondiversity Introduction The Certificate of Environmental Clearance Rules (2001) is generally consideredthemosteffectivemechanismforregulatingdevelopmentin Trinidad and Tobago. For developments that may have significant environmental impacts, the regulating agency, the Environmental

30

Management Authority (EMA), typically requires an Environmental Impact Assessment (EIA), before issuing or denying a Certificate of Environmental Clearance (CEC). The agency issues Terms of Reference (TOR), which are a blueprint for completing the EIA (EMA, 2010). These TORsprovidedeveloperswithanindicationofwhatstudiesarerequired foranEIA. Typically, developers can satisfy the TOR by undertaking rapid ecological assessments (EMA, 2010). Such assessments provide a snapshot of the biological diversity in the area earmarked for development, and include a description of the species richness and diversityofmajortaxonomicgroupssuchastrees,birds,fish,andbenthic macrofauna (EMA, 2010). This assessment provides a portrait of the degree of disturbance already experienced at the site. From a development perspective, the more disturbed a site is found to be, the easieritwouldbetoobtainaCEC,andtheCECitselfshouldnotrequire intensive conservation efforts to protect the biological resources at the site. However, while the most diverse taxa on Earth are insects (Wilson, 1988), it is notable, that this taxon has never been identified as objects for study in TORs by the EMA (EMA, 2010). Insects are probably just as overlooked on the local stage as they are on the international level (Meyers et al. 2000; Clark and May, 2002; and Leather, Basset, and Hawkins, 2008). If this taxon was to be considered in the future by the EMA,onekeyquestionremainstheidentificationofthegroupofinsects tobeusedasindicatorsofhabitatdisturbance.Termites(Vasconcelloset al, 2010; Lawton et al. 1998; Eggleton et al, 1995), beetles (Scheffler, 2005; Rainio and Niemel, 2002; Davis et al. 2001) and ants (Philpott, Perfecto, and Vandermeer, 2006; Watt, Stork, and Bolton, 2002; Roth, Perfecto,andRathcke,1994)areexamplesofarthropodgroupsthathave beenusedinhabitatdisturbancestudies. Butterfliesareanotherwellstudiedgroupthathavethepotentialto serve as indicators of disturbance (Sundufu and Dumbuya 2008), and have been widely used in habitat disturbance studies elsewhere (Beck andSchulze2000;WoodandGillman1998;DeVriesetal.1997;Hilletal. 1995;Sparrowetal.1994;Spitzeretal.1993;Kremen1992;Brown1991; Lovejoy et al. 1986;). These insects have also been suggested as good environmentalindicatorsduetotheirsensitivitytomicroclimateandlight intensitychanges(WoodandGillman,1998;Erhardt,1985),andbecause of their complex life history (Kremen, 1992; Ehrlich, 1984). Butterfly taxonomy, distribution and natural history (Brown, 1997) is also well

31

knownandmanyspeciescanbereliablyidentifiedinthefield(Woodand Gillman,1998). This paper examines the suitability of butterflies as indicators of disturbanceinTrinidad,andtheirutilityinrapidecologicalassessmentsin the local EIA process. This was done by comparing butterfly diversity under natural forest, cocoa plantations and openfield agriculture land usetypes. Methods StudySite ThisstudywasconductedatGrandeRiviere,aremotevillagesituatedin northeastTrinidad(Figure1).Here,threetreatmentsofvaryinghuman disturbancewerechosen:(undisturbed)forest(F);cocoaplantations(C); andopenfieldagriculture(A).Foreststudyareashad>70%canopycover and were dominated by tall trees of several species. Cocoa plantations had > 70% canopy cover, with approximately 90% dominance of Cacao spp. Finally, openfield agriculture areas had < 30% canopy cover, very fewtrees,andweredominatedbyagriculturalshortcropsandgrass.The agriculturalplotandthecocoaplotwereadjacenttoeachother,andthe forestplotwaslocated1.5kmawayfromthesetwo(Figure1).

Figure1:LocationofForest(F),Agriculture(A) andCocoa(C)DisturbanceAreas andSamplingStationsinGrandRiviere,Trinidad

32

Collection Butterflies were collected at the study sites, during the months May to September, which have been previously documented as the best times for collecting these insects (Barcant 1970). Butterflies were sampled for six days of each month (three days each at the start and end of each month). Baitedfruittrapswereusedtosamplebutterflycommunitiesateach site(MendzandFunes2007).Twentyoneinvertedconebutterflytraps were used in this study. Seven traps were set up in each of the three disturbancetreatments(F,C,andA).Oneachday,trapswerebaitedfrom 8amto12pm,andcheckedapproximately24hourslater.Allspecimens were removed for later identification, and bait was replaced if needed. The bait consisted of a fermented mixture of overripe/rotting bananas, brownsugar,andcanejuice. Analysis Butterfly diversity in the different habitats (F, A, and C), was estimated usingShannonsDiversity(H),andEquitability(EH),NumberofIndividuals (N), and Species Richness (S). These indices were calculated using the total results from F, A, and C treatments for each month from May to September to examine monthly changes in diversity. Nonmetric multidimensional scaling nMDS was also used to investigate differences in the diversity of butterflies among the three disturbance types. nMDS analysisofthetopfivespeciesfromeachareawasalsousedtoidentify indicatorspeciesforleastandmoredisturbedareas. Results Twothousandtwohundredandfortyfive(2245)individualsof45species of butterfly were captured during 609 trapdays of effort at Grande Riviere. Diversity indices per treatment for the sampling period are presentedatTable1,andTable2.
Table 1: Butterfly Diversity Indices for Forest, Agriculture and Cocoa Areas in GrandRiviere,Trinidad DISTURBANCEAREA Forest Agriculture Cocoa DIVERSITYINDICES H EH 2.48 0.76 1.56 0.44 1.99 0.61

N 443 1048 754

S 26 35 26

33

ThesedatasuggestthatthehighestHvaluewasrecordedintheforest, andlowestintheagriculturalareas.However,boththegreatestnumber of species and individuals were observed from the agricultural area, whichalsohadthelowestEHvalue.
Table 2: Monthly Butterfly Diversity Indices for Forest, Agriculture and Cocoa AreasinGrandRiviere,Trinidad MONTH DIVERSITYINDICES May June July August September H 1.72 1.98 1.05 1.99 2.15 EH 0.67 0.70 0.29 0.58 0.64 N 116 144 674 396 913 S 13 17 37 30 27

H values for the months of June, August and September were similar, with a slight decrease noted in May, and July showing the largest decrease. July also recorded the highest species richness with an intermediatenumberofindividuals,andthelowestEHvalue(0.29). ThenMDSanalysisofspeciesrichnessforeachsamplingstation ispresentedinFigure2.Thisfiguresuggeststhatthereisadistinctionin butterflydiversitybetweenthethreehabitattypes.Here,forestsampling stations were clustered on the left of the plot, agricultural sampling stationsweretotheright,andcocoainthecentre.Thesedatasuggest that the disturbance level may be related to changes in the butterfly diversity.


Figure2AnMDSPlotOfbutterflyspeciesrichnessateachSampleStation

34

While the nMDS analysis suggests that there were clear differences between the butterfly communities at each site, there were two agriculturalplots(A2andA3)thatclusteredwiththecocoaplots. The top five most abundant species for the three habitat types are presentedatTable3.E.penelope,T.virgilia,andColoburadirceoccurred in the top five for each of the study areas, while E. hermes was ranked secondintheopenagricultureandcocoaareas,respectively,butdidnot rank among the dominant forest species. M. insularis, C. Minor, and P. fusimaculataappearedonlyinforestsites. Wood and Gillman (1998) examined the effects of disturbance on forest butterflies in the Victoria Mayaro Reserve in southern Trinidad. They studied butterflies in disturbed (D) and undisturbed (U) areas of evergreen(E)andsemievergreen(SE)forest.Toallowcomparisonwith theirwork,diversityindiceswerecomputedfromtheirunderstorydata, andcomparedwiththeundisturbedareasinthepresentstudy(FtoUE andUSE).
Table3:ThetopfivemostabundantbutterflyspeciesfoundinForest,Agriculture andCocoaareasinGrandeRiviere,Trinidad Forest SpeciesName Number Euptchiapenelope 105 Taygetisvirgilia 55 Morpho peleides49 insularis Caligo eurilochus48 minor Coloburadirce 33 Pierella hyalinus33 fusimaculata Agriculture SpeciesName E.penelope E.hermes E.palladia T.virgilia C.dirce Number 574 272 27 23 21 Cocoa SpeciesName E.penelope E.hermes E.palladia C.dirce T.penela T.virgilia Number 353 81 70 64 38 38

Similarly,acomparisonbetweenthedisturbedareasintheirdata(C andAtoDEandDSE),andthecurrentstudy(F,UEandUSEtoC,A,DE, and D SE) was also undertaken. Diversity estimates for the Wood and Gillman(1998)studyarepresentedinTable4.Thisdatasuggeststhatin undisturbed evergreen forest (U E), H was higher than disturbed evergreenforest(DE),whileinundisturbedsemievergreenforest(USE), itwaslowerthandisturbedsemievergreenforest(DSE).

35

Table4ButterflyDiversityIndicesCalculatedfromWoodandGillman(1998). Area DiversityIndices H EH N S DE 1.78 0.81 66 9 UE 2.01 0.87 34 10 DSE 2.24 0.78 133 18 USE 1.68 0.65 86 13
DEDisturbedEvergreenForest UEUndisturbedEvergreenForest DSEDisturbedSemiEvergreenForest USEUndisturbedSemiEvergreenForest

Discussion This study set out to investigate the hypothesis that for tropical butterflies, species richness is positively correlated to vascular plant species richness, or HF > HA > HC (Simonson et al. 2001, Nyamweya and Gichuki,2000).However,itwasfoundthatforbutterfliesattheGrande RivieresitesthatHF>HC>HA. Thismaybetheresultoflandscapescalepatterneffectsofthecocoa samplingstations.Thestudyareawasnestledinalandscapeofsecondary forest with greater floral diversity than the cocoa area, while the agriculturalsamplingareawaslocatedjustnorthofthecocoaarea.The proximity of the cocoa area traps to areas richer in floral diversity may haveluredbutterfliesfromtheserichersurroundingareas. Perhaps, if Pollard Walks were used instead of baiting, the species richness and diversity in the cocoa sites would have been different. In addition, if the cocoa sample site was of a larger size, the source/sink dynamicsmayhavehadalessconfoundingeffectontheresults. However, the clear pattern of Shannon diversity (H) in the three disturbance treatments suggests that there was an effect of degree of disturbanceonspeciesrichnessandevenness. With the exception of Euptychia spp., no other species in the agriculturalplotswereobservedfeedingonfoodatthesite.Interestingly, twoagriculturalplotscouldnotbedifferentiatedfromforestplotsusing nMDS, suggesting that these plots may have been confounded by their relative position on the landscape relative to the other treatment type. Disturbanceoffloralcomposition,heightofcanopycover,andthetiming andintensityofphysicalalterationofthelandscapemayallcontributeto changesinbutterflydiversity.

36

ThemonthlyHvaluesobservedfromMaytoSeptembershowedno clear seasonal pattern. The lowest H value was in the month of July, which also recorded the highest S and lowest equitability values. The most abundant species captured in all months, wereE. penelope andE. hermes.Notably,Julywasalsothemonththatexperiencedtheheaviest andlongestperiodsofrainfall. Barcant(1970)liststhehostplantoftheEuptychiagenusasgrass, andnotesthatbutterfliesweremoreabundantfollowinganextremedry season (as was experienced in 2010). These conditions were realised in theagriculturalareaduringthemonthofJuly,andthismayaccountfor July having the highest species richness, and a population boom of the commonestspecies(E.penelopeandE.hermes). In decreasing order of abundance, the top five most abundant species for the forest sites were E. penelope, T. virgilia, M. peledies, C. minor,withC.dirceandP.fusimaculatatyingforfifth.Withtheexception of M. peledies, C. minor and P. fusimaculata, all the other species also featuredinthetopfivespeciesforthemoredisturbedareas.Thesethree specieswererecordedforeachofthetrappingmonthsmainlyfromthe forest, but they were also caught in lesser numbers in the cocoa. This makesthesespeciesidealasindicatorsofleastdisturbedsites(Sparrow etal.1994). The most noticeable difference between the forest and the other two areas are the presence of a tall canopy. This may well be the key factor affecting the distribution of these three butterfly species. Yet, during the time spent in the field, it was not unusual to observe M. peleidesflyingacrosstheagriculturalareas.However,nonewerecaught intrapsinthisagriculturaltreatment.BasedonthisitmaybesaidthatM. peleidespreferstofeedunderthecanopy. Similarly,whilethefoodplantforthelarvaeofC.minoristheMusa spp.(Barcant,1970),nonewereobservedflyingintheagriculturalarea, where Musa spp. was present, and only five individuals (N = 69) were capturedinthisarea.Thisisbecausethegenusisbothcrepuscularandits eyesaresensitivetobrightlight(FrederiksenandWarrant,2008).Despite this, when the conditions at dusk or at dawn were right for flight, the baitedtrapsintheopenagriculturalareastilldidnotattractC.minor.It may, therefore, be said that this species also prefers to feed under the forestcanopy. P. fuscimaculata was exclusively observed from canopyenclosed areas,andthisspeciesmaybeindicativeofundisturbedforestareas.E. penelopewasthemostabundantspeciesinallplots,andforeachmonth of sampling. This species was found to exhibit a greater affinity for the

37

disturbedareas(AandC),thanundisturbedarea(F).However,duetoits commonnessinallareas,relativeabundanceandnotpresence/absence datashouldbeconsidered,ifusingE.penelopeasinindicator. T. virgilia and C. dirce were also abundant in all of the disturbed areas. However, C. dirce, showed no meaningful pattern in its use of disturbedareas.EventhoughitshostplantCecropiapeltata(Barcant, 1970)favoursdisturbedareas(suchastheagriculturalandcocoaplots), comparativelyhighnumbersofC.dircewerefoundintheforestplot.T. virgilia exhibited a preference for disturbed areas, and was more dominantinthecocoa.However,thisspecieswasnotpresentinthedry season month, and 71% (N = 118) of the individuals were captured in September. These findings do not make T. virgilia a suitable indicator species. E.hermeswasthemostabundantspeciesforbothofthedisturbed areas(CandA),andexhibitedaclearpreferencefortheagriculturalarea, accountingfor69%(N=359)ofthetotalindividuals.Basedonthisand its presence during the entire sample period, this species is typical of disturbedareas.Inparticular,E.hermesmayalsobeindicativeofrecently alteredsites. T. penela was only recorded during the last three months of the study with July andSeptember peakpopulation periodsfor the species. Forthesereasons,T.penelaisnotanappropriateindicatorspecies. InGrandeRivieretheleastdisturbedarea(F)producedthehighestH value(2.48),andthemoredisturbedareashadlowerHvalues.However, this was not the case for both undisturbed areas used by Wood and Gillman(1998). Thetrapsintheirundisturbedevergreensites(UE)producedanHof 2.01, and traps in the disturbed evergreen (D E) sites produced an H of 1.78.ThisfollowedthefindingsatGrandeRivierewhereleastdisturbed areas had higher H value than disturbed areas. In contrast, undisturbed semi evergreen (U SE) areas produced an H value of 1.68, compared to the2.24valueforHinthedisturbedseasonalevergreen(DSE)areas.This findingisinconsistentwiththeprevioustwoobservations.Forbothforest types, undisturbed areas had higher numbers of C. minor than in their disturbedareas.Inthiscase,lightsensitivityofthisspecies(Frederiksen and Warrant, 2008) may play a key role in limiting their numbers at disturbed sites. However, almost the same number of M. peleides occurredineitherarea,andtoofewP.fusimaculatawerecaughttomake anyinferences.Ontheotherhand,E.penelopeandE.hermesweremore common in disturbed areas. This supports the inconclusiveness of rapid assessmentofbutterflyindicatorspeciesforassessingundisturbedareas.

38

WoodandGillman(1998)conductedarelativelyrapidassessmentof butterfly diversity, given that their 10 days of sampling occurred over a twentyday period. Since their H results did not consistently show that undisturbedareashavehigherHvaluethandisturbedareas,thissuggests thatrapidassessmentofbutterflydiversityusingShannonsIndexalone, wasnotsuitabletoidentifyareasofdisturbance. When Wood and Gillmans (1998) rapid assessment is compared to Barcants (1970) list of proposed butterfly indicators (Table 5), three of thefivespeciesappeareffective.
Table5OccurrenceofproposedbutterflyindicatorspeciesmodifiedfromWood andGillman(1998)andBarcant(1970). SpeciesName NumberofIndividuals DisturbedArea UndisturbedArea Barcant(1970) WoodandGillman(1998) SE E SE E INDICATORSOFLEASTDISTURBANCE P.fusimaculata P.hyalinus 0 1 0 0 M.peleides 3 9 6 5 C.minor C.eurilochus 2 0 6 2 INDICATORSOFMOREDISTURBANCE E.penelope Cissiapenelope 51 16 43 10 E.hermes C.hermes 6 1 1 0

Inthepresentstudy,itwasnotpossibletoobtaindataforafullwet season and a full dry season, which would have allowed for a more completeanalysis.Itwasalsolimitedtoareasinthelandscapethatwere generally safe and accessible. Ideally, larger areas of cocoa and the agricultural plots should have been sampled. This would reduce the contributionofthesource/sinkdynamicsfromnearbyhabitatsaffecting the results. Finally, due to manpower limitations it was not possible to includePollardWalksduringthisstudy. Conclusions ItwasfoundthatHF>HC>HA,andthatHisgreaterinlessdisturbedareas thaninmoredisturbedareasatGrandeRiviere.Variationinthemonthly HwasobservedfromMaytoSeptember.However,theresultsfromthis studyshouldnotbetakenasconclusivefromaseasonalorannualcycle. Where C. minor is relatively more abundant, such area(s) may be consideredleastdisturbed,whileareaswhereE.penelopeandE.hermes are relatively more abundant may be considered more disturbed. Rapid

39

assessment of butterflies, even when coupled with the use of indicator species,cannotbeusedtodeterminedisturbancelevelsinanarea. References
Barcant,M.1970.ButterfiesofTrinidadandTobago,1stedn.London:Collins. Beck. J., Schulze. C., 2000. Diversity of fruitfeeding butterflies (Nymphalidae) along a gradientoftropicalrainforestsuccessioninBorneowithsomeremarksontheproblemof pseudoreplicates,TransactionsoftheLepidopteraSocietyofJapan51:8998. Brown.K.S.,1997.Diversity,disturbance,andsustainableuseofNeotropicalforests:insects asindicatorsforconservationmonitoring.JournalofInsectConservation1:2542. Brown, K.S., 1991. Conservation of neotropical environments: insects as indicators, In The Conservation of insects and their habitats, 15th Symposium of the Royal Entomological SocietyofLondon,pp.50404,NewYork:AcademicPress. Clark. J. A., and May. R. M., 2002. Taxonomic bias in conservation research, Science 297: 191192. Davis, A. J., Holloway, J. D., Huijbregts, H., Krikken, J., KirkSpriggs, A. H. and Sutton, S. L. 2001.DungbeetlesasindicatorsofchangeintheforestsofnorthernBorneo.Journalof AppliedEcology,38:593616. DeVries. P.J, Murray. D., Lande. R., 1997. Species diversity in vertical, horizontal, and temporaldimensionsofafruitfeedingbutterflycommunityinanEcuadorianrainforest, BiologicalJournaloftheLinneanSociety62:34364. EggletonP,BignellDE,SandsWA,WaiteB,WoodTG,LawtonJH.,1995.Thespeciesrichness of termites (Isoptera) under differing levels of forest disturbance in the Mbalmayo ForestsReserve,southernCameroon,JournalofTropicalEcology11:8598 Ehrlich.P.R.,1984.Thestructureanddynamics ofbutterfypopulations,InThe Biology of Butterfies(R.I.VaneWrightandP.R.Ackery,eds),pp.2540,London:AcademicPress. Environmental Management Authority (EMA), 2010, National Register of Certificates of EnvironmentalClearance,accessedin2010. Erhardt. A., 1985. Diurnal Lepidoptera: sensitive indicators of cultivated and abandoned grassland,JournalofAppliedEcology,22:849862. Frederiksen. R., and Eric J. Warrant. J.E., 2008. Visual sensitivity in the crepuscular owl butterflyCaligomemnonandthediurnalbluemorphoMorphopeleides:acluetoexplain the evolution of nocturnal apposition eyes?, Journal of Experimental Biology 211, 844 851. Hill, J.K., Kramer, K.C., Lace, L.A., Banham. W.M.T., 1995. Effects of selective logging on tropicalforestbutterfliesonBuru,Indonesia,JournalofAppliedEcology32:754760. Kremen, C., 1992. Assessing the Indicator Properties of Species Assemblages for Natural AreasMonitoring,EcologicalApplications,Vol.2,No.2,pp.203217 Lawton.H.J.,Bignell.E.D.,Bolton.B.,Bloemers.F.G.,Eggleton.P,Hammond.M.P.,Hodda. M.,Holt,D.R.,Larsen.B.T.,Mawdsley.A.N.,Stork,E.N.,Srivastava,S.D.,andWatt,D. A., 1998. Biodiversity inventories, indicator taxa and effects of habitat modification in tropicalforest,Nature391,7276. Leather. S. R., Basset. Y., and Hawkins. B. A., 2008. Insect conservation: finding the way forward.InsectConservationandDiversity1:6769. Lovejoy, T.E., Bierregaard. R.O., Ryland, A.B., 1986. Edge and other effects of isolation on Amazonforestfragments,In:Soul,editor.Conservationbiology:thescienceofscarcity anddiversity,pp.257285.SinauerAssociatesInc. Mendz.M.,andC.Funes.,2007.InventariodeMariposasenSalamar,ColinasdeJucuarn, DepartmentodeUsutln,ElSalvador,SalvaNATURAInformedeConsultora.

40

Meyers.N.,Mittermeier.R.A.,Mittermeier.C.G.,daFonseca.G.A.B.,andKent.J.,2000. Biodiversityhotspotsforconservationpriorities,Nature403:853858. Nyamweya.N.N.,andGichuki.N.G.,2000.Effectsofplantstructureonbutterflydiversityin Mt. Marsabit Forest northern Kenya, African Journal of Ecology, 48: 304312. doi: 10.1111/j.13652028.2009.01151.x. Philpott.M.S.,Perfecto.I.,andVandermeer.J.,2006.EffectsofManagementIntensityand SeasononArborealAntDiversityandAbundanceinCoffeeAgroecosystems,Biodiversity andConservation,Volume15,Number1,139155. Rainio. J., and Niemel. J., 2002. Ground beetles (Coleoptera: Carabidae) as bioindicators, BiodiversityandConservation,Volume12,Number3,487506, Roth. S. D., Perfecto. I., and Rathcke. B., 1994. The Effects of Management Systems on GroundForagingAntDiversityinCostaRica,EcologicalApplications,Vol.4,No.3pp.423 436 Scheffler. Y. P., 2005. Dung beetle (Coleoptera: Scarabaeidae) diversity and community structure across three disturbance regimes in eastern Amazonia, Journal of Tropical Ecology,21,pp919 Sparrow. H.R, Sisk. T.D., Ehrlich. P.R., Murphy. D.D., 1994. Techniques and guidelines for monitoringneotropicalbutterflies,ConservationBiology8:800809. Spitzer. K., Novotn. V., Tonner. M., Lep. J., 1993. Habitat preferences, distribution and seasonality of the butterflies (Lepidoptera: Papilionidae) in a montane tropical rain forest,Vietnam,JournalofBiogeography20:109121. Sundufu. A., and Dumbuya. R., 2008. Habitat preferences of butterflies in the Bumbuna forest,NorthernSierraLeone.,JournalofInsectScience8:64. Vasconcellos. A., Bandeira. A.G., Moura. F.M.S., Araujo. V.F.P., Gusmao. M.A.B., Constantino.R.,2010.Termiteassemblagesinthreehabitatsunderdifferentdisturbance regimesinthesemiaridCaatingaofNEBrazil,JournalofAridEnvironments,74(2),pp. 298302. Watt. A. D., Stork, N. E., and Bolton. B., 2002. The diversity and abundance of ants in relation to forest disturbance and plantation establishment in southern Cameroon, JournalofAppliedEcology,39:1830. Wood. B., Gillman. M.P., 1998. The effects of disturbance on forest butterflies using two methodsofsamplinginTrinidad,BiodiversityandConservation7:597616. Wilson.E.O.,1988.Thecurrentstateofbiologicaldiversity.InBiodiversity,ed.EOWilson, pp.318.WashingtonD.C.,NationalAcademicPress.

41

Acomparisonofbeachmorphologyandphysicalcharacteristics ofTurtleBeach,TobagoandGrandeRiviere,Trinidadandits implicationsforturtlenesting 1,2 SheetalJankie andAndrewLawrence1


1

DepartmentofLifeSciences,TheUniversityoftheWestIndies,St.Augustine,Trinidad, WestIndies.Email:05773029@mysta.uwi.edu 2 CorrespondingAuthor

Abstract Globally, all seven species of marine turtles are threatened with extinction. Consequently, conservation efforts for these endangered speciesmustbeimproved.Oneapproachtoseaturtleconservationisby protection of their nesting habitat. Recent reports from Tobago have indicated a loss in stable nesting habitat at Turtle Beach, where sand mining and infrastructural developments have been suggested as the main contributors to habitat loss. This study investigated spatial and temporal changes in beach morphology and sediment composition of TurtleBeach,TobagoandGrandeRiviere,Trinidad,duringJuneOctober 2010. These data were compared to previously reported characteristics for these sites, to determine whether there were changes in these parameters and how the beaches differed from each other. Spearman rankcorrelationrevealedanegativecorrelationbetweenbeachareaand nesting activity at Turtle Beach. Research has suggested that turtles prefer to nest on smaller secluded bays and this correlation supported that theory. A KruskalWallis test revealed that the sediments on Turtle Beach were more coarse grained (0.5mm1.00mm) than they were 10 yearsagowithsignificancevaluesofH=7.02,P=0.008.In2000,sediments on Turtle Beach were classified as medium to fine grained (0.25mm 0.5mm)(0.025mm0.25mm). A KruskalWallis test also revealed that there was a significant difference (H=4.09, P=0.04) in Grande Rivieres grainsizecomposition(2010),whencomparedtoanalysesdonein1999. The beach however, remains classified as coarse grained. Other factors thatmayaffectnestingactivityincludeanthropogenicinfluences,photo pollution, changes in beach dynamics and bathymetry coupled with climatechangeandincreasedpoaching. Keywords Beach profiles, sediment composition, anthropogenic disturbance, particlesize,seaturtles,habitatpreference

42

Introduction Of the seven species of marine turtles in the world, three are listed as criticallyendangeredontheInternationalUnionfortheConservationof Nature (IUCN) Red List of Threatened Species. In addition, all seven speciesarelistedinAppendixIoftheConventiononInternationalTrade inEndangeredSpeciesofWildFaunaandFlora(CITES)(AbreuGlobois& Plotkins 2008). Marine turtles therefore, must be protected and conservationstrategiesfortheseanimalsstrengthenediftheirlongterm survivalistobeensured. Beaches play important roles in ocean ecosystems by providing key habitat for marine life, helping to balance marine food webs and facilitatingnutrientcyclingfromwatertoland(Wilsonet.al.n.d.).These habitatsalsoserveascriticalnestingsitesformarineturtles. The marine turtle species that most frequentlynest in Trinidad and TobagoareLeatherbackturtles(Dermochelyscoriacea),Hawksbillturtles (Eretmochelys imbricata) and Green turtles (Chelonia mydas) (Lalsingh 2008). Grande Riviere Bay in Trinidad, has the second highest recorded nestingfrequencyofleatherbackturtlesintheCaribbean(LeeLum2005). InTobago,TurtleBeachisalsoanimportantbeachfornestingseaturtles (Clovis, 2006) with approximately two thirds of all nesting activity in Tobagotakingplaceatthatbeach(Lawetal.2010). TherehavebeenrecentreportsofdecliningnestingactivityonTurtle Beach and it has been suggested that beach sand mining coupled with changes in the beach structure and morphology are responsible for this decline (Butler 1998). Beach sand mining in Tobago is still prevalent despite being illegal (Lalsingh 2010). Nesting turtles have also increased thepopularityofTurtleBeachasatouristattraction,andthishasledto an increase in infrastructure developments, anthropogenic influences, and modifications along the beach. This type of anthropogenic development may directly affect the structure and morphology of beaches, reducing stable nesting habitats and posing further threats to thespecies(Butler1998). Hawksbillturtlesprefertonestonlowenergy,steepbeachesbutwill alsousehighenergybeaches(EarnstandLovich2009).However,steep, highenergy beaches are thought to be preferred by leatherbacks and loggerheadsastheyreducetheamountofenergyrequiredforterrestrial locomotion by gravid females and hatchlings (Pritchard 1971). Hendrickson and Balasingam (1966) report that in Malaya, leatherbacks preferredtonestonbeacheswithpronouncedslopesandcoarsersands. Inaddition,ahighcorrelationbetweensedimentparticlesizeandmean

43

elevation of beach was noted by these authors (Hendrickson and Balasingam1966). While, up to 18,000 turtle nests have been recorded at Grande Riviere Bay in Trinidad in one nesting season (Wildlife Section, unpublisheddata),only300nestsofthisspecieshavebeenrecordedat Turtlebeach(Clovis,2005).TurtleBeachandGrandeRiviereBayareboth highenergy beaches (IMA 2004, Forestry Division et al. 2010). Both are steep beaches, with Turtle Beach spanning a greater area, and each beach hosts the highest number of nests on their respective islands. Although Turtle Beach provides a greater expanse of turtle nesting habitat, it is much more easily accessible than Grande Riviere. Grande Riviere is more isolated and less subjected to exploitation, which may makeitamoresuitablenestinghabitat. Thisstudycomparedthedifferencesinbeachsedimentcomposition, morphology, and physical characteristics of Turtle Beach and Grand Rivieretoassesshowthesefeaturesmayinfluencethesuitabilityofeach beachfornestingmarineturtles. Methods StudyAreas Turtle Beach, also known as Great Courland Bay, is 1.8km long and is locatedonthesouthwesternendofTobago.GrandeRiviereisnestledin the northeastern corner of Trinidad and spans approximately 1.3km. Bothbeachesareboundbyriversthatseasonallybursttheirbermsand flowintothesea.Atotalof1.1kmofTurtlebeachand0.65kmofGrande Riviereweresurveyedduringthisstudy(Figure1.0). GrandeRiviere Turtle Beach
Figure1.0LocationofGrandeRiviereBay,TrinidadandTurtleBeach,Tobago.

44

Beachprofiles GrandeRiviereandTurtleBeachwerestudiedbetweenJuneandOctober of 2010. Each beach was topographically surveyed and sediments were collected for analysis. Twentytwo beach profiles were performed on Turtle Beach, including 3 at permanent benchmarks previously established by the Institute of Marine Affairs (IMA, unpublished a). Thirteen beach profiles were performed for Grande Riviere. The first 12 profiles were 50m apart and included three IMA benchmarks. A thirteenth profile was performed at a 4th benchmark (IMAGR1), 250m awayfrombenchmark12.BenchmarksonTurtleBeachweresurveyedby theIMAfrom19992008whereasthoseonGrandeRiviereweresurveyed from19902008. Profiling was conducted using methodologies adopted by the IMA (IMA,unpublishedb).ProfilesweremeasuredusingaSokkiasurveylevel, a survey staff extendable to 4m, and a measuring tape. Profiles were performedoneachbeachat50mintervals.WheretheIMAsbenchmarks were within 2m of the 50m interval location, these were surveyed instead.Measurementsweretakenatthevegetationline,high,midand the low shore positions. Beach profiles were conducted at low tide to capturethemaximumtransectdistance. For each profile, the maximum elevation, and gradient were determined. These were tabulated against the same parameters from previously conducted profiles and analysed using Spearman Rank correlation tests. KruskalWallis tests were carried out using Statistix 7 (Analytical Software, 2000) statistical software to determine whether there were any significant changes in each parameter within and betweenbothbeaches. Sedimentanalysis Sediment samples were collected monthly at the upper, mid and low shore marks along each transect, using an 11.25cm x 20cm PVC corer. Approximately 8001000g of each sample was extracted using random bulk sampling. Sediment analyses were conducted using ASTM D6919 standard methods for particle size distribution of sediment using sieve analysisandsandgrainsizeanalysis(Barzani2009,Zeeman2009).These techniquesweresupplementedbymethodologiesoutlinedinKnodeletal (2007),andcharacterisedusingFolkandWards(1957)classificationfor skewness, kurtosis and sorting. Samples were air and sun dried, and samples taken at the low shore were oven dried at 100C for 24 hours. Driedsampleswereweighedandsievedintosixfractions,usinganASTM

45

mechanicalsieveshakerfor10minutes.Thefractionsineachsievewere calculatedasapercentageofthetotalsampleweight. Foreachprofileandsamplingpoint(upper,midandlowshore), anaverageofthepercentagegrainfractionswasderivedfortheperiodof study. Cumulative frequency curves of percentage grainsize versus phi ()valueswereplotted,andmean,median,skewness,kurtosis,standard deviation and sorting were determined using Minitab. Here, Phi representsthenegativelogarithmtothebasetwo,ofthevariousparticle sizes expressed in mm (Bowen 1986). These results were compared to sediment analyses previously conducted by the IMA using nonmetric Multidimensional Scaling Plots in Primer, KruskalWallis tests in Statistix 7, descriptive statistics and cumulative frequency plots in Minitab. The sedimentswereclassifiedaccordingtotheUddenWentworthGrainSize ClassificationScheme(Knodelet.al2007). NestingData Nesting data was acquired from Save our Seaturtles (SOS) Tobago and the Wildlife Section, Forestry Division of the Ministry Housing and the Environment.UnpublishednestingdataforGrandeRivierewereobtained from the Wildlife Section. These data were also used to comparatively assessdifferencesinnestingactivitybetweenbothbeaches.Nestingdata wereusedincorrelationanalysestoestablishpossiblelinkagesbetween beachconditionandnesting. Results Profiling KruskalWallistestsbetweenprofiledatafromthisstudy(2010)anddata fromIMA(2000&1990)respectivelyforTurtleBeachandGrandRiviere revealed significant changes over time in beach width, maximum elevation, and area. However, the tests indicated no significant changes overtimeinthegradientofeitherbeach. When profile parameters (2010) of both beaches were compared usingKruskalWallisandMultidimensionalScalingPlots,itwasfoundthat there were significant differences in the morphology of the beaches (area: H=28.28, p<0.001; width: H=31.85, p<0.01; maximum elevation: H=47.9,p<0.01;Gradient:34.75,p<0.01).TurtleBeachoccupiesagreater area,iswiderandgentlerinslopethanGrandeRiviere.Duringthisstudy, TurtleBeachwasalsofoundtoaccretetohigherelevationsthanGrande Riviere. Throughout the period JuneOctober 2010, both beaches underwentchanges,whichincludedchangesinarea,elevationandwidth, whichsuggestthattheyareverydynamicandhavedifferentpatternsof

46

sediment erosion and accretion. Rivers emptying through the beaches contributedtothedynamicnatureofbothbeaches. SedimentAnalysis Sediment analyses suggest that the mean grain size of Turtle Beach has changed over time. Turtle Beach now comprises of larger sand grains thanitdid10yearsago(Table1a)andiscoarsegrained.Itsmeanparticle size composition ranges from 0.3 to 0.8mm. In 2000, sediments on this beachweremediumgrainedandmoderatelysorted.By2010,sediment analyses revealed that the beach was medium to coarse grained and moderatelywellsorted. Grande Riviere now comprises of slightly larger sediments when compared to sediment analyses conducted in 1999 (Table 1b). It has a mean particle size distribution, which ranges from 0.6mm to 1.1mm. There were significant changes also in Grande Rivieres mean grain size composition. The beach is now well sorted and coarse grained throughouttheentirebeachasopposedtobeingwelltomoderatelywell sorted, and ranging from medium to coarsegrained 11 years ago. Non metric multidimensional scaling (nMDS) plots revealed two distinct groupsofsamplesindicativeofdifferencesingrainsizebetweenGrande Riviere and Turtle Beach as well as similarity in mean grain size among sites within each study area indicating near uniform distribution of sedimentsacrosseachbeach.Outliersoftheseanalysesforbothbeaches werefoundwhererockyoutcropswerepresent. 1a 1b
Table 1. Changes in sediment mean grain size composition of 1a) Turtle Beach Tobago from 20002001 and 1b) Grande Riviere from 19992001. KEY MWS moderately well sorted; WS well sorted; Ppoorly sorted; FGfine grained; MG mediumgrained;CGcoarsegrained;VCGverycoarsegrained

47

NestingData The number of turtles nesting reported on Grande Riviere appears far greaterthanthenumberofturtlesreportednestingonTurtleBeach.The maximum number of nests recorded in Turtle Beach is 300 in 2005 (Clovis,2005).Sincethenthereappearstohavebeenaslightdeclineto the present number of 271 (Clovis, 2005). This is much lower than the 18,000 nesting turtles recorded in Grande Riviere in 2005 (Wildlife Division, unpublished). However, no data were available for nesting on TurtleBeachin1999orGrandeRivierein2009and2010andthefigure for2008representsonlythenumberofturtlesthatweretaggedatthat beach. Discussion Turtle nesting data collected by SOS and the Forestry Division indicate that turtles nest at much higher densities on Grande Riviere, Trinidad, than Turtle Beach, Tobago. Nesting data shows that there has been a temporal increase in the number of nesting turtles on both beaches. However,priorto2000monitoringstrategiesinGrandeRivierewerenot assystematic(Livingstone&Downie2004)asthosecurrentlyemployed bythemonitoringinstitutions,andsomaynotbeasaccurate(Livingstone 2004).Asasmallenvironmentalgroup,SOSTobagoonlybegancollecting datain2000(Clovis,2007). This study revealed that both beaches have undergone extensive morphological change over time. This is probably due to natural and seasonal patterns of sediment erosion and accretion in each area (Lee Lum2005).ThesedimentgrainsizecompositionofTurtleBeachhasbeen altered from mediumgrained sand to coarsegrained sand. A coarser beachmeansthatfinerparticleshavebeenwashedaway,leavingbehind the larger, coarse grains (Pinet 2003). The composition of coarser sediments may be associated with erosion of the beach (IPCC 2007). Erosionisknowntobecorrelatedwithwaveenergy(Bird2008),aswell as sea level rise (IPCC 2007). The Intergovernmental Panel on Climate Change (IPCC 2007) and Singh (1997 a,b) have indicated that beach erosion rates have been high in Trinidad and Tobago over the last 15 years and sealevel rise is considered a contributory factor (Mc Leod 1999,Singh,1997a,b). Erosionandtheremovalofsandoftenleadtosignificantchangesin beach morphology and slope, and where the rate of removal exceeds natural replenishment, a negative sediment balance occurs (Cox & Embree 1990). Sediment particle size was found to be positively

48

correlated to beach elevation, gradient, and width in this study. According to Bird (2008), larger grains tend to accrete better than finer grains.Inaddition,HendricksonandBalasingam(1966)reportedthatthe steepness of beach slope is correlated to grain size. Consequently, this indicates that the observed changes in beach structure reported here mayberelatedtothesedimentchangesrecordedinthisstudy. In contrast to green turtles and hawksbills, which tend to nest in vegetated areas where trees shade the sand and keep nests cool (Horrocks & Scott 1991), leatherbacks nest on open sands (Hendrickson andBalasingam1966).Theporosityandabilityofanimalstoburrowinto a sandy beach are affected by the sediment particle size. Fine sand beacheshavegreaterwaterretentionwhereascoarsesandbeachesdrain well and dry out quickly (Karleskint et.al 2010). Grain size determines characteristics such as heat retention, compaction, water retention, aerationandporosity(Knodeletal.2007). Knodel et al. (2007) also reported that the degree of sorting in sedimentsaffectcharacteristics,suchasporosityofthesand.Sortingisa measureoftheevennessofsandparticledistribution.Wellsortedsandis more porous, aerated, well drained and has better heat regulation (Pfannkuchundated,Xieetal.2010).Turtlesgenerallyselectundisturbed beachesthatarereasonablysteepwithsandthatisneithertoofinenor compact(Mc.Lachlanetal.2006).Thesedimentanalysesconductedhere indicated that Grande Riviere has better sorted sediments than Turtle Beach, is fineskewed and near symmetrical. Turtle Beach on the other handwasfoundtobecoarseskewed,andmoderatelywellsorted.Coarse skewedsandhasmoresedimentfinerthanitmeangrainsize(Schwartz 2005). In this study, it was found that the area of the beach was not a limiting factor for nesting on Turtle Beach, which is longer, wider and covers a greater area than Grande Riviere. In addition, the sediment analyses indicated that the composition of Turtle Beach has changed to thosemorefavouredbynestingturtlesandyetthenumberofnestshas been relatively stable/slightly reduced over the last 5 years. No correlations were found between grain size composition and nesting activitieswhenSpearmanRankcorrelationtestswereperformedondata from either beach. Therefore, the hypothesis that changes in turtle nesting activities are due to sediment changes cannot be statistically confirmedfromthisstudy. However,grainsizecandeterminetheshapeandslopeofbeaches (Karleskint et al. 2010). Studies have revealed that Hawksbill turtles prefer to nest on steep, narrow beaches (Earnst and Lovich 2009).

49

Leatherbacks have an affinity for nesting on coarsegrained beaches (HendricksonandBalasingam,1966).Wellsortedsandalsoseemstobe favouredbyloggerheadturtles(Karavasetal.2005).Consequently,these turtles seem to select suitable nesting sites in terms of optimum conditionsprovidedbybeachsands(Demetropoulos2001).Indeedithas been suggested that a combination of interacting ecological factors including temperature, particle size, water content, salinity, sand softness,lagoonpresence,beachlength,andbeachheight,influencenest siteselection(VarelaAcevedoetal.2009). Changes in sediment composition may, therefore, affect the suitability of Turtle Beach as a nesting site. However, differences in nestingactivitiesbetweensitesmightnotbelimitedtothecharacteristics of the beach alone but may also be due to geographic, demographic, historical, natural and anthropogenic factors. Consequently, a more detailedstudyonpropertiesofthesediments,otherfactorsandnesting patterns is required to explain trends in nesting activities recorded for bothbeaches. Grande Riviere is a smaller, narrower, steeper beach with larger, bettersortedsandgrainsthanTurtlebeach.Thesecharacteristicsmay,in part, be responsible for the differences in nest densities between both beaches. Finegrained poorly sorted sand allows for compaction and water logging of beaches (Knodel et.al 2007). Sea turtles may nest on coarse grained, well sorted sands because it allows more ease in constructing nest chambers whilst at the same time being easily manipulated by emerging hatchlings (Mc Lachlan et al. 2006). Coarse, wellsorted,sandisnoteasilycompacted(Shatzel&Anderson2005)and thisisanadvantageforemerginghatchlingsasitmakestheirjourneyless strenuous. Coarser sands also allow better drainage, reduce water logging of nests, are more permeable, and so increase aeration of the sandandoxygenflowtotheeggs(Knodeletal.2007,Shatzel&Anderson 2005,Xieetal.2010.Thetemperatureofsandisbetterregulatedwhen thereisincreasedaeration.Notonlydoestemperatureaffectthesexof the hatchling, it also affects hatchling success and egg mortality. Successful incubation of turtle nests is possible within specific thermal limitsandisinhibitedbelow25Candabove35C(Gallegosetal.2009). Hightemperaturescanresultinhatchlingandnestmortality(Gallegoset al.2009).Thesecharacteristicsinfluencenestsiteselectionbyturtlesand iftheconditionsarenotfavourable,femalesmaymakefalsecrawlsand returntotheseawithoutnesting. Steeper beaches are preferred by turtles because it makes their ascenttostablenestinghabitatandthedescentoftheirhatchlingseasier

50

(McLachlanetal.2006).Leatherbacknestsareparticularlyvulnerableto erosion because the turtles' great size and tender skin force them to choose high energy, accessible beaches with a steep slope, which prevents them from travelling far inland to lay their eggs (Butler 1998). Steeper beaches also offer more protection to nests from wave action andarebetterdrainedthangentlyslopingbeaches(Anthoni2000),thus preventingwaterloggingofnestandmortalityofeggs. It has been noted that sea turtles prefer to nest on small beaches, and this is perhaps one of the reasons why so many turtles nest on GrandeRiviere.Smallbeachesinsecludedareasmaybeoptimalasthey are prone to fewer disturbances and are more sheltered. Open access beaches give room for human interference as is the case with Turtle Beach. Other factors that may potentially impact the nesting activities on TurtleBeacharemainlyanthropogenic.Itwasnotedthattherehasbeen an increase in developments along the beach as well as habitat modificationsbybeachfrontdcor.BoatingactivitiesaroundTurtleBeach aremuchmoreprevalentthaninGrandeRiviere.StudiesbyBacon(1973) inMatura,Trinidadconfirmedthatlightcausesdisorientationinnesting sea turtles. Due to the Turtle Beach hotel, photopollution and other activitiesmayactasadeterrenttonestingturtles. Leatherbacks almost never nest on beaches protected by reefs, as contactwiththereefsystemsatlowtideespeciallyinaroughseamaybe dangerous for this softskinned species (Pritchard 1971). Turtle Beach is one of the few beaches is Tobago not protected by reef systems. However, neighbouring beaches in Tobago are protected by reefs. This mayalsobeapossiblereasonfortheconsiderablylowernestingactivities onTurtleBeachwhencomparedtoGrandeRiviere.Trinidadhasfewreef systems,noneofwhichareinproximityofGrandeRiviere.Onlyonereef system,theSalybiaReef,isrecordedinTrinidad(Kenny2008). More indepth and extensive research is required to provide conclusive insight into the patterns of nesting being observed on Turtle BeachandGrandRiviere.Anthropogenicactivitiesareconsideredtohave significantimpactsonturtlenestingandthismaybeaparticularproblem forTurtleBeach.Assuch,researchandimplementationofmanagement andregulatorystrategiesshouldbeaddressedwithurgency. Conclusions The morphology and sediment composition of both Turtle Beach and GrandeRivierehavechangedoverthepast20years.Reductionsinbeach width, elevation and area have been noted for both beaches and

51

sedimentanalysesrevealedlargermeangrainsizecompositionatTurtle Beachthannoted10yearspreviously.Specifically,thisbeachwasfound to be medium to coarse grained, and moderately well sorted. Grande Rivierehascoarsegrainedsandthroughouttheentirebeachandchanged frombeingmoderatelywellsortedtowellsorted.Thesechangesmaybe related to natural and seasonal patterns of sediment erosion and accretion. The changes may also be enhanced by changes in beach dynamics and bathymetry as a result of sea level rise associated with climatechange. Both beaches are steep, highenergy locations, which are characteristics preferred by nesting turtles. More turtles however, nest onGrandeRivierethanonTurtleBeach.GrandeRiviere,whencompared to Turtle Beach, has higher energy, is narrower and steeper, and has coarser,bettersortedsands. In addition to morphological and sediment changes, other factors, not examined in this study, may affect nesting activities. These include presence of vegetation, infrastructure, photopollution, pollution from runoff and litter. Much more research, therefore, is required to determinethecausefordifferencesinnestingactivitiesbetweenthetwo beaches. Acknowledgements Aboveall,IthankGodforguidingmewithhisgraceandglory.Ithankfor theirsupport,myparentsandfamilywhoworkedhardalongsidemeas my team on the field, and Mr. Shiva Heerah who lent support and assistance in the analyticalcomponent of this research. I also thankthe Institute of Marine Affairs, namely Mr. Lester Doodnath and Mr. Sunil Ramnath. Many thanks to the Wildlife Division, SOS Tobago and Mr. Giancarlo Lalsinghall for providing me with data that was vital to the completion of this research. Many thanks are extended to all the Life Sciences and Soil Sciences staff who assisted especially the laboratory technicians;Raj,DexterandAnton,Dr.Wuddivira,Dr.Eudoxie,Professor ShawandMs.Atwell. References
AbreuGrobois,A.andPlotkin,P.2008.Lepidochelysolivacea.In:IUCN2010.IUCNRedList ofThreatenedSpecies.Version2010.4.Accessedfromwww.iucnredlist.org23/11/2010 AnalyticalSoftware.2000.Statistix7.Tallahassee,Florida,USA. Anthoni.J.F 2000. Disappearing Beaches: drying in the tide accessed from http://www.seafriends.org.nz/oceano/beachdry.htm(17.11.2010) Bacon.P.R.,1973.TheOrientationCycleinthebeachAscentCrawloftheLeatherbackTurtle Dermochelyscoriacea,inTrinidad.Herpetologica,Vol.29,pp343348

52

Bird. C.F. Eric, 2008.Coastal Geomorphology: An Introduction 2nd Edition, John Wiley and SonsLtd.England.pp174 Butler.K.R.,1998.CoastalProtectionofSeaTurtlesinFlorida.FloridaStateUniversityJournal of land Use and Environmental Law accessed from http://www.law.fsu.edu/journals/ landuse/vol132/butl.htm Clovis, T.2005. SOS Tobago Annual Report March November 2005, Save Our Sea Turtles (SOS),Tobago Clovis, T., 2006. Save our Sea Turtles Summary Report 2006, Save our Sea Turtles (SOS), Tobago Demetropolos. A.. 2002 ed. Colloquy on Marine and Coastal Ecological Corridors; proceedingsLlandudno2021June2002.CouncilofEuropePublishing,pp98 EarnstC.,LovichJ.,2009.TurtlesofUnitedStatesandCanada,TheJohnHopkinsUniversity Press,pp87 FAO,2004.Paperspresentedattheexpertconsultationoninteractionsbetweenseaturtles andfisherieswithinanecosystemcomplex,FAOFisheriesReportNo.738 Lancaster.N,1995.GeomorphologyofDesertDunes.Routledge,London.pp105 Gallegos.J.,Fish.M.,Drews.C.2009.TemperatureMonitoringManualWorld WildlifeFund (WWF),Central America accessed from http://assets.panda.org/downloads/ temperature_manual_one_pager__final__1.pdf(20.10.2010) Hendrickson and Balasingam, 1966 in Pritchard. P.C.H.1971. The Leatherback or Leathery TurtleDermochelyscoriacea.IUCNSwitzerland.pp1720 Horrocks, J.A. and N.M. Scott, 1991. Nest site location and nest success in the hawksbill turtle Eretmochelys imbricata in Barbados, West Indies. Marine Ecology Progress Series 69:18 Institute of Marine Affairs (IMA). 2004. A Guide to Beaches and Bays of Trinidad and Tobago,.InstituteofMarineAffairs,pp8087 InstituteofMarineAffairsGrainSizeanalysesandProfileData(unpublished) Karavas.N., Georghiou.K.,Arianoutsou.M.,Dimopolous.D.,2005.Vegetation and Sand CharacteristicsinfluencingnestingactivityofCarettacarettaonSekainiaBeach,Biological Conservation121,pp177188 Karleskint.G.,Turner.R.,Small.J.2010.IntroductiontoMarineBiology.Brooks/ColeCengage Learning.U.S.A.pp375 Kenny.J.S., 2008. The Biological Diversity of Trinidad and Tobago: a naturalists notes. ProspectPress,PortofSpain.pp164. Kndel. K., Lange. G., Voigt HansJrgen. 2007. Environmental geology: handbook of field methodsandcasestudies.SpringerVerlagBerlinHeidelberg,pp710724 Lalsingh G. 2008 Summary Report of marine Turtle Nesting Activity 2008 Save our Sea turtles(SOS)Tobago, LalsinghG,2010.SummaryofMarineSeaTurtleActivity2010.SaveourSeaturtles(SOS). Tobago Law. A, Clovis, T, Lalsingh. G.R. and Downie J. R 2010 The Influence of Lunar, Tidal and Nocturnal Phases on the Nesting Activity of Leatherbacks (Dermochelys coriacea) in Tobago,WestIndies.MarineTurtleNewsletter2010. Lee Lum.L, 2005. Beach dynamics and nest distribution of the leatherback turtle (Dermochelys Coriacea) at Grande Riviere Beach Trinidad and Tobago, International JournalofTropicalBiology53:239248. Marine Turtle Specialist Group 1996. Caretta caretta. In: IUCN 2010. IUCN Red List of ThreatenedSpecies.Version2010.4.accessedfromwww.iucnredlist.org(23.11.10) Mc. Lachlan. A., Brown. A.C., Brown, Alexander.C.,2006. The Ecology of Sandy Shores. ElsevierIncorporated,U.S.A.pp215218

53

MinistryofLand,HousingandtheEnvironment,WildlifeSection,19992007.NestingData (unpublished) NationalResearchCouncil(U.S.)CommitteeonSeaTurtleConservation,1990.Declineofthe seaTurtles:CausesandPrevention.NationalScienceAcademy Pfannkuch.H.O.,Paulson,R.n.dGrainsizedistributionandhydraulicproperties accessed from http://faculty.njcu.edu/wmontgomery/Coastal_Zone/ Grain%20Size%20Distribution.htm(01.09.2010) Pinet.R.Paul,2003.Invitation to Oceanography 3rd Edition. Jones and Bartlett Publishers, London,U.K.pp368 Pritchard. P.C.H.1971. The Leatherback or Leathery Turtle Dermochelys coriacea. IUCN Switzerland.pp1720 Sand Grain Size Analysis. University of New England. Accessed from http://faculty.une.edu/cas/szeeman/oce/lab/sediment_analysis.pdf(21.06.2010) Seminoff, J.A. 2004. Chelonia mydas. In: IUCN 2010. IUCN Red List of Threatened Species. Version2010.4.Accessedfromwww.iucnredlist.org>(23.11.2010) ShatzelandAnderson,2005.Soil:GenesisandGeomorphology,UniversityPress,Cambridge, pp17 Schwartz.M.L.2005EncyclopaediaofCoastalScience,Springer.Netherlandspp178 Singh, B., 1997a: Climaterelated global changes in the southern Caribbean: Trinidad and Tobago.GlobalandPlanetaryChange,15,93111. Singh, B., 1997b: Climate changes in the greater and southern Caribbean. International JournalofClimatology,17,10931114. VareldaAcevedo.E., Eckert K.L., Eckert.S. Cambers.G., Horrocks.G.A.,2009. Sea Turtle Nesting Beach Characterization Manual, p.4697. In:Examining the Effects of Changing Coastline Processes on Hawksbill Sea Turtle(Eretmochelys imbricata) Nesting Habitat, MastersProject,NicholasSchooloftheEnvironmentandEarthSciences,DukeUniversity. Beaufort,N.CarolinaUSA.pp97 Wilson. E.G, Miller. K.L., Allison. D., Magliocca. M. n.d. Why healthy oceans need sea turtles: the importance of sea turtles to marine ecosystems. Accessed from http://na.oceana.org/sites/default/files/reports/Why_Healthy_Oceans_Need_Sea_Turtle s.pdf(06.12.2010) XieZ.,Wang.Y.,ChengG.,MalhiS.S.,VeraC.L.,Guo.Z.,Zhang.Y.,2010.Particlesizeeffects on soil temperature, evaporation, water use efficiency and watermelon yield in fields mulchedwithgravelandsandinsemiaridLoessPlateauofnorthwestChina.Agricultural WaterManagement,Vol97:6,pp917923

54

SpatialDistributionofintertidalbenthicmacrofaunainthree sandybeachesinTrinidad LanyaFanovich1,2,HowardP.Nelson2andAndrewLawrence2


2

DepartmentofLifeSciences,TheUniversityoftheWestIndies,St.Augustine,Trinidad, WestIndies.Email:lfanovich@gmail.com 1 CorrespondingAuthor

Abstract While sandy beaches in Trinidad and Tobago are important recreational areas, they are often perceived as ecological deserts, since most of the organisms in these habitats are inconspicuous. However, these beach habitatscanbebiologicallydiverse,andtheirfauna,whileinconspicuous, is potentially susceptible to anthropogenic disturbance. This study investigated the spatial distribution of the benthic macrofauna in the intertidal zone on the beaches of Salybia, Guayaguayare and Chagville, Trinidad,betweenJuneandAugust2010.Fivetransectsweresampledon eachbeach,at50mintervals.Benthicandsedimentsamplesweretaken at the high, mid and low tide levels on each transect. Descriptive and multivariateanalysesrevealedthatallthreebeachesvariedinthelength oftheirintertidalzone,slope,grainsizecompositionandpH.Ofthese,pH and grain size may have the greatest influence in the patterns found. Salybia and Guayaguayare were different from each other whilst Chagville shared biotic similarities with both these beaches. Species abundancewashighestatlowtideforGuayaguayareandChagvilleandat the driftline for Salybia. The deposit feeder and filter feeder guilds of polychaetes dominated each intertidal zone. Nereis sp. was the only carnivorepresentinthemidtidalzonesofSalybiaandChagville.Spionid polychaetes were the dominant species in the midtidal and low water markinGuayaguayareandChagville. Keywords grainsize,intertidalzonation,Nereis,pH,salinity,tropicalcoastlines Introduction Sandybeacheshaveoftenbeendescribedasecologicaldesertsdueto their lack of visible fauna and flora. However, a large percentage of a sandybeachsdiversityisfoundinthesand.Theseorganismsaresmall, inconspicuous and often found in high densities (Brown and McLachlan 1990, Jones et al. 2004, Schlacher et al. 2008, Defeo et al. 2009,

55

Pallewatta2010).Invertebratesintheintertidalzonemaybeclassifiedas eithermeiofaunaormacrofauna.Meiofaunaarethoseinvertebratesthat aresmallerthan0.5mmwhilemacrofaunaarethosespecieslargerthan 0.5mm (Jones et al. 2004, Pallewatta 2010). Polychaete worms, bivalve molluscs and crustaceans have been previously described as the dominantmacroinvertebratesofsandybeachesandtheirintertidalzones (Jonesetal.2004,NybakkenandBertness2005,Defeoetal.2009). Theintertidalzoneisatransitionbetweenthemarineandterrestrial ecosystem, and as a result is a harsh environment for many organisms due to the fluctuation of environmental conditions caused by tidal movement. As a result, this zone displays the highest variability in environmentalconditionscomparedwithanyothermarineenvironment (NybakkenandBertness2005). While, several authors have proposed intertidal zonation schemes (Jaramilloetal.1993,JaramilloandMcLachlan1995,Barrosetal.2001, DefeoandMcLachlan2005,NevesandBemvenuti2006),mostintertidal zonesexhibitsimilarcommunityassemblages.Unlikerockyshores,sandy shores do not have easily identifiable zones, and the distribution of macrofauna can be patchy. Particle size, wave action and slope are the important defining factors of sandy shore ecosystems, affecting distributionoforganisms(JaramilloandMcLachlan1993,Jaramilloetal. 1993, McLachlan 1996, Ricciardi and Bourget 1999, Nybakken and Bertness2005). It has been proposed that intertidal zonation allows organisms to avoid interspecific competition while feeding in the harsh conditions presentinsandyshores(Newell1979).However,thespaceprovidedby sandy shores makes overcrowding unlikely. In addition, motility allows the organisms to avoid interspecific competition unlike most sessile organismsonrockyshores(NybakkenandBertness2005).Mostspecies are opportunistic feeders rather than obligate feeders. As a result, competitionforfoodisunlikely(NybakkenandBertness2005). Most information on sandy beaches comes from research in the temperatezoneandinthecaseoftropicalcoastlines,thereisadearthof research in these areas, particularly the southern Caribbean (Gobin 2010). Published coastal benthic studies conducted in Trinidad and Tobagos sandy beaches rarely attempt to place the macrofauna into zones, and no published comparative studies of these beaches are known.Thisstudyaddressedthisinformationgap,bycomparingzonation in three sandy beaches. The observed zonation patterns were then comparedtothegeneralisedschemeproposedbyBrownandMcLachlan (1990)todetermineitsapplicabilitytoTrinidadandTobago.

56

Materialsandmethods StudySite ThreesandybeachesontheislandofTrinidadweresampledduringthe low spring tide. Salybia Bay, on the northeast coast of Trinidad, was sampled on June 7th to 8th, 2010. Guayaguayare Beach, on the south coast, was sampled between the 14th and 15th July 2010. The final sampling site was Chagville Beach located on the western coast, which wassampledbetweenthe8thand9thofAugust,2010(Figure1).

Figure 1. Location of Salybia, Chagville and Guayaguayare sandy beach sampling sites,Trinidad.(Source:AdaptedfromInstituteofMarineAffairs2004)

Salybia Bay is approximately 700m in length, and is exposed to the northeast trade winds and subjected to high wave energy. The eastern partofthebeachisprotectedbyafringingreef,andthebeachsediment is coarsegrained and composed of quartz and carbonate particles (Institute of Marine Affairs 2004, Office of Central Statistics, 2007). GuayaguayareBayisinfluencedbymoderateenergywavesandriverine outputfromthenearbySouthAmericanmainland.Itismorethan5kmin length and composed of brown finegrained quartz sand (Institute of MarineAffairs2004,OfficeofCentralStatistics2007).Liningthecoastline is a seawall approximately 1.5m in height and 30m from the lowwater mark,uponwhichmanyhousesarebuilt.

57

Facing the Gulf of Paria, Chagville is one of the beaches making up CarenageBayandlikeSalybia,isfrequentedbyvisitors.Itisa600mlong manmade beach composed of sand, gravel and broken coral. On the eastern side of this beach, the mangrovelined Cuesa River, empties freshwaterintothebay(InstituteofMarineAffairs2004,OfficeofCentral Statistics2007). Datacollection The methodology of Jaramillo, McLachlan and Coetzee (1993) was adapted for this study. Five transects, 50m apart were sampled at each beach, with random selection of the location of the initial transect. At each transect, topographical profiles were taken from the top of the scarp at each beach to the waterline at low tide (Neves and Bemvenuti 2006). The profiles were measured using a Sokkisha TM6 optical theodolite.GPSreadingsweretakenatsamplingstationsatthehightide driftline, the lowwater mark and midway between these two points usingaGarminGPSmap76ShandheldGPS. Three samples of benthic macroinvertebrates were taken at each station, on each transect, amounting to fortyfive samples from each beach. Samples were taken usinga15cmdiameter(0.02m2)PVCcorer,pushedtoadepthof25cm,or as deep as possible (Neves and Bemvenuti 2006, Schlacher et al. 2008). Thefirstsampleateachstationwastakenatthepointoftopographical measurement.Theremainingtwosamplesateachstationweretaken1m totheleftand1mtotherightofthefirstsample(NevesandBemvenuti 2006). The samples were then sieved using a 1.0mm metal mesh. The residuewastransferredtoZiplocbags,containing10%formalinfixative, to which Rose Bengal stain was added (Neves and Bemvenuti 2006, Schlacheretal.2008).Theorganismsfoundwereidentifiedtothelowest taxonomic unit and quantified to determine density of individuals per squaremeter(NevesandBemvenuti2006). Water temperature, salinity, pH and conductivity were recorded in triplicate at each transect, at the time of biological sampling, using a YSI63 meter. In addition, one sediment sample was collected at each sampling station for sediment analysis. The sample was collected as previouslydescribedbuttotheleftofthemiddlebenthicsample.Intotal, threesedimentsampleswerecollectedpertransectathigh,midandlow water.Inthelaboratory,thesedimentwasdriedinanovenfortwenty four (24) hours between 80C and 85C. The dried sediment was then sievedusinggraduatedsievesrangingfrommeshsizeof4mmto63m, such that six (6) sediment fractions were obtained. Each fraction was weighedtodetermineitsproportioninthesample.

58

Dataanalysis Analyses were performed using descriptive statistics and PRIMER v5 (Clarke and Warwick 2001) for multivariate analyses. A fourthroot transformationofbiologicaldensitydatawasconductedandBrayCurtis similarityandnonmetricmultidimensionalscaling(nMDS)analysesused todeterminerelationshipsbetweenthebiologicalcommunitiessampled (ClarkeandWarwick2001). Squareroottransformationwasinitiallyperformedonthegrainsize data and subsequently a principal components analysis (PCA) of all environmental variables was then undertaken. To test for colinearity among the environmental variables a draftsman plot was produced, in ordertoremoveanyvariableswhichwerehighlycorrelated.Inorderto linkthebiologicalpatternsfromthenMDStotheenvironmentalpatterns from the PCA, a BIOENV analysis was undertaken, to determine which combinationofenvironmentalvariablesbestexplainedthebioticpattern (ClarkeandAinsworth1993,ClarkeandWarwick2001). To determine horizontal and vertical zonation patterns within each beach,thedensitiesextrapolatedtoindividualspersquaremeter(ind.m 2 ) was used. Those species that were present in the highest densities amongthesamplingstationswereconsideredasdominantandtherefore representativeofthedifferentzones. Results Topographicalprofiles,grainsizeandbasicenvironmentalcharacteristics showed that each beach was highly variable. The steepest slopes were found at Salybia, whereas Guayaguayare was much flatter. Salybia also hadtheshortestintertidaldistancewhileGuayaguayaresintertidalzone was much longer than the other two beaches. Profiles of Chagville and Salybia were highly undulating unlike Guayaguayares almost even surface. Chagville was extremely uneven in the length of its intertidal zone,unliketheothertwosites. Sediment texture varied immensely between the three sites. The largestgrainsizefoundatall3beacheswasbetween501mand2.36mm (Figure 2), which is classified as very coarse to coarsegrained on the WentworthScale(InstituteofMarineAffairs2004).SalybiaandChagville had mostly cobble and coarsegrained sand, while Guayaguayare had a higherproportionofmediumtofinegrainedsand.

59

Proportion

0.600 0.500 0.400 0.300 0.200 0.100 0.000 0.100

Salybia Guayaguayare Chagville

Grainsize

Figure 2. Comparison of proportion of each grain size category from Salybia, GuayaguayareandChagvillebeaches,Trinidad

The grain sizecomposition at each tidal level also varied within the beaches. At midtide levels, coarse grain size greater than 501m, accounted for over 50% of the sediment at Guayaguayare (60.3%) and Chagville (81%). The high tide levels at both beaches were comprised mainlyofthemediumtofinergrainedsand.ChagvilleandGuayaguayare differed in the grain size proportions at the low tidal region, however, withChagvillepredominantlycobbly(76.1%),whereasGuayaguayarewas mostly medium to finegrained (59.4%). In contrast, Salybia had an almostequaldistributionofcoarsegrainedsedimentacrossallthreetide levels,withover80%ofthesedimentgreaterthan501m. Temperature,pH,conductivityandsalinityofthewaterwerelargely similarbetweenbeaches,withtheexceptionofsalinityandconductivity in Chagville in which there was a sharp decrease in both variables at Transect5. The PCA ordination of abiotic parameters showed tight clustering among the stations at each beach, suggesting that very little variation occurred within the sites at each beach (Figure 3). Chagville and Guayaguayares environment, while quite distinct from each other, still hadhigherlevelofsimilaritytoeachotherthantoSalybia.AtChagville, Transect5(C5)showeddeviationduetoextremelylowsalinity(0.13ppt) andconductivity(0.387ms).

60

Figure 3. Principal component analysis ordination of abiotic factors on each transectonsandybeachesatSalybia,GuayaguayareandChagville,Trinidad

Thetestofcolinearitybetweenenvironmentalvariables,revealedno colinearitybetweenanypairofenvironmentalparameters.
14 12 10 8 6 4 2 0 Salybia Guayaguayare Phyla Chagville

Numberof species

Figure4.Numberofspeciesofeachphylafoundintheintertidalzoneofthethree beachesatSalybia,GuayaguayareandChagville,Trinidad

Atotaloffortyonemacroinvertebratespecieswereidentifiedduring thisstudy.ThephylumAnnelida,representedbypolychaeteworms,was themostspeciesrichgroup(Figure 4),withmostspeciesbeingfoundin

61

Chagville. Only at Salybia was this group surpassed in richness by the arthropods,consistingofisopodsandamphipodswithelevenspecies,in comparison to ten species of annelids. Porifera and Cnidaria were also presentonlyatSalybia.

36.28

36.20

SALYBIA GUAYAGUAYARE CHAGVILLE

57.56

Figure 5 Comparison of the density of macroinvertebrates found at Salybia, GuayaguayareandChagvillebeaches,Trinidad(Density=numberm2).

Despite the presence of more phyla in Salybia, this beach only had 36.2individualsm2( Figure 5) in comparison to Guayaguayare which had fewer species but at a density of 57.56 individuals m2. The organisms exhibited a preference for the midtidal level and the lowwater mark at Chagville and Guayaguayare. However, Salybia had a higher density of macroinvertebratesatthedriftline(54.04indm2)andthelowestdensity wasatthelowwatermark(21.37indm2). Nonmetricmultidimensionalscaling(nMDS)analysisoftheintertidal macrobenthos showed that Salybia and Guayaguayare had distinct community assemblages whilst Chagville shared similarities with both other beaches (Figure 6). Specifically, the composition at the Chagville driftlinewassimilartothatatSalybia,whilethemidtidallevelandlow watermarkofChagvillewassimilartothatatGuayaguayare.Thespecies assemblagesatthedriftlineofallthreebeachesweremarkedlydifferent from the other tidal ranges in the intertidal zone. It can therefore be deducedthattherewereatleasttwozonespresentateachbeach,with verylittledistinctionbetweenthemidlevelandlowwatermark.

62

The BIOENV analysis suggested that pH and grain size had the highest correlation of (0.728) with the biotic data. Thus, these two environmental variables accounted for the majority of the pattern produced by the nMDS. The pH levels were mostly constant within the beachesbutdifferentamongthesamplingsites.ChagvilllespHwasmuch lowerthanthemorealkalinelevelsatSalybiaandGuayaguayare.

Figure 6. Nonmetric multidimensional scaling (nMDS) of intertidal macroinvertebrates in the sampling stations of the three beaches at Salybia (s), Guayaguayare(G)andChagville(C),Trinidad.H=highwater,M=midwaterandL =lowwater.

Thedistributionofspeciesoneachbeachindicatedthatatleasttwo juxtaposedzonessharedsimilardominantspecies.However,atleastone or two species were found that distinguished each zone (Table 1). Chagville and Guayaguayare both had Spionid polychaetes dominating the midtidal level and lowwater mark. Polychaete worms were the dominantfaunapresentatthesetwobeaches.However,Salybiashowed a greater diversity across different phyla. The amphipod Parhyale hawaiensiswasthesoledominantspeciesatthelowwatermark.While each zone had at least one dominant species, the driftline in Guayaguayarewasanexception,withnodominantspeciesandveryfew invertebratesinthiszone. All three beaches had an almost equal proportion of polychaete species among the various feeding guilds. Salybia had eight deposit feeders,GuayaguayaressevenandChagvilleten.Filterfeederswerevery

63

uncommonwithonespecieseachatSalybiaandGuayaguayareandtwo at Chagville. Herbivorous polychaetes were absent in Guayaguayare. However,SalybiaandChagvillehadtwoherbivorespecies.Thiswasalso thecaseforthecarnivorousspeciesatthesetwobeaches.Guayaguayare had a single carnivore species present. Some species of polychaetes occupied more than one feeding guild such as the spionid polychaetes, Nereisspp.andLumbrinerisspp.
SamplingStations Zones* Samplingsites Salybia Guayaguayare Chagville Hightide High Notomastussp. Nospecies Unidentifiable (driftline) Intertidal Nematodessp. dominant juvenile Zone polychaetes Midtidal Mid Nematodes Spio1 Nereissp. Intertidal Nereissp. Donax Apoprionospiosp. Zone Sipunculid2 denticulatus Unidentifiable juvenile polychaetes Lowwatermark Low Parhyale Spio1 Spio1 Intertidal hawaiensis Nereissp. Zone Capitellasp. Prionospio Table 1. Zonation of macroinvertebrates in the intertidal zone based on density (ind.m2)*ZonesasmodifiedfromSouzaFilhoet.al.(2003)

Discussion Thisstudyinvestigatedwhetherintertidalzonationpatternswerepresent onthethreebeachesofSalybia,GuayaguayareandChagvilleinTrinidad. The results appear to support this hypothesis with two distinct zones apparent,basedondominantspecies.Inaddition,eachbeachappeared to support a different composition of macrofauna in its intertidal zone, withrespecttothespeciespresentandtheirtrophiclevel. Several authors have proposed that grain size may influence beach slope, with coarsegrained sediment more likely to produce a steeper slopethanfinegrainedsediment(Jaramilloetal.1993,Knox2000,Rodil andLastra2004,NybakkenandBertness2005,WieserUndated).Thiswas foundtobethecaseforSalybiaandGuayaguayare.Salybiahadasteeper slope than Guayaguayare and comprised mostly of coarsegrained sand and cobble. This may in turn affected the length of the intertidal zone. Thus, the steeper the slope of the beach, the higher the proportion of coarsegrainedsandandtheshorterthelengthoftheintertidalzone.

64

TheBIOENVindicatedthatgrainsizehadaninfluenceonthebiotic patternproducedbythenMDS.Apredominantlycoarsegrainedbeachis more porous than finegrained beaches. Therefore, water cannot be retained, leaving intertidal invertebrates susceptible to desiccation. This coarsegrained sand is also unfavourable for burrowing organisms. However,thisdoesnotnegatethepossibleinfluencetheothervariables such as nutrients, salinity, and oxygen content of the sediment, on the speciesassemblages.Forexample, thelargedecreaseinthesalinityand conductivity levels measured at site C5 in Chagville may have occurred duetotheproximityofthetransecttotheCuesaRiver,whichemptiesa substantialvolumeoffreshwaterintothebay.Despitethis,however,the biotafoundatthistransectwasnotaffectedsuggestingtheirtoleranceto thefreshwater.Thiswasnotsurprisingsincesomeofthesespecieshave beenfoundinestuariesthathavebrackishwater. Salybia had the highest proportion of coarsegrained sand and the highestspeciesdiversity.Yetithadthelowestnumberofindividualsm2. Whiletherewasasignificantnumberofspeciesofpolychaetespresent, thenumberofcrustaceanswasmuchhigherthanattheotherbeaches. Theisopodsmaybebettersuitedtomovingthroughthecoarsegrained sand and cobble as well as more tolerant of the low interstitial water withinintheintertidalzone. Guayaguayarehadthehighestproportionoffinegrainedsands,with extremely low species diversity, but a density of individuals that was much higher than Chagville and Salybia. It is possible for some species dominancetooccurinbeaches,becauseofenrichmentandotherfactors. Thismayfavourtheproliferationofthespecies.TheSpionidpolychaete species were notably absent from Salybia, yet highly abundant at the othertwosites.Itisquitepossiblethatthisfamilyofpolychaetesprefer finegrained sediment. Low salinity and conductivity levels may also be desired.Thesameconditionsmaybepreferredbybivalvemolluscssuch as Donax denticulatus, which was found only at Guayaguayare in this study. There was a significant lack of bivalves in Salybia with the exceptionofasingleDiplodontapunctata. The sipunculid worms were rock or coral borers. This diagnostic featuremayreadilyexplaintheirabsencefromGuayaguayareduetothe lackofcobbleorbrokencoral.However,theymighthavebeenexpected in Chagville, since there were quite a number of coral fragments at this site. Higher salinity may be an important factor accounting for their presence in Salybia than Chagville, despite both beaches having a river feeding into the saline waters. Another reason for the absence of

65

sipunculid worms from Chagville, is that this phylum maybe sensitive to wastewaterandothercontaminants. Nereis was also absent from Guayaguayare. This polychaete has parapodiathataremodifiedtoallowthemtonegotiatethesurfaceofthe coarsegrained sand and cobble. The very unstable nature of the fine grainedsedimentsinGuayaguayaremaynotbeidealforthisspecies. ThesimilaritiesinChagvillesenvironmenttotheothertwobeaches mayhaveallowedittohavespeciessharedbybothbeaches.Inshort,the varying conditions of the beaches possibly accounted for the presence and absence of different species. However, other factors that were not testedcouldhavealsoinfluencedthepresenceofvariousspecies. McLachlan and Jaramillo (1993) in reviewing zonation patterns in sandybeachesconcludedthatallzonationstudiesrevealedthreezones supralittoralzone,littoralzoneandthesublittoralzone.However,these zonesarenotspatiallyortemporallyfixedandthefaunaadjustwiththe daily changes in the shore gradient (Knox 2000, Nybakken and Bertness 2005). This movement caused by the waves and swash transporting sedimentandorganismsfromoneareatoanothercausesthepatchiness oftenfoundonsandyshores(Knox2000,NybakkenandBertness2005). None of the organisms suggested by McLachlan and Jaramillo (1993) as diagnostic of the supralittoral zone, such as ocypodid crabs, talitridamphipodsandisopods(BrownandMcLachlan1990,Jaramilloet al. 1993, Knox 2000, Nybakken and Bertness 2005) were found at the threeTrinidadbeaches.Thisindicatesthatthesupralittoralzonewasnot surveyed in this study. Future sampling above the driftline may yield someofthefaunalfamiliessuggestedbyMcLachlanandJaramillo(1995 inKnox2000),NybakkenandBertness(2005). The midlittoral zone contained different polychaete species including the Spionid species, and bivalves at Chagville and Guayaguayare.Ithasbeenpreviouslysuggestedthatthiszonemayhave greaterdiversitythanotherzonesintheintertidalarea(Knox2000).This study indicates that the polychaetes, identified as dominant species at these three study sites, were either deposit feeders or filter feeders. Depositfeeders are organisms that ingest sediment to obtain their nutritionandareabundantinfinesedimentswithahighclayandorganic content(Fauchald1979,Newell1979,Knox2000,NybakkenandBertness 2005).Thefilterfeedersontheotherhandobtaintheirfoodintheform of phytoand zooplankton, detritus and organic particles, suspended in water. These polychaetes often proliferate in coarsegrained sediment (Fauchald1979,Newell1979,Knox2000,NybakkenandBertness2005).

66

The presence of these organisms may suggest the nature of the sedimentatthethreebeaches.Salybianotablylackeddominantfilteror depositfeeders, with the exception of Notomastus sp. in the high tide zone. Chagville and Guayaguayare, however, had some depositfeeders and filterfeeders within the littoral zone. While the predominant grain size can be considered coarsegrained, Guayaguayare and Chagville did have a much higher proportion of finegrained sand than Salybia. This mayhaveallowedforthedominanceofthesetwofeedingguilds.Nereis wasthelonecarnivorethathadsomedominanceinChagvilleandSalybia, favouringtheslightlydrierconditionsofthelittoralzone. One interesting feature in the distinction of feeding guilds was that some organisms occupied more than one feeding guild. Most of the polychaetesthatdisplayfilterfeedingordepositfeedingbehaviourscan switch between these two modes, based on flow rates of the water. These organisms were called interface feeders by Dauer et al. (1981 in Knox 2000). Other species like Nereis spp. which can be considered an omnivore (Fauchald 1979), may be opportunistic in nature and feed on anyorganicparticlesthatmaybeavailableatthetime. Most intertidal zonation studies focus on vertical zonation patterns in which differences between the various zones are reported. However, veryfewstudieshaveaddressedhorizontalspatialdistributionwithinthe sandybeach(GimenezandYannicelli2000).Thedatainthisstudyfailed to detect any substantial difference on the longitudinal axis of the beachesstudied. Sandy beaches are important providers of ecosystem services to peoplebutaresensitivetonaturalandanthropogenicdisturbances.They create important functional links between primary producers and large consumers (McDermott 1983, Ricciardi and Bourget 1999, Jones et al. 2004, Defeo et al. 2009). Filter feeders feed on phytoplankton and grazers such as amphipods feed on stranded algae. Wastes from these organismsreturntotheenvironmentasuseablenutrients.Commercially important fishes feed on surface depositfeeders and filterfeeders (RicciardiandBourget1999,Jonesetal.2004,Defeoetal.2009).Whileit is possible for a beach to recover from natural disturbances over time, anthropogenic degradation can become permanent (Linton and Warner 2003,Schlacheretal.2008,Defeoetal.2009,Pallewatta2010). Intertidal species tend to be sensitive to pollution and changes in their environment (Ulfig 1997, Salvo and Fabiano 2007, Defeo et al. 2009).Therefore,theycanactasexcellentbiologicalindicatorssincethey aremostlysessileorsedentaryinnature,andeasytoidentify(Clarkeand Warwick2001,Fujii2007,Schlacheretal.2008).Diversityestimationsin

67

benthic ecology have become increasingly important in regulatory policiesandconservationdecisions(Carney2007,Schlacheretal.2008). Consequently, studies on intertidal macrofauna can be beneficial to TrinidadandTobagoandthewiderCaribbean,sincetheycanprovidean index to the condition of the beach environment (Linton and Warner 2003)andallowforimprovedcoastalmanagementstrategies.InTrinidad and Tobago, approximately 80% of urbanised land is located within or adjacent to coastal areas and approximately 50% of the roads pass throughcoastalareas(OfficeofCentralStatistics2007). Temporal benthic studies are particularly important for understandingthepossibleconsequencesthatclimatechangemayhave onbeachecology(Saizar1997,Sagarin2003,Jonesetal.2004,Cambers etal.2008,Schlacheretal.2008,Defeoetal.2009).Whilethefullimpact of climate change on sandy beaches remain unknown, some of the predicted changes include decreased physiological performance, geographic movement of fauna to cooler latitudes, increased invasive species,decreaseddiversityanddensities,changingsedimentsupplyand changingparticlesizeandslope(Jonesetal.2004,Cambersetal.2008, Schlacheretal.2008,Defeoetal.2009). Inresponsetosealevelrise,theremaybeanincreaseinhard engineering structures to protect coastlines and human settlements againsttheencroachingsea.Thisconsequentlypreventsthemigrationof the beach and thus reduces available beach habitat. This may in turn affect the assemblage of fauna present on the beach (Sobocinski 2003, DuganandHubbard2006,Dugan2008,Schlacheretal.2008,Defeoetal. 2009). AtGuayaguayare,thehightidelevelwas establishedatthebaseof theseawall.Macrofaunawasnoticeablyabsentatthissite.Observations during this study suggest that coastal squeeze may already be taking effectatthissite.However,furtherresearchisrequiredtodemonstrate thiseffect. Conclusion Differencesintherichnessandabundanceofbenthicmacrofaunainthe intertidalzoneatSalybia,GuayaguayareandChagvilleappearrelatedto differentabioticenvironmentsateachbeach.Amongtheenvironmental variablessampled,pHandgrainsizemayhavethegreatestinfluencein thepatternsfound.Eachbeachhadadifferentverticalzonationpattern, whichwasconsistentwiththeintertidalzoneinthegeneralisedscheme proposedbyJaramilloandMcLachlan(1993).Nohorizontalzonationwas found and it was assumed that the data collected might not have been

68

adequate to show a clear pattern. The information produced from this study can form the baseline to future temporal studies, which can be beneficialtoTrinidadandTobagoandassistinassessingthestateoflocal beaches and developing suitable management strategies to counter damagingeffectsonthecoastlines. Acknowledgements Theprimaryauthorwouldliketogivespecialthankstoherparentswho assisted with much of the field and lab work as well as Kerron Castillo. Thanks also to Mark Charran and Shiva Maharaj for giving their time when possible to assist in processing the benthic samples for identification. Thanks to Anuradha Singh, Professor John Agard and Dr. Judith Gobin for their assistance with the identifications of the benthic organisms, invaluable comments and advice. The assistance of Rajesh Ragooforhissupportwithdataanalysisisgratefullyacknowledged.The assistance of Raabia Hosein, Rajindra Mahabir, Christine Fraser, Anton Manoo, Professor Paul Shaw andMelissa Atwell in theacquisition ofall equipment and materials utilised for this study, is gratefully acknowledged. Special thanks to Lee Ann Beddoe for her help and patiencewithvariousaspectsofthisproject.Finalthankstoallthosewho encouraged, motivated and supported the primary author during the research. References
Barros F, Borzone CA, Rosso S (2001) Macroinfauna of six beaches near Guaratuba Bay, southernBrazil.BrazilianArchivesofBiologyandTechnology44:351364 Bousfield EL, Quesnel VC (19891990) The beach fleas and sandhoppers (Amphipoda; Talitirdae) of Trinidad. Living World Journal of Trinidad and Tobago Field Naturalists' Club:4345 BrownAC,andMcLachlanA(1990)Ecologyofsandyshores,Vol.Elsevier,Amsterdam CambersC,ClaroR,JumanR,ScottS(2008)Climatechangeimpactsoncoastalandmarine biodiversityintheinsularCaribbean.ReportNo.382,CANARI Carney R (2007) Use of Diversity Estimations in the Study of Sedimentary Benthic Communities.In:OceanographyandMarineBiology.CRCPress Clarke KR, Ainsworth M (1993) A method of linking multivariate community structure to environmentalvariables.MarineEcologyProgressSeries92:205219 Clarke KR, Warwick RM (2001) Change in marine communities: an ecological approach to statisticalanalysisandinterpretation,Vol.PRIMERE,Plymouth Defeo O, McLachlan A (2005) Patterns, processes and regulatory mechanisms in sandy beachmacrofauna:amultiscaleanalysis.MarineEcologyProgressSeries295:120 Defeo O, McLachlan A, Schoeman DS, A. ST, Dugan J, Jones A, Lastra M, Scapini F (2009) Threatstosandybeachecosystems:Areview.Estuarine,CoastalandShelfScience81:1 12 Dugan J (2008) Ecological effects of coastal armouring on sandy beaches. Marine Biology 29:160170

69

DuganJE,HubbardDM(2006)Ecologicalresponsestocoastalarmouringonexposedsandy beaches.ShoreandBeach74 FauchaldK(1979)Thedietofworms:astudyofpolychaetefeedingguilds.Oceanographic MarineBiologyAnnualReview17:193284 FujiiT(2007)Spatialpatternsofbenthicmacrofaunainrelationtoenvironmentalvariables in an intertidal habitat in the Humber estuary, UK: Developing a tool for estuarine shorelinemanagement.Estuarine,CoastalandShelfScience75:101119 Gimenez L, Yannicelli B (2000) Logshore patterns of distribution of macroinfauna on a Uruguayan sandy beach: an analysis at different spatial scales and of their potential causes.MarineEcologyProgressSeries199:111125 Gobin JF (2010) Freeliving marine polychaetes (Annelida) inhabiting hardbottom substratesinTrinidadandTobago,WestIndies.RevBiolTrop58:147157 Institute of Marine Affairs (2004) A guide to beaches and bays of Trinidad and Tobago. InstituteofMarineAffairs JaramilloE,McLachlanA(1993)Communityandpopulationresponsesofthemacrofaunato physicalfactorsoverarangeofexposedsandybeachesinsouthcentralChile.Estuarine, CoastalandShelfScience37:615624 Jaramillo E, McLachlan A, Coetzee P (1993) Intertidal zonation patterns of macroinfauna overa rangeofexposedsandybeachesinsouthcentralChile.MarineEcologyProgress Series101:105118 Jones A, Gladstone W, Hacking N (2004) Sandybeach ecosystems and climate change: potentialecologicalconsequencesandmanagementimplications.In:TheSecondDecade Coastal Planning and Management in Australia towards 2014 Proceedings of Coast to Coast2004,Australia's6thNationalCoastalManagementConference,2004April1923, Hobart KnoxG(2000)SoftShores.In:TheEcologyofSeashores.CRCPress LintonDM,WarnerGF(2003)BiologicalindicatorsintheCaribbeancoastalzoneandtheir roleinintegratedcoastalmanagement.OceanandCoastalManagement46:261276 McDermottIJ(1983)FoodwebinthesurfzoneofanexposedsandybeachalongtheMid Atlantic coast of the United States. In: McLachlan A, Erasmus T (eds) Sandy beaches as Ecosystems.JunkPubl.,TheHague,p579538 McLachlanA(1996)Physicalfactorsinbenthicecology:effectsofchangingsandparticlesize onbeachfauna.MarineEcologyProgressSeries131:205217 NevesFM,BemvenutiCE(2006)Spatialdistributionofmacrobenthicfaunaonthreesandy beaches from northern Rio Grande do Sul, Southern Brazil. Brazilian Journal of Oceanography54:135145 NewellRC(1979)BiologyofIntertidalAnimals,Vol.MarineEcologicalSurveysLtd.,Kent Nybakken JW, Bertness MD (2005) Marine Biology: an ecological approach, Vol. Pearson Education,Inc.,SanFrancisco OfficeofCentralStatistics(2007)FirstCompendiumofEnvironmentalStatisticsTrinidadand Tobago.In:DevelopmentMoPa(ed),PortofSpain PallewattaN(2010)ImpactsofClimateChangeonCoastalEcosystemsintheIndianOcean Region. In: Michel D, Pandya A (eds) Coastal Zones and Climate Change. The Henry L. StimsonCenter,Washington,p316 Ricciardi A, Bourget E (1999) Global patterns of macroinvertebrate biomass in marine intertidalcommunities.MarineEcologyProgressSeries185:2135 SagarinRD(2003)AChecklistforHistoricalStudiesofSpecies'ResponsestoClimateChange. In.CRCPRess Saizar A (1997) Assessment of impacts of a potential sealevel rise on the coast of Montevideo,Uruguay.ClimateResearch9:7379

70

SalvoVS,FabianoM(2007)MycologicalassessmentofsedimentsinLigorianbeachesinthe northwestern Mediterranean: pathogens and opportunistic pathogens. Journal of EnvironmentalManagement8:365369 Schlacher TA, Schoeman DS, Dugan J, Lastra M, Jones A, Scapini F, McLachlan A (2008) Sandy beach ecosystems: key features, sampling issues, management challenges and climatechangeimpacts.MarineEcology29:7090 Sobocinski KL (2003) The impact of shoreline armouring on supratidal beach fauna of CentralPugetSound.MScThesis,UniversityofWashington SouzaFilho,P.W.M.;Tozzi,H.A.M.andElRobrini,M.(2003)Geomorphology,landuseand environmentalhazardsinAjuruteuamacrotidalsandybeach,northernBrazil.Journalof CoastalResearch,35,580589. Ulfig K (1997) General assessment of the occurrence of keratinolytic fungi in river and marine beach sediments of Caledonian waters (Spain). Water, Air and Soil Pollution 94:275287

71

GeneticDiversityandStructureoftheNeotropical MonodominantSpeciesMoraexcelsa(Benth.)inFive NaturallyFragmentedPopulations NigelAustin,MichaelOathamandPathmanathanUmaharan


DepartmentofLifeSciences,TheUniversityoftheWestIndies,St.Augustine,Trinidad,West Indies.Email:nigel.austin@sta.uwi.edu

Abstract Mora excelsa is an evergreen Neotropical tree prized for its durable lumber,andthespeciesiswellknownforitsabilitytoformalmostpure monodominantstands.Inspiteofthiseconomicimportanceverylittleis known of its genetics characteristics. In this paper the genetic diversity andstructureoffiveM.excelsapopulations(fourinTrinidadandoneon theSouthAmericanmainland)wasexaminedusingRAPDs.DNAfrom100 leafsamplesofM.excelsawasamplifiedwith9decamerprimerswhich yielded 113 RAPD markers, 78% of which were polymorphic. The populationsexhibitedvaryinglevelsofdiversitywiththeVictoriaMayaro populationinTrinidadhavingthehighestdiversity(%P=64.6%,I=0.376 0.028 and UHe = 0.264 0.020) and the most isolated population in Paria,Trinidadhavingtheleast(%P=45.13%,I=0.2600.028andUHe= 0.1820.020).TheAMOVAandShannonIndexpartitionedthevariation similarly, with greater variation found within populations than among populations, which is typical of Neotropical trees. These results suggest thatthereissignificantdifferentiationbetweenpopulations(Gst=0.2535 and PT = 0.210, P<0.001). However, the PCA, Nm and high similarity betweenpopulationssuggeststhatthisdifferentiationisrecent.Thedata suggests that the two smaller Mora sp. populations in Trinidad were founded,andthetwolargerpopulationswereheavilyconnected,bygene flowinthepast. Keywords percentage polymorphism, Shannons diversity, Trinidad, tropical tree, unbiasedheterozygosity,Venezuela Introduction Tropical forests are regarded as amongst the worlds most biologically diverse and threatened terrestrial ecosystems (Myers, et al. 2000; Achard, et al. 2002; Mayaux, et al. 2005). The greatest threats to these forests are from the increasing demand for forest resources by growing

72

populationsandtheexpansionofurbanandagriculturalareas(Murawski, et al. 1994; Wright 2005). These activities result in reduced forest area and subsequent fragmentation, which notonly affect the functioning of biologically important processes carried out by forests but also the genetic diversity and survival of individual species (Achard, et al. 2002; Krishnapillay, et al. 2003; Schaberg, et al. 2008). The genetic consequences to tropical forests of smaller population sizes and fragmentation, are genetic drift and disruption of gene flow, both of whichmayleadtoinbreedingandlossofalleles(BiscaiadeLacerda,etal. 2008). Additionally, in allogamous species, significant loss of genetic diversity and consequent inbreeding are associated with decreased productivity, fecundity and ability to respond to biotic and abiotic changes. Such changes may be disastrous, especially if the species is exploited as a natural resource (Hamrick 1994; Newton, et al. 1996; Young,etal.1996). Mora spp. (Fabaceae) is a genus of large trees consisting of seven species(Bisbyetal.2010).ThespeciesMoraexcelsaisnativetoGuyana, the proposed centre of diversity, from where it is believed to have expanded to Suriname, Venezuela and Trinidad (Beard 1946). The tree produces valuable and durable timber used for industrial flooring, ship building, railroad and roof construction (Chudnoff 1984). Amerindians havealsobeenknowntouseitsseedsasafoodsource(Beard1946). Thisevergreencanopyspeciescanexceed40minheightandhasa girth of up to 3.6 m above the buttresses. In Suriname, Venezuela and Guyana M. excelsa forms monodominant stands confined to alluvial floodplainsnearstreams(Henkel,2002).InTrinidadM.excelsaseemsto have a wider topographical range as it colonizes hills and ridges (Beard 1946; Rankin 1978; Torti, et al. 1997). At maturity, M. excelsa produces spiked inflorescences each with numerous small white bisexual flowers. Theseflowersareproducedontheterminalendofbranchesevery1.5to 2 years. Flowering is asynchronous in most populations and usually occursbetweenJanuarytoAprilandlaterbetweenJulyandSeptemberof thesameyear(terSteege1990).Vastquantitiesoflargebuoyantseeds eachweighingupto0.5kgand15cminlength,usuallyfallaboutthree monthsafterfloweringusuallyinthewetseason(Beard1946;terSteege 1994). Seeds are water dispersed, but away from streams, they can germinateintheshadeoftheparenttreewithlowmortality(Beard1946; OathamandJodhan2002). InTrinidad,M.excelsaisthemostabundanttreespeciesexistingin naturally fragmented populations (Bell 1971) (Figure 1). The two largest populationsinTrinidad,VictoriaMayaroandMatura,arebelievedtobe

73

established by natural means whereas the other smaller populations (Paria and Cedros) may owe their existence to dispersal by man (Beard 1946). In Trinidad, M. excelsa usually outcompetes and decreases the presenceofothercommontreespeciestoadensitylessthan15%(Beard 1946). Much of the work done on M. excelsa has focused on its ecology, physiology and its management as a timber resource, while no informationexistsonthegeneticdiversityandpopulationstructure.The objectives of this study were to measure the genetic variation present within five populations of M. excelsa (four in Trinidad and one in Venezuela) and to determine how this diversity is structured and estimate the level of gene flow between populations. This information wouldallowforbetterdecisionsonitsconservationandmanagement.
Figure1M.excelsasamplelocationsinTrinidadandSouthAmerica

Methods StudypopulationsandLaboratoryProcedures Fivepopulations,includingfourfromTrinidad(VictoriaMayaro,Matura, CedrosandParia)andonefromVenezuela(Orinocodelta)weresampled (Figure1andTable1).InTrinidad,theshortestdistancebetweenanytwo populationswas20km.AllVenezuelansampleswerecollectednearthe CaoTucupita(TucupitaCanal)intheOrinocoDelta(smallpopulation).


74

Table 1 M. excelsa locations in Trinidad and Venezuela and population descriptions. Population Paria Matura Mayaro Cedros Cao Tucupita Coordinates Latitude 10.78331 10.65917 10.20759 10.11834 Longitude 61.2530 61.0774 61.1528 61.7667 Description Moraforest Moraforest Moraforest Disturbed Mora forest Single dispersed individuals and clumps with few individuals Site Size ha) *490 *10,000 *21,750 *600 **800

9.4427

61.6753

*SizebasedonBeard(1946) **Estimatedfromareasurveyed

WiththeexceptionoftheOrinocoandCedrospopulations,allother sampleswerelocatedinmonodominantMoraforests.Individualsinthe Orinoco delta were found in small patches of few individuals or as isolated individuals, while the Cedros population was highly fragmented withseveralindividualsbeingfoundnearsmallagriculturalplots. Youngtofullyexpandedleafsamplesfrom20Moraspp.individuals per population were collected. Individuals were sampled at least 20 m aparttominimizesamplingofsiblingsandhalfsiblings.Theleafsamples were stored in ice during transport to the laboratory, and DNA isolated using the Kobayashi et al protocol (1998) with the following modifications: approximately 0.4g of leaf tissue was used for each extraction, and all extractions were carried out in 2mL micro centrifuge tubes.Tothecrushedtissue,1mLofBuffer2togetherwith1.5%PVPwas added. In addition, the mercaptoethanol concentration of the buffer was increased to 0.2%. After incubation at 60oC, the sample was centrifugedfor5minutesat12,000rpmandthesupernatantdecanted. The extracted DNA was resuspended overnight in 1 mL 50mM Tris: 10mM EDTA after which it was washed with a phenol: chloroform: isoamyl alcohol (25:24:1) mixture and again with chloroform. The supernatantcontainingtheDNAwasprecipitatedusing1/10volume3M sodium acetate pH 8 and an equal volume of icecold isopropanol. The dried DNA pellet was resuspended in50L 10mM Tris: 1mM EDTA and quantifiedbyspectrophotometry.Workingdilutionswerepreparedin10 mMTrisforfurtheranalysis.

75

Levietals(1993)PCRprotocolwasoptimizedbyadditionofgelatin to the PCR mix. The mix contained 10.6 l millipore water with 0.024% gelatin,5lprimer(3ng/l,Operon,U.S.A.),3lDNA(0.15ng/l),2.5l dNTP(2mM,Gibco,U.S.A.),2.5lBuffer(10x,Klentaq,ABpeptidesInc, U.S.A.), 0.25 l DNA polymerase (Klentaq, 25 U/l, AB peptides Inc, U.S.A.)andwascarriedoutin25lreactions.PCRwascarriedoutona Techne Thermocycler (Techne Co. UK) with amplification conditions as follows:94oCfor5minutes,37oCfor30seconds,72oCfor1minutefor40 cycles and a final extension time of 72oC for 5 minutes. Twentytwo microlitresoftheamplifiedproductweremixedwith3Lloadingdyeand separatedona1.4%agarosegelcontaining0.1ng/mLethidiumbromide in1XTBEbuffer.Thegelwasrunat5V/cmandphotodocumentedusing theUVPImagestore7500GelDocumentationsystem(UVPInc.,U.S.A.). DataAnalysis Reproducible amplicons were scored as present (1) or absent (0) for analysis. Genetic diversity was determined for each population, for all samples collected and for samples collected in Trinidad only. Three indices of diversity were calculated using GenAlEx version 6.3: percentage polymorphism (%P), Shannons diversity index (I), and unbiasedheterozygosity(UHe)(SchlterandHarris2006). The genetic structure within and among the study populations was explored via Analysis of Molecular Variation (AMOVA) (Excoffier, et al. 1992), using the GenAlEx v 6.3 software. Neis (1972) expected heterozygosity in the total population (HT) and the withinpopulation meanexpectedheterozygosity(HS)werealsocalculatedusingPOPGENE v.1.32.Thecoefficientofdifferentiation,GstwascalculatedinPOPGENE and gene flow, Nm estimated. Mantel (1967) tests using 9999 permutations were conducted between pairwise parameters to determinewhetheranyrelationshipexistedbetweenpopulationsizeand genetic diversity, geographic distance and genetic distance and geographicdistanceanddivergence(Table2). Jaccards similarity coefficients were calculated for each population andadendrogramwasconstructedusingtheneighbourjoininginFAMD v 1.25 (Schlter and Harris 2006). The phylogram visualized in FigTree v1.3.1 (Rambaut 2009). Principal coordinate analysis (PCoA) was also undertaken using the similarity and UPGMA algorithms in GenAlex version6.3.

76

Results GeneticDiversity OnehundredandthirteenRAPDreproducibleampliconsweregenerated using the 9 primers for the six M. excelsa populations. The number of amplicons produced per primer ranged from seven in P14 to 25 in BO5 withanaverageof12.5ampliconsperprimer(Table3).
Table 2 Pair wise Neis genetic distance, PT, and Geographic distance for TrinidadandmainlandpopulationsofM.excelsa. Population1 Population2 Unbiased PT Neis genetic distance 0.925 0.250 0.889 0.166 0.872 0.245 0.902 0.230 0.876 0.252 0.903 0.109 0.909 0.209 0.914 0.205 0.881 0.272 0.900 0.172 Geographic Distance(km) 92.594 68.001 75.444 96.325 23.617 50.636 149.849 64.624 155.485 102.390

Cedros Cedros Cedros Matura Matura Matura Matura Paria Paria VictoriaMayaro

Paria VictoriaMayaro CaoTucupita Cedros Paria VictoriaMayaro CaoTucupita VictoriaMayaro CaoTucupita CaoTucupita

Of the 113 RAPD amplicons, 94 (78.76%) were found to be polymorphic among all the populations studied. The percentage polymorphism within populations ranged from 45% in Paria to 65% in VictoriaMayaro (Table 4). When considering all the individuals from Trinidad, there was more variation in the summed samples than any single population in Trinidad (Table 4). The trends observed for all diversityindicesamongpopulationsweresimilar.Ingeneral,thesmaller populations in Trinidad (Paria and Cedros) had lower diversity than the largerpopulations(MaturaandMayaro)withrespecttoallthediversity indices(Table4).Althoughrelativelysmall,theM.excelsapopulationin CaoTucupitaexhibiteddiversitylevelssimilartothatofthemuchlarger Maturapopulation. ThegeneticdistanceamongM.excelsapopulationsestimatedbythe NeisUnbiasedGeneticIdentityindexvariedfrom0.872to0.925.There was no significant (P > 0.05) correlation between pairwise geographic distance and PhiPT values for all populations (R2 = 0.0911) likewise between Neis unbiased genetic distance and geographic distance (R2 = 0.0005)basedtheManteltest(Table2).

77


Table 3. Oligonucleotide sequence and degree of polymorphism detected by primersinTrinidadandVenezuelanpopulations. PrimerName OPA08 OPB05 OPB07 OPC16 OPF09 OPF12 OPP09 OPP14 OPG06 Nucleotide Sequence 5GTGACGTAGG 5TGCGCCCTTC 5GGTGACGCAG 5CACACTCCAG 5CCAAGCTTCC 5ACGGTACCAG 5GTGGTCCGCA 5CCAGCCGAAC 5GTGCCTAACC No.Amplicons No.Polymorphic Scored Amplicons 8 25 14 9 13 8 11 7 18 8 19 13 7 3 7 11 7 13


Table4GeneticdiversitywithinpopulationsofM.excelsafromTrinidadandthe Venezuelan mainland. N = number of individuals sampled, #P = number of polymorphic markers, %P = percentage of polymorphism, I = Shannon Diversity IndexandUHe=UnbiasedHeterozygosity.SE=StandardError. %P I(SE) UHe(SE) Population N #P Paria 20 51 45.13 0.260(0.028) 0.182(0.020) Salybea 20 59 52.21 0.304(0.029) 0.214(0.021) VictoriaMayaro 20 73 64.60 0.376(0.028) 0.264(0.020) Cedros OrinocoDelta Trinidad All 20 20 80 100 52 57 86 94 46.02 50.44 76.11 78.76 0.264(0.029) 0.185(0.021) 0.293(0.029) 0.207(0.021) 0.444(0.026) 0.306(0.018) 0.453(0.025) 0.311(0.018)

The partitioning of the genetic variation by AMOVA suggested significant(P<0.001;9999permutations)geneticdivergenceamongthe populations (Table 5), although most of the variation was within populations.TheresultsforShannonDiversityIndicesweresimilar(Table 6).

78

Table 5 Molecular variance (AMOVA) of six Mora excelsa populations from VenezuelaandTrinidad. Source df SS 272.700 MS Est.Var. % 21.0 Stat Value Prob PT 0.210 0.0001

AmongPops 4 WithinPops 95 Total 99

68.175 2.870

1024.250 10.782 10.782 79.0 1296.950 13.6551 100

ThePhiPTvalue(0.210)obtainedfromAMOVA,whichrepresentsthe average divergence among populations, was similar to the Gst value obtained from partitioning based Neis gene diversity (0.2535). All pair wise PhiPT comparisons derived from AMOVA were significant at the P<0.001level,emphasizingahighdegreeofdifferentiationamongthese populations.
Table6StructuringofvariationofpopulationsofMoraexcelsainTrinidadand VenezuelameasuredusingtheShannonDiversityandNeisDiversityIndices. Neis(1987)GeneDiversity Shannons All Trinidad Allpopulations(SE) Trinidad(SE) Index populations Hpop 0.299 0.301 Hs 0.2050(0.0250) 0.2059(0.0259) Hsp 0.453 0.444 HT 0.2747(0.0372) 0.2718(0.0376) Hpop/Hsp 0.660 0.678 Gst 0.2535 0.2425 (HspHpop)/Hsp 0.340 0.322 Nm 1.4723 1.5618

The PCoA showed that the individuals in the VictoriaMayaro population could not be separated from the Matura population, while there was significant spatial separation evident with the other populations(Figure2).

79

Matura Axis2 Cedros Paria VictoriaMayaro CaoTucupita Axis1

Figure2PCoAforfivepopulationsofM.excelsastudiedusingRAPD.

In this regard, the Cedros population was proximal to the Victoria Mayaro and Cao Tucupita populations on the PCoA plot, while the Cedros and Paria populations appeared as a continuum. The phylogram showed two bifurcating groups (VictoriaMayaro and Matura and Cao TucupitaandParia)andbetweenthemwastheCedrospopulation(Figure 3). The tree also suggests a high degree of similarity between the VictoriaMayaroandMaturapopulationandbetweentheCaoTucupita andPariapopulation.
Figure3UnrootedphylogenetictreeofpairwiseJaccardssimilarityindexvalues andneighbourjoiningamongM.excelsapopulations.

80

Discussion The M. excelsa populations in this study represent the northern most edge of the natural range of the species (Beard, 1946). Typically, island populations are generally thought to exhibit low genetic diversity since colonization events are believed to be followed by loss of variation by founder and bottleneck effects (Barrett and Shore 1989; Husband and Barrett 1991). Remarkably, for a population on the periphery of the speciesrangeandonanisland,thediversityofM.excelsainTrinidadwas foundtobehigh(%P=76%).Asimilarlyhighgeneticdiversity(%P=74%) has also been reported in another tropical forest tree species, Pterocarpusofficianalis,inTrinidad.ThediversityforM.excelsa(Shannon diversity index, Hsp = 0.453) in this study is similar to that reported for other Neotropical tree species including Cedrela odorata (Hsp = 0.450), Swieteniamacrophylla(Hsp=0.450),Plathymeniareticulata(Hsp=0.396), Terminalia amazonia (Hsp = 0.380), Mabea fistulifera, (Hsp = 0.426) and Eremanthuserythropappus(Hsp=0.455)(Gillies,etal.1997;Gillies,etal. 1999;Lacerda,etal.2001;Pither,etal.2003;Goulart,etal.2005;Freitas, etal.2008).ItcanbeconcludedthatthediversityexhibitedbyM.excelsa ishighandistypicalforlonglived,outcrossing,Neotropicaltreespecies (Dick,etal.2003;Lemes,etal.2003;Ruschel,etal.2007;Cardosoetal. 1998;Santosetal.2008). ThehighdiversityofM.excelsapresentinTrinidadsuggeststhatthis species must have colonized the island with a relatively large founder population(Beard1946).UnlikeotherislandsintheCaribbean,Trinidadis a continental island, believed to have been part of the South American mainlandasrecentlyas10,000yearsago(Kenny2000).Thefactthatthe seeds of this species show high mortality when exposed to sea water makes it unlikely that colonization occurred following the separation of TrinidadfromtheSouthAmericanmainland(Beard1946). ThelackofcompetitionfromotherrainforestspeciesandTrinidads seasonal climate seems to have provided ecological release for Mora speciesinTrinidad(Beard1946),accountingforthelargepopulationsize (e.g. VictoriaMayaro) and diversity. Further evidence for ecological release of Mora in Trinidad comes from the following observations: firstly, Mora forests have extended beyond their ancestral flat alluvial habitats and exist as large fasciations in Trinidad; secondly, M. excelsa densities in Trinidad are more than twice that found in Guyana and thirdly, M. excelsa is invading and outcompeting the species in the adjacentCarapaEschweileraforest(Marshall1939;Beard1946;Oatham andJodhan2002).

81

Typically,infragmentedpopulationstheamountofgeneticdiversity found within a population is proportional to its size (Travis, et al. 1996; White, et al. 1999; Lowe et al. 2003). This was observed in Trinidad, where the larger populations (VictoriaMayaro and Matura) were more diverse than the smaller populations (Paria and Cedros). The regression analysis of population size and genetic diversity also supported this hypothesis.Smallisolatedpopulationstendtobelessdiversesincethey are prone to effects of inbreeding and genetic drift (Barrett and Kohn 1991; Ellstrand and Elam 1993). The high degree of monomorphism (> 50%) exhibited within the Paria and Cedros populations may represent founder effects, followed by genetic drift and inbreeding in these populations. IftheM.excelsapopulationsstudiedwerepanmictic,nodivergence among populations would be expected. The significant population divergence among geographically isolated populations in this study therefore suggests restricted gene flow among populations. The lack of waterways between populations makes dispersal by seed unlikely. Furthermore, since the maximal distance travelled by pollinators of Neotropical trees is considered to be not more than 19km (Ward et al. 2005),pollenmediatedgeneflowbetweenM.excelsapopulationsisalso unlikely as the populations were at least 23km apart. Hence, it appears that gene flow between populations is restricted and presently the populationsstudiedmaybegeneticallyisolated. The results from the AMOVA showed that although there was significantdivergenceamongthepopulations,mostofthevariationwas duetodiversitywithinpopulationsratherthanamongpopulations.Thisis consistentwiththegeneralruleforwoodytropicallonglivedspecies,in whichinsectpollinationandallogamyarecommon(Hamrick,etal.1992; Nybom 2004). The gene flow between these populations appears moderate(Nm=1.4723),whichistypicalfortropicaltreesandissimilarto othervaluesreportedforoutcrossinganimalpollinatedspecies(Hamrick andGodt1990). The significant PhiPT and Gst values also suggest that the Mora populations were highly differentiated. Differentiation results from restricted gene flow between populations and can be quite strong in populations that have been fragmented or founded (Loveless and Hamrick1984;Austerlitzetal.1997).Thelackofcorrelationobservedin M. excelsa populations with respect to divergence and diversity and geographic distance suggests that historical and anthropogenic factors may have contributed to their current genetic structure (Castillo Cardenas,etal.2005).

82

Phylogenetic analysis showed limited genetic divergence between the VictoriaMayaro and the Matura populations. The scatter plot from the PCoA showed that the two populations formed a continuum with some degree of overlap. These observations suggest that these populations may historically have been a single population, which divergedfollowingrecentfragmentation.Basedonthesizeandgeological evidence,Beard(1946)believedthattheVictoriaMayaropopulationwas the first established population in Trinidad, which founded the Matura populationbyrangeexpansion. DespitebeinggeographicallyisolatedbytheGulfofParia,theCao Tucupita population and the VictoriaMayaroMatura complex (VMM) were not considerably diverged based on the unbiased Neis genetic distance and PhiPT values. The data suggests that the differentiation observedbetweentheCaoTucupitapopulationandtheVMMcomplex is somewhat recent. A logical explanation for this phenomenon may be that both populations may have been founded by the northern range expansionofMorapriortotheseparationofTrinidadfromthemainland. The existence of some unique polymorphisms in each population suggestssomedifferentiationmayhaveoccurredsincetheseparationof Trinidadfromthemainland. NeisunbiasedgeneticdistancesuggeststhattheCedrospopulation is most similar to the VictoriaMayaro and Cao Tucupita populations. The PCoA further suggests that the VM and Cao Tucupita populations contributed to the Cedros population. This is also supported by the phylogram, which positions Cedros between the two major clades. One explanationforthisobservationmaybethatbothpopulationshavehad geneticinputintotheformationoftheCedrospopulation.Beard(1946) believed that the Cedros population originated from the Orinoco Delta eitherbyalandbridgeorthroughseedsbeingwashedashoreduringthe seasonal floods. Evidence for his theory is supported by the fact that threeotherspeciesfoundintheOrinocoDelta(Saccoglottisamazonica, Quassia amara and Astrocaryum aculeaum) are confined on Trinidad to the Cedros peninsula where M. excelsa also occurs (Beard, 1946). The present data, however, suggests that after separation of the Cao TucupitaandCedrospopulations,theCedrospopulationwasstillableto access genetic input from the VictoriaMayaro population. This could havebeenbecauseofahistoricalrangeexpansionoftheVMpopulation. The M. excelsa population in Paria was the most isolated of the populations studied. This population is located primarilyalong the Paria river which is nested in a valley on the northern side of the Northern Range separated from the nearest natural population by at least 22 km

83

and a mountain ridge about 600m high. The unbiased Neis genetic similarityvaluesandphylogeneticanddivergencedatasuggestthatthis population is most similar to the Cedros population strongly suggesting that this population was founded by individuals from the Cedros population.Ifthispopulationwastheproductofnaturalfragmentationit would have been expected to share more similarity with its nearest neighbouring population Matura. This evidence supports the theory thatthispopulationwasfoundedbyanthropogenicmeans(Beard,1946). The small size of the founding population may explain the low genetic diversityandthedivergencefromtheCedrospopulation. Insummary,thisstudysuggeststhattheVictoriaMayaropopulation is the most diverse and the earliest founding of the five M. excelsa populationsstudied.ThedatasuggeststhatVictoriaMayaroandMatura populationsmayhavebeenintimatelyconnectedbygeneflowuntilthe recent past, supporting the theory of a VMM complex. It appears as thoughtheCedrospopulationwasfoundedfromtheCaoTucupitawith relictualgeneflowfromtheVMMcomplex.Thestudyindicatesthatthe VMM complex is of immense conservation value due to its genetic diversity in Trinidad. The Cedros population with some unique genetic inputfromCaoTucupitawouldalsobeofvalue.TheCedrospopulation however appears to be most vulnerable because of its low genetic variationandthehighlevelofanthropogenicdisturbanceinthearea.This populationshouldbeconsideredasahighpriorityareaforconservation inadditiontotheVMMinthefuture. References
Achard, F., H.D. Eva, H.J. Stibig, P. Mayaux, J. Gallego, T. Richards, and J.P. Malingreau. 2002. Determination of Deforestation Rates of the Worlds Humid Tropical Forests. Science297:9991002. Austerlitz, F., B. JungMuller, B. Godelle, and P.H. Gouyon. 1997. Evolution of coalescence times,geneticdiversityandstructureduringcolonization.TheoreticalPopulationBiology 51(2):148164. Barrett, S.C.H., and J.R. Kohn. 1991. Genetic and evolutionary consequences of small populationsizeinplants:Implicationsforconservation.InGeneticsandconservationof rareplants,editedbyD.FalkandK.E.Holsonger.Oxford,UK:OxfordUniversityPress. Barrett, S.C.H., and J.S. Shore. 1989. Isozyme variation in colonizing plants. In Isozymes in PlantBiology,editedbyD.E.Soltis.Portland,Oregon:DioscoridesPress. Beard,J.S.1946.ThemoraforestsofTrinidad,BritishWestIndies.TheJournalofEcology33 (2):173192. Bell, T.I.W. 1971. Management of the Trinidad mora forests with special reference to the MaturaForestReserve.PortofSpain,Trinidad:GovernmentPrintery. Bisby, F.A., Y.R. Roskov, T.M. Orrell, D. Nicolson, L.E. Paglinawan, N. Nailly, P.M. Kirk, T. Bourgoin,andG.Baillargeon.2010.Species2000andITISCatalogueofLife:2010Annual Checklist.

84

BiscaiadeLacerda,A.E.B.,M.Kanashiro,andA.M.Sebbenn.2008.Effectsofreducedimpact logging on genetic diversity and spatial genetic structure of a Hymenaea courbaril population in the Brazilian Amazon forest. Forest Ecology and Management 255 (3 4):10341043. Cardoso,M.A.,J.Provan,W.Powell,P.C.G.Ferreira,andD.E.DeOliveira.1998.Highgenetic differentiation among remnant populations of the endangered Caesalpinia echinata Lam.(LeguminosaeCaesalpinioideae).MolecularEcology7(5):601608. CastilloCardenas, M.F., N. ToroPerea, and H. CardenasHenao. 2005. Population genetic structure of neotropical mangrove species on the Colombian Pacific Coast: Pelliciera rhizophorae(Pellicieraceae).Biotropica37(2):266273. Chudnoff,M.1984.Tropicaltimbersoftheworld.Remagen,Germany:KesselHouse. Dick, C.W., G. Etchelecu, and F. Austerlitz. 2003. Pollen dispersal of tropical trees (Dinizia excelsa: Fabaceae) by native insects and African honeybees in pristine and fragmented Amazonianrainforest.MolecularEcology12(3):753764. Ellstrand,N.C.,andD.R.Elam.1993.PopulationGeneticConsequencesofSmallPopulation Size:ImplicationsforPlantConservation.AnnualReviewofEcologyandSystematics:217 242. Excoffier, L., P.E. Smouse, and J.M. Quattro. 1992. Analysis of molecular variance inferred frommetricdistancesamongDNAhaplotypes:applicationtohumanmitochondrialDNA restrictiondata.Genetics131:479491. Freitas, V.L.O., J.P. LemosFilho, and M.B. Lovato. 2008. Contrasting genetic diversity and differentiation of populations of two successional stages in a neotropical pioneer tree (Eremanthus erythropappus, Asteraceae). Genetics and Molecular Research 7 (2):388 398. Gillies,A.C.M.,J.P.Cornelius,A.C.Newton,C.Navarro,M.Hernandez,andJ.Wilson.1997. Genetic variation in Costa Rican populations of the tropical timber species Cedrela odorataL.,assessedusingRAPDs.MolecularEcology6(12):11331145. Gillies, A.C.M., C. Navarro, A.J. Lowe, A.C. Newton, M. Hernandez, J. Wilson, and J.P. Cornelius.1999.GeneticdiversityinMesoamericanpopulationsofmahogany(Swietenia macrophylla),assessedusingRAPDs.Heredity83:722732. Goulart, M.F., S.P. Ribeiro, and M.B. Lovato. 2005. Genetic, morphological and spatial characterization of two populations of Mabea fistulifera Mart.(Euphorbiaceae), in different successional stages. Brazilian Archives of Biology and Technology 48 (2):275 284. Hamrick, J.L. 1994. Genetic diversity and conservation in tropical forests. Paper read at Internationalsymposiumongeneticconservationandproductionoftropicalforestseed, atChiangMai,Thialand. Hamrick, J.L., M.J. Godt, and S.L. ShermanBroyles. 1992. Factors influencing levels of geneticdiversityinwoodyplantspecies.NewForests6(14):95124. Hamrick, JL, and MJ Godt. 1990. Allozyme diversity in plant species. In Plant Population Genetics,Breeding,andGeneticResources, editedbyA.H.D.Brown, M.T.Clegg,A.L. KahlerandB.S.Weir.Massachusetts,USA:Sinauer,Sunderland. Husband, B.C., and S.C.H. Barrett. 1991. Colonization history and population genetic structureofEichhorniapaniculatainJamaica.Heredity66:287296. Kenny, J.S. 2000. Views from the Ridge: Exploring the natural history of Trinidad and Tobago.PortofSpain,TrinidadandTobago:ProspectPress. Krishnapillay,B.,M.I.Adenan,A.R.M.Ali,andS.Nimura.2003.Tropicalrainforest:Acradle forbiologicalresourcesandtheMalaysianpoliciesonCBD.Actinomycetologica17(2):50 53.

85

Lacerda, D.R., M.D.P. Acedo, J.P.L. Filho, and M.B. Lovato. 2001. Genetic diversity and structureofnaturalpopulationsofPlathymeniareticulata(Mimosoideae),atropicaltree fromtheBrazilianCerrado.MolecularEcology10(5):11431152. Lemes,M.R.,R.Gribel,J.Proctor,andD.Grattapaglia.2003.Populationgeneticstructureof mahogany (Swietenia macrophylla King, Meliaceae) across the Brazilian Amazon, based on variation at microsatellite loci: Implications for conservation. Molecular Ecology 12 (11):28752883. Loveless,M.D.,andJ.L.Hamrick.1984.Ecologicaldeterminantsofgeneticstructureinplant populations.AnnualReviewofEcologyandSystematics15(1):6595. Lowe,A.J.,B.Jourde,P.Breyne,N.Colpaert,C.Navarro,J.Wilson,andS.Cavers.2003.Fine scale genetic structure and gene flow within Costa Rican populations of mahogany (Swieteniamacrophylla).Heredity95:268275. Marshall, R.C. 1939. Silviculture of the trees of Trinidad and Tobago British West Indies. Oxford,UK:OxfordUniversityPress. Mayaux,P.,P.Holmgren,F.Achard,H.Eva,H.J.Stibig,andA.Branthomme.2005.Tropical forest cover change in the 1990s and options for future monitoring. Philosophical TransactionsoftheRoyalSocietyB:BiologicalSciences360(1454):373384. Murawski, D.A., I.N. Gunatilleke, and K.S. Bawa. 1994. The effects of selective logging on inbreeding in Shorea megistophylla (Dipterocarpaceae) from Sri Lanka. Conservation Biology8(4):9971002. Myers, N.J., R.A. Mittermeier, C.G. Mittermeier, G.A.B. da Fonseca, and J. Kent. 2000. Biodiversityhotspotsforconservationpriorities.Nature403(6772):853858. Newton, A.C., J.P. Cornelius, P. Baker, A.C.M. Gillies, M. Hernandez, J.F. Mesen, and A.D. Watt. 1996. Mohagany as a genetic resource. Botanical Journal of the Linnean Society 122(1):6173. Nybom,H.2004.ComparisonofdifferentnuclearDNAmarkersforestimatingintraspecific geneticdiversityinplants.MolecularEcology13(5):11431155. Oatham,M.P.,andD.Jodhan.2002.Ismoratakingover?Testingthelimitstotheinvasive ability of Mora excelsa Benth. A pilot study. Journal of the Trinidad and Tobago Field Naturalists'Club. Pither,R.,J.S.Shore,andM.Kellman.2003.GeneticdiversityofthetropicaltreeTerminalia amazonia(Combretaceae)innaturallyfragmentedpopulations.Heredity91:307313. Rambaut, A. 2009. FigTree: Tree figure drawing tool, v1.2.2. University of Edinburgh, UK: InstituteofEvolutionaryBiology. Rankin,J.M.K.1978.Theinfluenceofseedpredationandplantcompetitionontreespecies abundancesintwoadjacenttropicalrainforestcommunitiesinTrinidad,W.I.,University ofMichigan,AnnArbor,MI. Ruschel,A.R.,R.O.Nodari,andB.M.Moerschbacher.2007.ThegeneticstructureofSorocea bonplandii in Southern Brazilian forest fragments: AFLP diversity. Silvae Genetica 56 (2):5157. Santos, RP, PCS ngelo, PTB Sampaio, RC Quisen, MC Leite, and CL Oliveira. 2008. Geographic pattern of genetic diversity in natural populations of Rosewood (Aniba rosaeodora),intheCentralAmazonia.ActaAmazonica38:459466. Schaberg, P.G., D.H. DeHayes, G.J. Hawley, and S.E. Nijensohn. 2008. Anthropogenic alterationsofgeneticdiversitywithintreepopulations:Implicationsforforestecosystem resilience.ForestEcologyandManagement256(5):855862. Schlter, P.M., and S.A. Harris. 2006. Analysis of multilocus fingerprinting data sets containingmissingdata.MolecularEcology6(2):569572. ter Steege, H. 1990. A monograph of wallaba, mora and greenheart, Tropenbios Series. Wageningen,TheNetherlands:TropenbosFoundation.

86

. 1994. Flooding and droughttolerance in seeds and seedlings oftwo Mora species segregated along a soil hydrological gradient in the tropical rain forest of Guyana. Oecologia100(4):356367. Torti,S.D.,P.D.Coley,andD.P.Janos.1997.Vesiculararbuscularmycorrhizaeintwotropical monodominanttrees.JournalofTropicalEcology13(4):623629. Travis,S.E.,J.Maschinski,andP.Keim.1996.AnanalysisofgeneticvariationinAstragalus cremnophylax var. cremnophylax, a critically endangered plant, using AFLP markers. MolecularEcology5(6):735745. Ward, M., C.W. Dick, R. Gribel, and A.J. Lowe. 2005. To self, or not to self...A review of outcrossingandpollenmediatedgeneflowinneotropicaltrees.Heredity95:246254. White, G.M., D.H. Boshier, and W. Powell. 1999. Genetic variation within a fragmented populationofSwieteniahumilisZucc.MolecularEcology8(11):18991909. Wright,S.Joseph.2005.Tropicalforestsinachangingenvironment.TrendsinEcologyand Evolution20(10):553560. Young, A., T. Boyle, and T. Brown. 1996. The population genetic consequences of habitat fragmentationforplants.TrendsinEcologyandEvolution11(10):413418

87

TheconservationstatusofMetastelmafreemani GayatrilakshmiRaghavaSingh1andMichaelOatham
DepartmentofLifeSciences,TheUniversityoftheWestIndies,St.Augustine,Trinidad,West Indies.Email:gamma_rae_star0020@hotmail.com 1 CorrespondingAuthor

Abstract Metastelma freemani (ApocynaceaeAsclepiadoideae) is an endemic perennialfoundonthenortheastcoastofTrinidad.Currentlythespecies does not occur in any protected area and coastal development may threaten its habitat, endangering its existence. To enable greater protection of M. freemani, its conservation status and current distributionneedstobebetterunderstood.Fortyrandomsamplepoints were chosen along the northeast coast, and at each point a hundred metertransectwassampled.Hereabiotic(substratetype,substratepH, inclination,elevation,distancefromshore,aspect,canopycover,canopy openness and land use) and biotic (dominant vegetation) parameters wererecorded.WhereM.freemaniwasidentifiedalongthetransect,the extent of its inland distribution was measured via subsequent perpendiculartransectsfromthecoast.Usingpresencedata,Maxentwas usedtomodelM.freemanisgeographicdistributionalongthenortheast coastanddeterminemajorpredictorsofthispattern.Logisticregression was used to facilitate intrasite comparison and development of predictive formulae for observed distributions. Nonmetric Multidimensional Scaling (nMDS) was used to investigate correlations betweenbiologicalcommunitiesandM.freemanisites.Itwasfoundthat northeastern Trinidad was a hotspot for M. freemani, with geology emergingasamajorpredictorofthespeciespresence.However,onthe microscale,noneoftheparametersanalysedviaLogisticRegressionwas significant.Additionally,nMDSrevealednovegetativecompositionsthat predicted M. freemani distribution. M. freemani appears to be a poor competitorandthusinhabitsareasdevoidofothervegetation.Thisplant appears adapted to the harsh abiotic environments found on rocky coastal areas. Unfortunately, its dispersal success seems limited since it has not been observed at sites other than those where it was first collected. Keywords Asclepiadoideae,distribution,endemic,habitat,Maxent,Trinidad

88

Introduction Metastelma freemani is a Trinidadian endemic belonging to the Apocynaceae (dogbane) family and the Asclepiadoideae (milkweed) subfamily. Williams et al. (19281992), describes the plant as prostrate, highlybranched,formingmatsontherockycliffsbetweenBalandraand Toco,(Figure1).Itsleavesareopposite,baseofstemwoody,andcorolla tubeandlobesbothlongandpaleyellow. According to APGII (2003), M. freemani initially belonged to the Asclepiadaceae family. However, recent taxonomic consolidation has demoted this family to three subfamilies (Periplocoideae, Secamonoideae, Asclepiadoideae) and these were added to the family Apocynaceae. This plant was also originally included within the genus CynanchyumbuttheNewWorldspecieshavenowbeenseparatedintoa newgenus,Metastelma(LiedeandMeve,2004). The generalized destruction and degradation of natural habitats in theWestIndies,bothduringthecolonialperiodandinrecenttimes,has ledtothelossofseveralplantspecies.However,thetrueextentofthis lossremainsunknown(AcevedoRodriguezandStrong,2007).Currently, thetotalnumberofnativeandnaturalisedplantsofTrinidadandTobago is reported to be 2361 species (EMA, 2002). A recently revised list of endemicplantsforTrinidadandTobagosuggeststhatbothislescontain approximately 59 endemics, 39 of which are exclusive to Trinidad (Van den Eynden et al. 2008). Being a continental island, Trinidad has a low rateofplantendemismthatstandsslightlylessthan3percent(Vanden Eynden et al., 2008). However, it has high species richness, sharing the majority of its flora and fauna with the South American mainland (CBD, 2010).Inspiteofthis,theconservationstatusofTrinidad'sfloraispoorly known, as is the case with other developing nations. Indeed, Van den Eynden et al. (2008) note that Trinidad only has seven plant species on the2007IUCNRedListofThreatenedSpecies,noneofwhichincludeits endemicspecies. Endemics are important contributors to both local and global biodiversity and, given their restricted global range, are usually of high conservation interest. While a local legislative framework exists in Trinidad for protectionto threatened species (Environmentally Sensitive SpeciesRules,2001),noplanthasyetbeenaddedtothislist. Thepurposeofthisstudywastodeterminethedistribution,ecology and conservation status of M. freemani, and so determine the most efficientmeansofprotectionforthisplant.

89

Methods Studyarea The study was conducted in the northeastern region of Trinidad, from MatelottoMatura,withsamplingconfinedtothecoastalzone(Figure1). Within this 60km long region, 40km of transects were surveyed in addition to several random, exploratory surveys. BHP Billiton (2002) notesthevarietyofcoastallandformsexistingwithinthisareaincluding exposed rocky shores, wave cut platforms, exposed tidal flats, coarse sandbeaches,gravel/pebblebeachesandcobble/gravelbeaches.

Figure1MapofstudyareaofM.freemanonthenortheastcoastofTrinidad.

DataCollection Accordingtoherbariumrecords,M.freemaniisfoundonelevated,rocky formations along the coastline. An initial reconnaissance trip was conducted to the location where herbarium voucher specimens were previously collected, to confirm that the species was still extant at this site. Thus, sample sites were chosen from both rocky and nonrocky locations to investigate the hypothesis, that these plants prefer rocky substrate. A BHP Billiton coastal classification system (BHP 2002) was used to create a thematic layer in the ArcMap 9.3 (ESRI 2005) GIS environment. The study area was classified as ESI1 (exposed rocky shores), ESI2 (wave cut platform), ESI4 (coarse sand beaches), ESI5 (exposedtidalflats)andESI6(gravel/pebbleandcobble/gravelbeaches), using this coastal classification system. Herbarium records indicated preference of rocky substrate. To investigate this ESI1 and 2 were

90

combinedunderrockysubstratewhilstESI4,5and6undernonrocky,in selection of sample sites. Soil type (alluvial, beach, terrace and upland) and geology (alluvium, limestone, metabasalt, metasandstone and slate/metalimestone) were also used as other measures of substrate preference. Forty random sample points were generated using this GIS, which included twenty on rocky substrate and twenty on nonrocky substrates. At each sample point, a 100m line transect was laid out parallel to the coast, and sampling took place at five 25m intervals along each transect.Atallsitesbiotic(vegetationcomposition)andabioticvariables including substrate type, substrate pH, inclination, elevation, distance from shore, aspect, canopy cover, canopy openness and land use) were recordedateachinterval. Where M. freemani was present, transects were sampled perpendiculartothecoastfromtheindividualmats,to20mbeyondthe lastM.freemaniindividual.Microenvironmentalvariableswererecorded at 20m intervals where M. freemani was detected. Stretches of coast were also surveyed if they were thought to be potential sites for M. freemani.Inthisinstance,datawasonlyrecordedforpositivesites. DataAnalysis Maxent (Phillips et al., 2004) was used to model the geographic distribution of M. freemani. This habitat mapping technique predicts a species distribution by estimating the maximum uniform entropy function,assumingthattheexpectedvalueofeachfeaturematchesthe measured average for those features (Phillips et al., 2004). In this case, occurrence localities and environmental data provide the measured values for the model (Phillips et al. 2006; Phillips et al. 2004). In this study,presencedataandGISlayerswereusedtogeneratethedataused in Maxent. A jackknife analysis was also performed to ascertain which environmental factors had the greatest influence on M. freemani distribution. Logistic Regression was used to determine if any of the abiotic variables were correlated to the presence of M. freemani. Logistic regression allows for the creation of fitted models, which allow mathematical estimation of presence or absence (Berwick et al., 2005). Anintrasitecomparisonwasconductedusingpresence/absencedatafor eachverticalintervalalongthehorizontaltransect. Descriptive statistics were used to examine the data for normality. Correlations between independent variables were assessed via Spearmans Rank Correlation. Significant correlation between pairs of

91

variables was defined when P values were less than 0.05 and R2 values greater than 0.7. One member of correlated pair was then removed to prevent over/under weighting in subsequent analyses, fourth root transformations were carried out to normalise the dataset, and logistic regressionperformed. Nonmetric multidimensional scaling (MDS) was performed on the random sample points. The floristic datasets were semiquantitative because presence/absence data was recorded at each transect interval. MDSgroupswerecreated,whichincludedApotentialM.freemanisites (thatis,siteswithinthepredictedhotspotregionofMaxentsmodel),B actual M. freemani sites and C all other sites. Bubble plots of co occurring species were overlaid onto the MDS figures, to allow explorationoftrendswithineachgroup(A,BandC). Results Of the forty randomly selected survey sites, M. freemani was only detected at one location, where five individual mats of the plant were present. The result of the Maxent model of geographic distribution of MetastelmafreemaniispresentedatFigure3.Hereavalueof1suggests aprobabilityofM.freemanipresence,whileavalueof0indicatesthatits presence is highly unlikely. This model predicts that the northeastern cornerofTrinidadisapotentialhotspotforM.freemanioccurrence.

Figure2ShowingMaxentPredictedModelofGeographicDistributionof Metastelmafreeman.

92

Jackknifemodelling,ofintrasiteconditionswhereM.freemanioccurred, suggested that the environmental variable with the greatest predictive power(Phillips,2005)forthisspecieswasgeology(Table1). Table1ShowingJackknifeAnalysisofVariableContributions
Variable Geology BHPCoastal Classification SoilType LandUse Slope Exposure Percent Contribution 80.6 12.3 3.7 3 0.5 0 Permutation Importance 58.6 2 31.4 7.7 0.3 0

The analysis of descriptive statistics of the abiotic and biotic factors associated with M. freemani occurrence, indicated that all of the variableswereskewed,kurtosedorboth.ASpearmansrankcorrelation wasperformedonthesedataandthissuggestedthatRandiaaculetaand Jaquina armillaris (r21.0, p= 0.0); substrate type and substrate pH (r2 0.9903, p=0.0); location and elevation (r20.8434, p=0.0); distance from shoreandlocation(r20.8128,p=0.00)weresignificantlycorrelatedpairs. Randiaaculeta,substratetypeandlocationwerethusremovedfromthe dataset and fourth root transformation performed on the remaining variables. All of the p values were greater than 0.05, thus it was concluded that none of the independent variables were significantly correlatedwiththepresenceofM.freemani. Itwasnotpossibletoconstructfittedlogisticregressionmodelsfor thepredictionofM.freemanidistribution. Vegetation found at the sample points within the hotspot area for M. freemani included Clusia rosea, Philodendron sp, Languncularia racemosa, Manilkara bidentata, Mikania micrantha, Hedychium gardnerianum, Bromelia plumier, Erythroxylum havanense, Erythrina pallid,angiferaindica,Jacquinarmillaris,Pteridophytesp,andCoryanthes sp. Plant species present at both hotspots and confirmed M. freemani sites included Pitcairnia integrifolia, Coccoloba uvifera, and Cocos nucifera. Plant species present at confirmed M. freemani site only was themossBryopsidaspp.

93

Figure3ShowingMDSPlotwhereSampleSitesAccordingtoFactors. The semiquantitative presence/absence data was recorded at each transectintervalwasconstantthroughout(valuesbetween05),andasa result no transformations of the data were required. However, non metricmultidimensionalscalingofbioticandabioticfactorsrevealedno predictivepatternforeitherabioticorbioticfactors. Discussion The choice of environmental factors used in the predictive models (Maxent,logisticregressionandnMDS)wasbasedonobservationsofthe range of environmental characteristics at sample locations. Chapman (1967)notesthatsubstrateplaysamajorroleinpresenceorabsenceof plant species, and as a result, three factors were used to represent this variableinmodellinginthisstudy. Field observations suggest that M. freemani preferred to be above thesplashzonebutwithinthevicinityofseaspray,indicatingsomesea spraytolerance.Chapman(1967)alsonotesthatthelevelofsaltpresent in coastal environments is an importantdeterminant ofvegetation type inthesehabitats. Chapman(1967)alsocontendsthatharshnessofenvironmentisalso reflected in landscape stability. Degree of slope was another variable considered in this study because steeper slopes affect runoff and subsequenterosion.Thus,theabilityoftheplanttopersistinadynamic environmentwouldbeassessed.Exposure,asdeterminedbyaspect,was

94

used as a measure of stability and harshness of the environment (with respect to seaspray and erosion). Level of exposure of the coast was determinedbyprevailingwindsandcurrentsinthearea(CSO,2004).For this study area, north and west facing segments of the coast were designated as exposed, while south and east were sheltered. Land use was also modelled to determine whether M. freemani was located in higherriskareasofanthropogenicactivity(residentialandagriculture)or more naturalised conditions (forest, broken forest, coconut estate and scrubland). Maxent predicted that the northeast corner of the island was a hotspotforM.freemani,andthedominantenvironmentalvariableshere included a geology of Galera Grit (metasandstone), coastal classification ofrock,terracesoilformationandlanduseofcoconutestates. The random sample points, which fell within the vicinity of the hotspot, were found to be devoid of M. freemani. Phillips et al. (2006) suggest that human interference, biotic interaction and geographic barriers can explain such discontinuity in Maxent modelling. Human interference can include destruction of habitat and trampling. Biotic interactioncouldbecausedbycreationofanunfavourableenvironment (such as shade) or competition. Geographic barriers, such as habitat heterogeneity,wouldalsopreventM.freemanifromcolonisingasuitable habitat if the plant were unable to disperse across these boundaries. Errors in the model may have also resulted from other environmental factorsthatwerenotincludedinthemodel,suchaselevation.Phillipset al. (2006) noted that a minimum threshold requirement for occurrence localities has not yet been determined, nor has the degree of regularisation, to prevent model overfitting of deficient presence data, beenfinalised.GiventhesmallnumberM.freemanipresencesitesthere isastrongpossibilitythatthemodelwasoverfitted,potentiallyaffecting theaccuracyofresults. NocorrelationwasfoundbetweenthepresenceofM.freemaniand the betweensite abiotic variables recorded. From field observations M. freemani appears to colonise the face and edge of cliffs. This indicates that elevation and rocky substrate may be important factors. It may be able to survive under harsh conditions (rocky substrate, sea spray) and becomes confined to this area due to its poor competitive ability and shade intolerance. The Maxent model confirms the preference of rocky substrate. However, it also adds two new dimensions to this variable soiltypeandgeology.Apparently,M.freemanispresenceiscorrelatedto theoccurrenceofterracesoilandametasandstoneformationknownas Galera Grit. The former substrate type gives an indication of the

95

landform, but not the physiochemical properties of the substrate. Metasandstone is rock formed by metamorphosis of separate clades of sand grade rock, and Ali (1983) indicates that Galera Grit comprises of indurated quartzitic grit with calcite and quartz veins and joints. The soft (calcitic) veins and fissures may allow the roots of the plant to invade,enablingM.freemanitocolonisethehardrockysubstrate. Bubble plots identified Bryopsida spp. as the only species found exclusivelyatpresencesitesforM.freemani.However,examinationof intrasitedata,revealsthatthismossdidnotoccurconcurrentlywithM. freemani(thatis,bothspecieswerenotfoundwithinthesametransect interval). TheabsenceofbioticcorrelationmaybeduetoM.freemanispoor competitive ability, suggested by its location on the lower cliff faces. It creepsupinsomeinstances,reachingthetopofcliffsthatareclearedor sparselyvegetatedandrelativelyundisturbed.Remotesensingmayaidin habitatidentificationbymappingofthespectralsignatureofbarerocky land near the coast (Xie 2008). It should be noted that Maxent also correlatedcoconutestateswiththepresenceofM.freemani.Thetreesin a coconut estate are usually well spaced from their neighbours thus shade and competition would not prove to be an issues in this sense. Unfortunately, Cocos nucifera cannot be used as an indicator species givencoconutswidespreaddistributionalongthecoast. Several methods exist for assessment of the degree of threat to a species. Krupnick et al. (2009) give a preliminary conservation assessment algorithm that uses data from herbaria to perform an abundance,temporal,andspatialassessmenttoassessthreatstoagiven plant species. In the case of M. freemani, specimens have only been collectedafter1900,andfromfewerthan6localities.Giventhatitwas only collected three times prior to this study and two of those were beforethe1960s,thereisahighlikelihoodthatM.freemaniwarrantsa listingaspotentiallythreatened,usingKrupnicketal.s(2009)criteria. M. freemani also satisfies a considerable number of the characteristics of threatened species outlined by Primack (2006). Specifically,itoccupiesanarrowgeographicalrangeduetoitsspecialised niche requirements, has poor dispersal ability and lack of competitiveness. Field studies identified only a few extant populations, some of which may be in decline due to proximity to anthropogenic activity. This is further exacerbated by the fact that M. freemani is a perennial and may have a low rate of population increase. Thus, M. freemanihasahighlikelihoodofbecomingpronetoextinction.

96

During this study there was difficult to assess whether M. freemani wasthreatenedaccordingtoIUCNs(2008)criteriabecausemuchofthe data to apply these criteria was not available. Using best available evidence,toaddresstheIUCNcriteriaB2a(Areaofoccupancyestimated to be less than 500 km2) and B2biii (inferred decline of area, extent and/orqualityofhabitat),suggeststhattheclassificationofM.freemani as endangered is warranted. From field observations, there may be less than 2500 mature individuals of this species in northeastern Trinidad, but longterm population demographics comparisons are required to confirmthisestimate. Conclusion M. freemani has managed to hold on to its coastal niche in northeast Trinidad for the last 80 years, with collection sites remaining relatively unchanged, at Balandra, Galera and Toco. Its distribution appears to be limitedbygeologyandconfinedtothelittoralzone.Itmaybeadaptedto harsh conditions (rocky substrate and sea spray) due to its poor competitive ability. Its low abundance coupled with the increasing anthropogenic activity along the coast (in the form of pollution, recreationalactivity andcoastal development) may result indestruction ofitslimitedhabitat,potentiallymovingM.freemanisstatustocritical. Acknowledgements The primary author wishes to recognise the efforts of her parents who provided support both financially and in the field; Department of Life Sciencesfieldandtechnicalstafffortheirprovisionofresourcesandfield help;andmostimportantlyGodwithoutwhosegracenothingwouldhave beenpossible. References
AcevedoRodriguez, P. and Strong, M. 2007. The Flora of the West Indies. Smithsonian National Museum of Natural History. http://www.botany.si.edu/antilles/WestIndies/#floristic.Accessed6thDecember,2010. Ali,W.1983.NewsLetter5:GeologicalFieldTriptotheTocoDistrict.GeologicalSocietyof TrinidadandTobago.http://www.gstt.org.Accessed28thNovember,2010. APG II. 2003. An update of the Angiosperm Phylogeny Group classification for orders and familiesoffloweringplants:APGII.Bot.J.Linn.Soc.141:399436. Berwick, V., Cheek, L., Ball, J. 2005. Statistics Review14: logistic regression. Crib Care 9(1): 112118. Broken Hill Proprietary Company (BHP) Billiton. 2002. Atlas of Environmentally Sensitive Areas:EastCoastTrinidad. Central Statistical Office (CSO) of Trinidad and Tobago. 2004. Environmental Statistics Chapter9.http://www.cso.gov.ttAccessed28thNovember,2010.

97

Convention on Biological Diversity (CBD).2010. Country Profile Trinidad and Tobago. http://www.cbd.int/countries/profile.shtml?country=tt#status . Accessed 20th November,2010 Chapman,V.1976.CoastalVegetation,2ndend.Pergamon,Oxford. EnvironmentalManagementAuthority.2002.StateoftheEnvironmentalReport2001and 2002.http://www.ema.co.ttAccessed20thNovember,2010. EnvironmentallySensitiveSpeciesRules(ESS)2001,EnvironmentalManagementAct2000. TrinidadandTobago. ESRI.2005.ArcGIS.Redlands,CA:EnvironmentalSystemsResearchInstitute. Krupnick,G.,Kress,J.,Wagner,W.2009.AchievingTarget2oftheGlobalStrategyforPlant Conservation: Building a Preliminary Assessment of Vascular Plant Species using Data fromHerbariumSpecimensBiodiversityandConservation,18(6):14591474. InternationalUnionofConservationofNature(IUCN).2001.IUCN RedListCategoriesand Criteria : Version 3.1. IUCN Species Survival Commission. IUCN, Gland, Switzerland and Cambridge,UK. Liede, S. and Meve, U. 2004. Revision of Metastelma (ApocynaceaeAsclepiadoideae) in SouthwesternNorthAmericaandCentralAmerica.Ann.MissouriBot.Gard.91:3186. Phillips, S., Dudik, M., Schapire, R. 2004. A Maximum Entropy Approach to Species DistributionModeling.In:ProceedingsoftheTwentyFirstInternationalConferenceon MachineLearning.ACMPress,NewYork,pp.472486. Phillips, S. 2005. A brief tutorial on Maxent. ATand T Research http://www.cs.princeton.edu/~schapire/maxent/tutorial/tutorial.doc. Accessed 5thOctober,2010. Phillips, S., Anderson, R., Scharpire, R. 2006. Maximum Entropy Modeling of Species GeographicDistributions.Ecol.Model.190:231259. Primack, R. 2006. Essentials of Conservation Biology. Fourth Edition. Sinauer Associates, Sunderland,Massachusetts. Van den Eynden, V., Oatham, M., Johnson, W. 2008. How free access internet resources benefit biodiversity and conservation research: Trinidad and Tobago's endemic plants andtheirconservationstatusOryx42:400407CambridgeUniversityPress. Williams, R., Cheesman, E. and Philcox, D. 19281992. Flora of Trinidad and Tobago, Volumes13.GovernmentPrinter,PortofSpain,TrinidadandTobago. Xie,Y.2008.RemoteSensingImageryinVegetationMapping:AReview.JPlantEcol.1(1):9 23.

98

FireintheAripoSavannasEnvironmentallySensitiveArea: CausesandConsequences AditiBisramsingh1,2andMichaelOatham2


2

DepartmentofLifeSciences,TheUniversityoftheWestIndies,St.Augustine,Trinidad, WestIndies.Email:diti_78@hotmail.com 1 CorrespondingAuthor

Abstract The Aripo Savannas Environmentally Sensitive Area (ASESA) is the last tropicalgrasslandinTrinidadandTobagostillinitsnaturalstate.Manyof theorganismsfoundthereareuniquetotheareaandofgreatecological and scientific importance. While the Aripo Savanna remains relatively intact, it is today beginning to face anthropogenic disturbances. One of the more significant forms of anthropogenic disturbance is fire. These fires are caused by squatters using fire to clear land for agriculture or burning debris, and hunters using smoke to drive animals out of their burrows.Thisstudyinvestigatespatternsoffireoriginanddamageatthe ASESA and its effects on plant community composition in the open savanna.UsingForestryDivisionfirerecordsinaGISplatform,sixtyfour 50cm by 50cm quadrats in burnt and unburnt areas sampled to determine their plant species composition. Data analysis using the statistical software PRIMER showed that of all the plants species collected, Paspalum pulchellum and Rhynchospora barbata were the mostabundantspeciesatthesamplesites.Mostuniquecommonspecies suchasDroseracapillarisandUtriculariasp.werefoundinboththeburnt and unburnt areas. Two endemic species in the Polygalaceae family, Polygala exserta and Polygala adenophora, were found in the burnt areas. Burning may, therefore, induce flowering in these species. Other factors such as depth of hardpan, soil type, distance to savannah edge and the time since a fire last burnt the area, did notappear to have an effectontheplantcommunitiesinthesampleplots. Keywords Open savannas, fires, species composition, edaphic; squatting, hunting, managementplan. Introduction The Aripo Savannas Scientific Reserve (ASSR) lies in North East Trinidad (1035N,6112W)betweenArimaandSangreGrandeinalowlandarea

99

atanaltitudeof3540m,eastoftheCaroniPlain.Itisgenerallyflat,and gently rising to the north towards the town of Valencia. The savannas coveranareaofapproximately1788hectares,including250hectaresof open savanna habitat, which consist of ten separate savannas (Comeau 1990,EMA2007a).TheASSRlieswithintheLongStretchForestReserve (Laughlin2004). The ASSR, is the largest remaining natural savanna ecosystem with endemic flora in Trinidad and Tobago. This ecosystem also includes a marshformationthatconsistsofamixtureofmarshforest,palmmarsh andsavanna.Itprovidesahabitatforseveralrareandthreatenedspecies of plants and animals. In 2007, this site was designated as an Environmentally Sensitive Area, (the Aripo Savanna Environmentally SensitiveAreaASESA)undertheEnvironmentalManagementAct2000 (EMA2008a). Of the 457 species of plants identified in these savannas, 38 are restrictedtotheAripoSavannas,with16to20thatareconsideredrare orthreatened,and6thatarethoughttobeendemic(EMA2006).These uniquespeciesincludestheendemicsedgeRhynchosporaaripoensisand insectivorous plants such as the sundew and bladderworts. In addition, the ASESA contain the only palm/marsh forest in the country (Young 2006). Theedaphicconditionsintheopensavannahaveamajorinfluence on the vegetation. The soil is poorly drained and impoverished. During thewetseason,thesavannasarewaterlogged,givingrisetophysiological drought, causing restricted root growth and function due to lack of oxygen.Duringthedryseasonthesandysoildryoutrapidly,givingriseto a severe physical drought. As the subsurface claypan and fragipan dry out,theybecomeextremelyhardsothatrootscannotpenetratetothese lowerhorizons(Richardson1963). Savannas fires have been identified in a management plan for the ASESAasbeingoneofthemajoranthropogenicdisturbancesatthissite. Firescanhavedirectandindirecteffectsonseveralecologicalprocesses in tropical savannas, by affecting plant community composition and physiognomy (Silva et al. 1991). Fires in open savannas can alter the species composition by allowing fire resistant species to survive more than other nonfire resistant species (Comeau 1990). Fire is also an important source of mortality for plant seedlings and for small grasses and can affect the structure and functioning of the savanna (Silva et al. 1990). IntheAripoSavannas,allthefiresthatoccurareofanthropogenicorigin. The cause of fire is a result of humanactivities such as land clearing by

100

squattersforcultivation,unauthorizedprivatelandfires,negligencewith incendiary materials, hunting, mischief and land clearing to plant marijuana (Schwab 1988). The fires in the savannas also pose a serious problemtotheForestryDivisionwithregardtoimplementationoftheir management plan for the area, since the effects of fires on the plant communitiesarenotwellknown.(EMA2008a). Assuch,themainobjectiveofthestudywastoidentifythepatterns offireoriginanddamagetotheecosystemsintheASESAanddetermine theeffectsoffireonplantcommunitycompositionintheopensavanna ecosystems. Methods StudyArea AreasoftheASESAthathavebeenburntsince1987wereidentifiedusing Forestry Division fire records. These records were transferred to a geographicinformationsystem(GIS)and,theareasthatfrequentlyburn were identified, and annual maps of fires generated. Stakeholders were shown these maps in order to independently verify accuracy and the mapswereupdatedasnecessary.
Figure1.MapofAripoSavannasEnvironmentallySensitiveArea(ASESA)andthe locationoftheopensavannaswithintheASESA.

Surveytechniquesanddataanalysis Once the burnt areas were identified using the GIS, 100 random points weregeneratedforeightopensavannasusingArcGIS9.3(ESRI2005),to select sample points in burnt and unburnt areas. Although the Aripo Savanna consists of ten open savannas, sampling for this project was

101

doneinSavannas1,2,3,5andSavanna7.Ateachsamplepoint,50cmby 50cm quadrats were used to sample the plants present. If specimens could not be identified in the field, they were taken the National Herbariumforidentification. At each quadrat, the depth of the hard pan, soil type and the distanceofthenearestmarshforestweredetermined.Thedepthofthe hardpanwasmeasuredusingasoilcorer.Thedistanceanddirectionto thenearestmarshforest(savannaedge)wasestimatedvisually.Soiltype wasderivedfromapreviouslyproducedsoilmap. Plant species diversity, richness and plant community compositionbetweenareasoftheopensavannathathaveneverburnt, according to Forestry Division records, and areas that have been burnt were compared using Primer. In Primer, dendrograms were used to clusterquadratsbasedonsitesimilarity.MultidimensionalScaling(MDS) was used to investigate site similarities, and differences in species compositionattheburntandunburntsites.PrincipalComponentAnalysis (PCA)wasusedtodetermineifanyoftheenvironmentalvariableshadan effectonthespeciescomposition. Results FirerecordsfortheASESAwereonlyavailablefortheperiodfrom1997 tothepresent.Theseshowedthatoutofthetensavannas,savannah1, savanna 2, savanna 3 and savanna 5 were burnt every year during this period.Norecordoffiresintheothersavannaswasfound,norwerethe foresters in charge of the ASESA, aware of any other fires at the site. However, retired foresters indicated that the savannas were completely burnt in 1987 but no formal records for that period were available. Subsequentto1987,thesavannaswereindividuallyburnt,thoughmost oftheburningonlytookplaceinportionsofthesavannas.During2010, Savanna 5 was burnt in its entirety while other savannas were burnt in patcheswithinthesavanna. The fire records also showed that that the majority of these fires weretheresultofhumanactivities.ThefiresthatoriginateinSavanna1 wereignitedbysquatters,wholivealongtheboundariesofthesavanna and use fire to clear land for agriculture. These fires then spread into savannah2and3.Otherfiresinthesesavannas,particularlyinsavannah 5, were caused by hunters using smoke to drive animals. Fires in these savannaswerealsoaresultofburningofdebrisneartotheboundaryof the ASESA, smokers and malicious intent by residents. In most of these cases, the abundance of shrubs, grasses and surface litter provided an excellentsourceofignition.Theseshrubsandgrassesattheboundaryof

102

the ASESA easily catch fire, which then spread to the interior of the savannas. Rainfalldata,alsorecordedintheannualfiresreports,revealedthat all the fires in the ASESA occurred in the dry season (January to May), with the highest fire frequency occurring in the month of April. The average temperature during this time is 34C. This was once again demonstratedwhenin2010theentireofsavanna5wasburntduringthe dryseason. Within the 64 quadrats, only 24 species representative of the open savannaecosystemweredetected(Table1). These data showed that the two species, Polygala exserta and Polygala adenophora, were only found in burnt areas while three other species,Hyptislantanifolia,MyrciaplatycladaandRhynchosporacurvula wereonlyfoundinunburntareas.Themajorityofspecies,however,were foundinburntareas.
Table1PlantspeciesoccurrenceinburntorunburntquadratsfromtheASESA. Species Burnt Unburnt Total Paspalumpulchellum 36 20 56 Utriculariasp 32 17 49 RhynchosporaBarbata 24 14 38 Rhynchosporafiliformis 21 12 33 Droseracapillaris 19 8 27 Lagenocarpusrigidus 14 12 26 Andropogonvirgatus 14 2 16 Panicumstenodes 10 4 14 Habenarialeprieurii 6 2 8 Acisantherabivalvis 3 3 6 Panicumnervosum 4 2 6 Comoliaveronicifolia 2 3 5 Sauvagesiasprengelli 3 1 4 Polygalaexserta 4 0 4 Rhynchosporatenuis 3 1 4 Chrysobalamusicaco 1 2 3 Polygalaadenophora 3 0 3 Thrasyatrinitensis 2 0 2 Hyptislantanifolia 1 0 1 Myrciaplatyclada 1 0 1 Peramahirsuta 0 1 1 0 1 1 Panicumparvifolium Andropogonsp 1 0 1 Rhynchosporacurvula 0 1 0

103

The environmental data was analysed using Principal ComponentAnalysis(PCA)(Figure2).ThePCAresultsforthesamplesites andthedepthofthehardpan(Figure2a)showedthatthedepthofthe hardpanhadthegreatestinfluenceonthosesiteslocatedontheleftof PC1axisandthisinfluencedecreasedmovingfromlefttorightalongthe PC1axis. Inaddition,thepresenceofthespeciesAndropogonvirgatuswasnot influencedbythedepthofthehardpansincethisspecieswasfoundat depthsrangingfrom20cmto60cm(Figure2b). The same pattern was repeated for all other environmental variables and for all species at the sample sites. Depth of hardpan and Andropogon virgatus was chosen for illustrative purposes but the same resultswereseenfortheother24speciessuperimposedontotheother environmental variables. None of the environmental variables tested showed any influence on species composition at each of the sites. Likewise,thepresenceofaparticularspecieswasnotdeterminedbyany oftheenvironmentalconditionsmeasuredateachoftheareas.
(a) (b) Figure2.(a)PCAordinationforASESAsamplesitesshowingwherethedepthof hardpanhastheleastandgreatestinfluence.Thelargerthebubblesthegreater the influence.(b) PCA ordination for dept of hard pan but this time the species Andropogonvirgatuswassuperimposedontotheenvironmentalvariable(depth ofhardpan).

104

Discussion The Aripo Savannas Environmentally Sensitive Area (ASESA), though a legally protected area, remains threatened by anthropogenic activities that can affect the plant communities found there. In the sample plots used in this study, it was evident that anthropogenic fires can quickly spreadintotheinteriorofthesavannas.Theresultsofthisstudyrevealed that the fires in these areas originate mainly through squatters clearing land for agriculture and hunters. These two major threats remain prevalentbecauseofanumberofreasons.First,thereisinsufficientstaff attheForestryDivisiontoproperlymanageandmonitortheseactivities; second,thereisineffectivecontrolofaccesstotheASESA;third,thereis alackofadequatecontrolofhuntingintheASESA;andfourth,thereisan ineffectiveandinappropriatefiremanagementsystem(Nelson2007). Squatting, highlighted as one of the major cause of fires in the savannas, could be controlled through the implementation of a proper management plan (EMA 2006). These squatters chose to live on the boundaryoftheASESAsincetheyareunabletoobtainlandlegallyanda lackofenforcementmakesthesiteaccessibletoanyone(EMA2008b). Hunting, another cause of fires in the savannas, is as a result of hunters commercially exploiting the rare orchids and vertebrate wildlife such as macaws and some reptiles (EMA 2007a). While the enabling legislation of the ASESA does not allow hunting in this area, the lack of enforcement and patrols, and a local demand for wildlife continues to facilitatehuntinginthesavannas(EMA2007a). This study reveals that fires in the ASESA all took place in the dry season.Thismaybeexpectedsinceduringthesetimesthetemperatureis highwithanaverageof34C,andthelackofanysignificantprecipitation atthesite.Duringthedryseason,therearealotofdrygrasses,shrubs and surface litter in and around the savannas, and as such when a fire startsitcaneasilyspreadtotheinterior.Inaddition,theblackenedsoils atAripocausemoreheattobelostsoproducingdrierconditions,which favourthespreadoffires(Comeau1990). AlltherecordeddatasuggeststhatfiresintheASESAarearesultof human activities rather than natural factors, such as lightning that are characteristics of other Neotropical savannas. Fire in these other Neotropical savannas is considered a prime factor in determining the savannastructureandfunctioningsincefiresareanimportantsourcefor seeding mortality or germination and can even influence flowering in someshrubsandherbs(Silvaetal.1990).Savannasthathavebeenburnt frequentlycanalsoseeachangeinsomeoftheplantspecieswheresome

105

of these plants will begin to evolve so to become fire resistant (Chang 1996). The Aripo Savannas however is not dependant on fire for its structure. These savannas rely on edaphic factors for their existence. Currentevidencesuggeststhatnaturalfireswerenotfrequentenoughin thepasttohaveinfluencedthestructureofthesavannas(EMA2007b). However, illegal fires set within and around the savannas, can have effects on the native vegetation, altering resprouting patterns and floweringcharacteristicsofspecies(Schwab1988). Thesedatasuggeststhatoutoftwentyfourspeciesdetectedin the quadrats, five were only found in the burnt areas, three were only found in the unburnt areas alone while twelve species had more individuals in the burnt areas compared to the unburnt areas. Four specieshadnearequalproportionsinbothburntandunburntareas.For thespeciesthatwereonlyfoundintheburntareasonespeciesPolygala exserta, was endemic. However, while these species were detected in burntconditionstheirhabitatpreferenceortheeffectoffireonthemhas notbeenconclusivelydetermined.Thesmallnumberofindividualswithin thequadrats,andtheindividualscollectedmaynotberepresentativeof theentirespecies. Only three species were unique to the unburnt area. For these species,firemaybedestroyingtheirseedlingsorpreventinggermination (Silvaetal.1990).However,onceagainduetolackofadequatesample size, it cannot be concluded that fires were directly responsible for the patternsobserved. Forthosespeciesfoundinbothburntandunburntareas,thesedata suggest that fires may not have much effect on the mortality of these plants.ForspeciessuchasPaspalumpulchellum,Rhynchosporabarbata, Rhynchospora filiformis and Utricularia sp, more individuals were found intheburntareasthanunburntareas. It should be noted that out of the 64 sample areas, 41 plots were burntwhileonly23werefromunburntsites.Asaresult,giventhatthere were more samples in burnt areas, the data set would have captured morespeciesintheburntareas.Nodefinitiveconclusioncan,therefore, be made about the role of fire in germination or flowering at the study area. However, Schwab (1988) showed that most of the open savannas vegetationspecieswereabletoresproutinburntareas.Speciessuchas Paspalum pulchellum flowered profusely in the burnt areas and their densities in both burnt and unburnt areas were near identical (Schwab 1988). The data presentedhere suggests that species such as Paspalum

106

pulchellumwerefoundmorefrequentlyintheburntareasthanunburnt areas.Therefore,someoftheseplantsmaybeadaptingandevolvingto copewithfiresintheASESA. Even though some of the data suggests that some species are beginningtofavourburntconditions,thesedatadonotrepresentmany speciesattheASESAsinceonly24ofthe86plantspeciesdescribedinthe open savannas, were detected in the quadrats. How fires affect these otherspeciesremainsunknown. Inthisstudy,itwasthoughtthatenvironmentalvariablessuchasthe distance and direction of the nearest marsh might have an effect since fires originated at the edge of the marsh forest or the boundary of the savannas.Thus,thecloseraplotistotheedge,thehighertheprobability of it being burnt. However, MDS analysis revealed that this variable did notinfluencethespeciescompositionatthesampleplots.Thedepthof the hardpan/clay pan has been previously proposed as an important environmentalvariablesinceitinfluencesthedrainageconditionsinthe savannas(Richardson1963).However,datainthisanalysisshowedthat hardpandepthdidnotinfluencetheplantspeciescompositionbetween the64sampleplots. Soiltypeoredaphicconditionshaveamajorinfluenceonvegetation. There are basically four types of soil characteristic to Aripo; Type A Aripofinesandonfineclay,TypeBAripofinesandorsiltonsiltyclay, TypeCAripofinesandorsandonsandytogravellyclayandTypeT Transitional Aripo fine sand silt. As such, the soils here tend to become verywaterloggedinthewetseasonandverydryinthedryseason.The plants therefore that grow here are very well adapted to the soil conditions present (Richardson 1963). However, this analysis found that no one species favoured a particular soil type as they were spread out throughallthevaryingsoiltypes. From this study, therefore, the environmental variables measured didnotinfluencethespeciescompositionintheASESA.Asforfireandits effectonplantspecies,moresamplingneedstobedonetodetermineits effect on all 86 species in the savannas. It may be possible that some plants, especially the more common ones are beginning to become adapted to burnt conditions. Anotheruseful approach to determine the fulleffectsoffiresonthesespeciesintheASESAistomeasurethechange over time in permanent plots rather than attempting to determine differencesinspeciescompositiononasingleoccasionatvaryingplaces (LonsdaleandBraithwaite1991).

107

Once the full effects of fires are known, those species needing conservation and added protection can be identified, and the Forestry Divisioncanbegintoimplementtheirmanagementplanforthearea. Acknowledgements WethanktheForestryDivisionforprovidinguswithpermitstoenterthe ASESA to carry out research as well as to the foresters attached to the ForestryDivisionsCumutoofficefortheirknowledgeoftheareaaswell asforfieldassistancewhenneeded.WealsothankstaffattheNational Herbarium for their help in identifying the plants species collected. We are grateful for the assistance of the technicians attached to the Universityforprovidingfieldassistanceduringsampling. References
Chang, ChiRu. 1996. Ecosystem Responses to Fire and Variations in Fire Regimes. Sierra NevadaEcosystemProject:FinalreporttoCongress,volII,AssessmentsandScientificbasis formanagementoptions. Comeau,P.1990.SavannasinTrinidad.LivingWorld:JournaloftheTrinidadandTobagoField NaturalistClub. Environmental Management Authority (EMA). 2006. Terms of Reference, Participatory ManagementPlanningforAripoSavannasEnvironmentallySensitiveArea. EnvironmentalManagementAuthority.2007a.AripoSavannasEnvironmentallySensitiveArea Literature Review to Facilitate the Preparation of Management Plans. Prepared by the CaribbeanNaturalResourceInstitute(CANARI). EnvironmentalManagementAuthority.2007b.AdministrativeRecordfortheEnvironmentally SensitiveArea:AripoSavannasScientificReserve.PortofSpain,Trinidad. Environmental Management Authority. 2008a. Managing together: A summary of the managementplansfortheAripoSavannas,anEnvironmentallySensitiveArea.Preparedby the Caribbean Natural Resource Institute for the Environmental Management Authority. PortofSpain,Trinidad. EnvironmentalManagementAuthority.2008b.AripoSavannasEnvironmentallySensitiveArea Resource Management Plan: A framework for Participatory Management. Prepared by Caribbean Natural Resource Institute for Environmental Management Authority. Port of Spain,Trinidad. ESRI.2005.ArcGIS.Redlands,CA:EnvironmentalSystemsResearchInstitute. Lonsdale, W.M and R.W.Braithwaite. 1991. Assessing the effects of fire on vegetation in tropicalsavannas.AustralianJournalofEcology. Nelson, H. 2007. Working Report Summary Report. Resource Management Focus Group. Prepare by Caribbean Natural Resource Institute for Environmental Management. Unpublisheddata. Richardson, W.D.1963. Observations on the Vegetation and Ecology of the Aripo Savannas Trinidad.JournalofEcology,Vol51,No.2. Schwab, S.I. 1988. Floral and faunal composition, phenology, and fire in the Aripo Savannas ScientificReserve,Trinidad,WestIndies.M.Sc.Thesis,UniversityofWisconsin. Silva, J.F., J. Raventos and H.Caswell.1990. Fire and fire exclusion effects on the growth and survivaloftwosavannagrasses.ActaEcologica.Vol.11.

108

Silva, J.F., J.Raventos., H.Caswell and M.C. Trevisan. 1991. Population Responses to Fire in a Tropical Savannas Grass, Andropogon Semiberbis: A Matrix Model Approach. Journal of Ecology,Volume79,Issue2. Young,J.L.2006.AripoSavannaScientificReserve:ADescriptionandShortHistory.Quarterly BulletinoftheTrinidadandTobagoFieldNaturalistClub.

109

BiodiversityandbiogeographyoflichensinTrinidadandthe implicationsforforesthealthandbiosensitivity. AndreaScobie


GeographyDepartment,TheUniversityoftheWestIndies,St.Augustine,Trinidad,West Indies. Email:an8scobie@yahoo.com

Abstract There is no reference material and information on the distribution of lichens of Trinidad, at the National Herbarium of Trinidad and Tobago. This study investigated the biogeography and biodiversity of lichens in Trinidad through field surveys, which mapped the distribution of these plantsandprovidedvoucherspecimenstotheNationalHerbariumandto theNaturalHistoryMuseumofLondon,England.Thestudyinvolvedthe survey of forested areas for lichens, verification of species identity, and theirsubstrates.FiftyfivesitesinTrinidadwerestudied.Alllichenswere identifiedtothegenericandsometothespecificlevel.Twohundredand fiftysevenspecimensof35generawerenamed.Limitationsinthestudy included a lack of resources and taxonomic expertise in Trinidad. The distributionoflichensatthe55siteswasmapped.Furtherstudiesshould be undertaken to determine the effect of pollution, disturbance and foresthealthinTrinidadandTobago.Thisstudyprovidesafoundationfor a national inventory of lichens, from which biomonitoring and forest healthstudiescanemerge. Keywords voucher specimens, forest, microlichens, diversity, distribution, macrolichen Introduction Trinidad is a tropical continental island, which possesses a range of topography,avariablerainfallpatternandarangeofforesttypes,which encouragearichdiversity.Oneorganismcommoninthenativeforestsof Trinidadistheepiphyticlichen.Thisisaninconspicuousorganismand,as such, has received little study. In the past, scientists, such as Vainio (1923) and Jermy (1963), made collections of lichens from Trinidad, but novouchersfromthesecollectionsnowexistattheNationalHerbarium of Trinidad and Tobago. The lichen flora of the country thus remains comparativelyunknown.

110

The objectives of this study were to develop a reference collectionof lichens of Trinidad at the National Herbarium, provide duplicates to an international archive (Natural History Museum, UK), develop maps of lichen distribution, produce maps of lichen distribution, and investigate the relationship between environmental conditions and presence and density of the lichens at the study sites. It was hoped that these data would provide the basis for use of lichens as bioindicators and in bio monitoring. Method Fiftyfive study areas were selected randomly in Trinidad and sample plotsof0.25km2 insize,weresurveyedforlichens. TheGPScoordinates, elevation, soiltype, canopy cover, leaflitter and presence of surface water, were recorded for all sites where trees were sampled (Figure 1). From a staked central position, four suitable trees closest to that point, onefromeachquadrant,wereselected.Distancesfromcentralpointand circumferences were recorded. Four quadrat ladders were tied to the tree,oneateachcardinalaspect,andlichentypeswerecountedineach quadrat. Samples were then taken from the trunks and transported to thelabforprocessing. Figure1Locationof55lichensampleplotsinTrinidad.

111

Results The geographical information for 35 sites were recorded. Numbers of lichens found in each site are presented at Table 1, andFigure 2 shows the relation between the number of samples of the various genera and theirfrequencies.Acompletelistofthegeneraidentifiedinthisstudyis presentedatTable2.
Table1:SummaryofLichensamplescollectedfromforestedareasinTrinidad

Samples Collected Identified Generaidentified Speciesidentified CuratedatNHM CuratedatTandT Number 287 216 35 17 C190 287

Two samples, Parmotrema austrosinense (Zahlbr.) Hale (Canas et al., 1997),andDirinariapicta(Sw.)Schaer.ExClem.(Aptrootetal.1997)are showninFigure3.
70 60
NUMBER OFLOCATIONSAND Series1 Numberoflocations Series2 Frequencyofgenera

50 40 30 20 10 0

GENERA

Figure2:Thenumberoflocationswhereeachgenuswasfoundandthefrequency ofeachgenusinalllocations.

112

a) b) Figure3Photographsofa)Parmotremaaustrosinense(Zahlbr.)Hale(Canasetal, 1997)andb)Dirinariapicta(Sw.)Schaer.ExClem.(Aptrootetal.1997)

Discussion TrinidadslocationintheTropicsaffordsahighspeciesrichness and diversity.The topography, rainfall and forest types lend themselves tosuchdiversity. The collections of lichen by Vainio (1923) and Jermy (1963) reflected a preponderance of macrolichens. However, this study noted mainlycrustosemicrolichens.Thesewereabundantonthetreetrunksin the forested areas, along with some squamulose and foliose types. The smallestrepresentationwasthefruiticosetypes,yieldingsuchgeneraas Usneasp.,Ramalinasp.,Teloschistessp.andCladoniasp. The latter, with the exception of Cladonia sp., were located at the site of highest elevation in this study, Morne Bleu (699m). At this elevation, all ten genera were found. This site also held the largest number of fruiticose types. This is consistent with previous studies that notedfruiticosetypeshavetheirmaximumrichnessathigherelevations thanfoliosespecies(Baniyaetal.,2009). Thefoliosetypesweremoreabundantonnonforestedtreespecies, whichwerenotamajorpartofthisstudy.Someweremoreprevalentin wetterforestedareas.OneexampleisLeptogiumsp.,whichwasfoundin areas that were waterlogged or experienced high humidity or under a thickcanopy. The areas of most abundant crustose species were well lit, i.e. the canopies were light and areas were drier. Sterile crusts were most abundant. Graphis sp. and Pyrenocarp sp. were prominent among the crustosetypes.

113

Table2Lichenfrequencyfrom55forestsitesofTrinidad Genera Leptogium Porina Bacidia Graphis Chapsa Arthonia Herpothallon Usnea Ramalina Teloschistes Parmotrema Phyllopsora Pyrenocarp Dirinaria Gyalecta Opegrapha Myriotrema Dimerella Catillarea Letrouitia Thelotrema Fissurina Arthathelium Coccocarpia sterilecrust sorediatecrust Canaparmiella Pyrenula Crocynia Eschatagonia Heterodermia Physcia Lechonidia Clathroporina Perusaria numberoflocations 5 17 5 12 1 5 15 1 1 1 3 6 9 3 1 9 1 1 2 7 1 6 1 1 20 2 1 2 1 2 4 1 1 1 5 Frequency:n/Nx100 14.5 49.3 14.5 34.8 2.9 14.5 43.5 2.9 2.9 2.9 8.7 17.4 26.1 8.7 2.9 26.1 2.9 2.9 5.8 20.3 2.9 17.4 2.9 2.9 58 5.8 2.9 5.8 2.9 5.8 11.6 2.9 2.9 2.9 14.5

Conclusion ThediversityoflichensinTrinidadiscomparativelyhighwith35genera (17 species of which were identified). This was especially true of the crustose type. The biogeography of lichens in this study was consistent

114

with other studies, as seen in the relationship between elevation and speciesrichness. Acknowledgements Thanksgotomysupervisors:PaulShaw,YasminBakshComeau,Patricia Wolseley; Dr. Holger Ths (Natural History Museum); Mr. W. Johnson, Ms.K.ManaureandMr.BradBharat(NationalHerbariumofTrinidadand Tobago);myhusbandandchildrenfortheirkeensupport. References
Vainio,E.A.1923:LichenesininsulaTrinidadaproferroreR.Thaxtercollecti ProceedingsoftheAmercanAcademyofArtsandSciences58:131147. Jermy, C. 1963. Collection of Lichens of Trinidad. Natural History Museum, CromwellRd.,LondonU.K..Unpublished. Lumbsch, Mc Carthy and Malcolm. 2001. Key to the Genera of Australian Lichens. Apothecial Crusts. Flora of Australian Supplementary Series II. AustralianBiologicalResources,Canberra. Baniya,C.B.,TorsteinS.,YngvarG.andM.W.Palmer.Theelevationgradientof lichenspeciesrichnessinNepal.TheLichenologist42no.1(2010):8396. WillWolf, S., C. Scheidegger, B. McCune. Monitoring scenarios, sampling strategies and data quality, in Monitoring with Lichens Monitoring Lichens, edited by P. L. Nimis, C. Scheidegger and P. A. Wolseley 147 162. Kluwer AcademicPublishers,Netherlands2002.

115

ElectricalEnhancementofCoralGrowth:APilotStudy 1,* 2 L.S.Beddoe ,T.J.Goreau ,J.B.R.Agard1,M.George3,andD.A.T. Phillip1


1

DepartmentofLifeSciences,TheUniversityoftheWestIndies,St.Augustine,Trinidad,W.I. E 2 GlobalCoralReefAlliance,37PleasantStreet,Cambridge,MA02139,USA 3 DepartmentofPhysics,TheUniversityoftheWestIndies,St.Augustine,Trinidad,W.I. *CorrespondingAuthormail:lbeddoe@gmail.com

Abstract It has been proposed that weak direct current increases the aragonite form of calcium carbonate deposition in some corals, resulting in increased growth in coral skeletons. The validity of this claim remains unsubstantiated because of weaknesses in the experimental design of previous studies. The objective of this study was to develop a scientific method for testing the effect of low direct current on the growth of corals.Thispaperreportsonthefirstphaseofa3phasestudyondirect current effects on aragonite deposition. During this first phase, the experimental design was developed and tested in small pilot laboratory andfieldexperiments.Theresultssuggestedthatclosedsystemsrequire furtherexperimentswhenusingthemineralaccretionmethod.Thefield experiments showed that Millepora alcicornis was the most suitable species for further experimental studies and that a modified version of the buoyant weighing method was the most precise approach for measuringcoralgrowth. Keywords electrolytic mineral accretion, Biorock, Millepora alcicornis, brucite, aragonite,buoyantweighing,directcurrent(DC) Introduction Pastresearchhasexploredtheuseofelectrolysisasameanstoenhance accretion and calcification processes in coral reef restoration (Schuhmacher and Schillak 1994; van Treeck and Schuhmacher 1999). This approach involved passing a weak direct current (DC) through a metalmeshcausingtheprecipitationanddepositionofreeflikematerial, such as aragonite, CaCO3; and brucite, MgOH2; from seawater (Hilbertz 1992).

116

Mineral accretion techniques were first applied to coral reef restorationbyTomGoreauandWolfHilbertz(Hilbertz1992).According to Hilbertz (1992), higher current densities result in faster growth but weaker material dominated by brucite [Mg(OH)2], while lower current densities produce slower deposition dominated by harder aragonite [CaCO3].Thistheorywassupportedbysubsequentexperimentalstudies done by van Treeck and Schuhmacher (1997), and Sabater and Yap (2004). However, the validity of this claim remains disputed due to weaknesses in the experimental design of previous studies and lack of scientificallyreviewedevidence.Inaddition,previousstudieshaveuseda range of methods to determine evidence of growth, which included measuringtape,Alizarindye,andVerniercallipers,allofwhichhavehad limitations. The objective of this study was to develop a method with a proper design of experimental treatments and controls, which provided more precise measurement of growth rates than those that have been previouslyattempted.Itishypothesizedthatlowdirectcurrentincreases calcium carbonate deposition in some corals, resulting in increased growth.Previouslypublishedpaperssuggestthatlowercurrentdensity resultsinmorearagonitebeingdeposited,thusthefollowinghypothesis isproposed:Alowdensitydirectcurrentrangeof23A/m2willresultin anincreaseinthedepositionofthearagoniteformofcalciumcarbonate. Thispaperdiscussessomeoftheresultsofphaseone,whichdeveloped andtestedtheexperimentaldesign. MaterialsandMethods Previousresearchers(Hilbertz1992;vanTreeckandSchuhmacher1997; Sabater and Yap 2002, 2004) have used a range of anode materials, including graphite, steel and lead, but have obtained best results with titanium (van Treeck and Schuhmacher 1997). Titanium is known to be highlyresistanttoadverseconditions,andinthisstudy,atitaniumanode wasused. During this pilot study, the goal was to develop the experimental design for investigating aragonite deposition by lowdensity direct current.Inthisregard,thepilotstudyconsistedofanumberoflaboratory experiments and a small pilot field experiment at 1109'26.01"N 6050'20.81"Winfrontofabreakwaterbarrier. Coralfragmentsoflocalcoralspeciesandfragmentsfromthesame colony were used to reduce potential variance in growth due to intra specificvariability.Thestudysitecontainedboththechargedandcontrol experiments,andreplicated.Physicalparametersmeasuredinthestudy

117

included water depth; height, size, and shape of the experimental structure; and wave energy; and nature of the sea floor. All environmentalvariableswerekeptidenticalforboththechargedandthe control experiments, to ensure that measurements of electricitys influenceoncoralgrowthratecouldbemoreaccuratelydetermined. Asmallscalelaboratoryexperimentwassetupusingcoralfragments of Porites porites, collected from Toco (1050'09.2"N 6055'25.5"W) on the north east coast of Trinidad. The first experiment consisted of two aquaria,measuring76.2cmx45.72cmx45.72cm.Oneaquariumactedas thecontrolandtheotherreceivedlowDCof6V4.24A.AlizarinRedDye, of concentration identical to that used by Sabater, was used initially to stainthecoralssothatgrowthwouldbevisibleandmeasurementsmade usingVerniercallipers(SabaterandYap2002).Thestainlesssteelcathode was 2.54cm squares and the anode measured 33.02cm x 38.1cm. PVC frameworkssimilartoSabaterandYap's(2004)designweresubmerged intheseaquaria,withamodificationinthepositioningoftheelectrodes, specificallypositioningtheanodebelowthecathode. The second laboratory experiment consisted of four aquaria measuring 76.2cm x 45.72cm x 45.72cm each, two of which were the controlandtheothertwobeingthetesttreatments.Cathode1received a 3V 0.615A DC charge during the day, 6am to 6pm, and cathode 2 receiveda3V0.615ADCchargeduringthenight6pmto6am.Thiswasto determinetheoptimumtimeforapplyingelectricityinthefield. Thethirdlaboratoryexperimentconsistedoffouraquariameasuring 76.2cmx45.72cmx45.72cmeach,twoofwhichwerethecontrolandthe other two being the test treatments. A fifth aquarium, measuring 15.24cmx45.72cmx45.72cm,wasusedtohousetheanodeandasalt bridgewasusedtoallowtheflowofionsfromthisaquariumtothetwo aquariacontainingcathode1andcathode2(Plate1.1).Thesaltbridge,a componentofanelectrochemicalcell,producesanelectricalcurrentasa product of an oxidationreduction or redox reaction. The salt bridge electrically connected the two test treatments while keeping them separate and allowed electrons to transfer between the anode and cathode 1 and cathode 2. For this experiment, the salt bridge was a U tubedesignfilledwithsodiumchlorideagar.Adirectcurrentof6V4.24A DCwasused.

118

Plate1.1UtubeSaltBridgeforlaboratory3experimentalsetup Only one species of coral, Porites porites, was used during these laboratory experiments. A rubble bottom, consisting of ground coral fragments of varying sizes, was created to imitate the seabed. Each aquariumusedAquaClearfilter70andaproteinskimmer.AnAtinicbulb was used to replicatenightlight and fluorescentbulbsto representday withalightingperiodof12hrseach.Directcurrentwassuppliedusinga NipponAmericaDVP515DCregulatedpowersupply.Theseawaterwas takenfromToco(1050'09.2"N6055'25.5"W),thesitewherethePorites porites colonies existed. Seawater was changed weekly and parametric readingstakencontinuously. Inthefield,anelectricallyconductivedomeshapedframewasbuilt fromBRCandservedasthecathode(vanTreeckandSchuhmacher1997; Sabater and Yap 2004). This was submerged and anchored to the sea bottom and a titanium mesh was used as the anode. A light sensitive circuit was used to allow current during the daylight and no current at night.Themaincurrentsourceforthisexperimentwasa12Vcarbattery, withacircuitcurrentof6A.Thebatterywasallowedtorechargeduring thenight.Duringtheday,itwasexpectedthatthebatterychargerwould beofforatminimalchargingrate,thusensuringasteadycurrentflowto theexperiment.Whenmeasureditwasapproximately4.85A,lowerthan thedesignedcurrentduetotheaddedresistancecausedbythedistance ofthecabletotheexperiment. Initially live coral fragments of varying species were attached using steel binding wire to the cathode. However, after several months of observations 20 replicates of Millepora alcicornis were added to the chargedandthecontrolexperiments.

119

BothvanTreeckandSchuhmacherhavefoundthatadirectcurrent rangebetween1to24Vatadensityof3Am2onthecathodicsurface gavethebestresults(vanTreeckandSchuhmacher1997).However,Dr. GoreauandProf.WolfHilbertz(pers.comm.)suggestedarangeof23A per m2 of surface area at 4.5V to be ideal. For this experiment, the amperagerangeusedwas4.7A7.1A. Results Thefirstexperimentresultedinovernightbleachingandgaseouschlorine fumesbeingreleasedintothelabenvironment.ApHof7.4wasrecorded and the total Cl ion concentration measured was out of range. This revealed thatthe direct current,6V4.24A, was too high. The amperage was lowered for the second experiment to 3V 0.615A to eliminate the potential for bleaching. However, within ten days of the experimental setup both cathodes developed rustcoloured bacteria, which subsequently colonized the entire aquaria. This suggested that the amperagewastoolow.WhentheDCwasappliedtotheexperiment,the pH,decreasedfrom7.54to6.88forthecoralschargedduringtheday. Asaltbridgewasusedforthethirdlabexperimenttoreducethe amount of ions being released and the DC was increased to 6V 4.24A. However,within18days,bleachingoccurredatthebaseofthecoralsand rustcoloured bacteria developed on the cathode of the control. There were visible signs of accretion on the charged cathodes (Figure 1), but eventuallythecoralswereallbleachedafterseveralmoredays.
Figure 1 Photograph of coral fragments following eighteen days of accretion on thecathodeusingthesaltbridgedescribedinexperimentthree.

120

In the field, live coral fragments of various species were initially attached so that the correctrange of direct current could be confirmed and the bestadapted coral species for the study site identified. It was determinedafterthreemonthsthatMilleporaalcicornisandMontastrea annulariswereideallysuitedforthisexperiment.However,Milleporasp. wasthefastestgrowingandeasiesttoobtainofthecoralspecies.Photo transects of the replicates of Millepora were taken to assess growth as measurementsusingVerniercallipersandmeasuringtapewerefoundto be inaccurate. Photo transects were also not effective because of the differentplanesofgrowth.Asaresult,onlyqualitativedatawascollected (Figure2).
Figure 2 A representative fragment of Millepora alcicornis showing growth from thepilotfieldstudyoversixmonths.

Discussion The bleaching process of corals in the laboratory experiments showed that in a closed system likean aquarium,a viable experimental method for investigating mineral accretion remains to be developed. Even with useofasaltbridge,itwasnotpossibletomaintainstableexperimental conditions.Asaresult,thelaboratoryexperimentswereabandoned. The pilot fieldstudy suggested that the thickness of accreted material was greater near the anode and cathode connections and declined as one moves further away from these connections. The electrolyticconfigurationusedinthispilotstudyyieldedvisiblysignificant changes in growth in the charged coral fragments compared to the controls. This study also showed that Millepora alcicornis was the most suitable species for the field experimental site, which had poor water quality.

121

ItwasalsonotedthatphototransectsofthereplicatesofMillepora sp. were the least effective form of growth measurement. Similarly, Vernier callipers measure length which is a two dimensional planar measurement, were also inappropriate. Since corals grow in various planes, volume or mass measurements were needed to give measurements in three dimensions. As a result it was deduced that buoyantweighingprovidedamoreprecisemeanstomeasureabsoluteor relative growth rates (Davies 1989). The buoyant weighing method was adaptedforfieldapplicationandwillbeappliedinfuturephasesofthis study.Coralsspendalotofenergybuildingskeletalmaterial,andinthe face of global climate change and its stressors of higher sea surface temperatures and ocean acidification, it is important that a better understanding of mechanisms for mineral accretion in coral growth is developed. Acknowledgement We would like to thank the late Prof. Wolf Hilbertz for his valuable trainingandexpertise,Mr.FaizulMohammed,Mr.AviBhagan,Dr.Azad Mohammed,Mr.DeosaranDocPersad,Mr.MarlonGeorgeandthestaff oftheelectronicsworkshop,PhysicsDept.,Dr.JennieMallela,TheDept. of Life Sciences, The Tobago House of Assembly, Undersea Tobago Dive Shop,CocoReefResort,TheUniversityoftheWestIndiesandvolunteers consisting of postgraduates, undergraduates, friends, family, locals and tourists. References
Davies, P.S. 1989. Shortterm growth measurements of corals using an accurate buoyant weighingtechnique.MarineBiology101(3):389395. Hilbertz,W.H.1992.Solargeneratedbuildingmaterialfromseawaterasasinkforcarbon. Ambio:126129. Sabater, M.G., and H.T. Yap. 2002. Growth and survival of coral transplants with and without electrochemical deposition of CaCO3. Journal of Experimental Marine Biology andEcology272(2):131146. Sabater, M.G., and H.T. Yap. 2004. Longterm effects of induced mineral accretion on growth, survival and corallite properties of Porites cylindrica Dana. Journal of ExperimentalMarineBiologyandEcology311(2):355374. Schuhmacher,H.,andL.Schillak.1994.IntegratedElectrochemicalandBiogenicDeposition ofHardMaterialANaturelikeColonizationSubstrate.BulletinofMarineScience,552 (3):672679. vanTreeck,P.,andH.Schuhmacher.1997.Initialsurvivalofcoralnubbinstransplantedbya new coral transplantation technologyoptions for reef rehabilitation. Marine Ecology ProgressSeries150(1):287292. vanTreeck,P.,andH.Schuhmacher.1999.Artificialreefscreatedbyelectrolysisandcoral transplantation: An approach ensuring the compatibility of environmental protection anddivingtourism.Estuarine,coastalandshelfscience(Print)49:7581.

122

PopulationdensityoftheagoutiDasyproctaleporinaatthe CentralRangeWildlifeSanctuary,Trinidad. HowardP.Nelson1,5,IndiraOmahMaharaj2,NadraNathaiGyan3, andAntonyRamnarine4.


DepartmentofLifeSciences,TheUniversityoftheWestIndies,St.Augustine,Trinidad, WestIndies.Email:howien@hotmail.com 2 WildlifeConservationCommittee,c/oForestryDivision,LongCircularRoad,St.JamesPort ofSpain,Trinidad,WestIndies 3 ZoologicalSociety,EmperorValleyZoo,St.Clair,PortofSpain,Trinidad,WestIndies 4 ForestryDivision,LongCircularRoad,St.JamesPortofSpain,Trinidad,WestIndies 5 CorrespondingAuthor
1

Abstract TheredrumpedagoutiDasyproctaleporina(Linnaeus,1758)isoneofthe mostheavilyhuntedgamemammalsontheislandofTrinidad.Resultsof a 2007 population survey of agoutis at the Central Range Wildlife Sanctuary in Trinidad are reported here. Thirty 1km transects were systematically surveyed within the 2,153 ha Central Range Wildlife Sanctuary using trained survey personnel from the Forestry Division. Surveys were conducted during the morning and afternoon over a one month period, and these data were pooled to estimate population densities for agoutis. Transect data were analysed using Distance 6.0 software and Kings estimator. Total transect distance surveyed during the study was 60 km and 20 agoutis were encountered. Agouti density was estimated at 10.43 individuals per km2 (95% C.I. 9.68 11.18 individualsperkm2)usingKingsestimator,whilethatusingDistancewas 8.38individualsperkm2(95%C.I.4.416.1individualsperkm2).These data were compared to reported densities in other Neotropical sites whereD.leporinaisknowntooccur.Thisstudysuggeststhatevenwithin the nominally protected Central Range Wildlife Sanctuary, D. leporina densitiesweresignificantlybelowthereporteddensitiesforthisspecies elsewhereintheNeotropics. Keywords game mammal, population estimate, distance sampling, line transect, Kingsestimator Introduction Wildmammalsprovideimportantfoodandrecreationalopportunitiesfor local peoples in the Neotropics (Redford and Robinson 1991). This is

123

especially so on the Caribbean island of Trinidad where the hunting of game mammals for subsistence, recreational and commercial activity is animportantforestuse(Asibey1984).Suchimportanceisreflectedinthe over 10,000 hunting licenses now issued annually by the countrys Forestry Division, for the harvest of game species (Wildlife Section, Forestry Division unpublished data). Among the mammals hunted on Trinidad,theredrumpedagoutiDasyproctaleporina(Linnaeus,1758)is oneofthemostimportanttargetsforrecreationalhuntingandthemost heavilyhuntedgamemammalsinTrinidad(Nelson1996). The redrumped agouti is a 36 kg Caviomorph rodent, which has a comparativelywidespreaddistributionintheNeotropics(Eisenberg1989; EmmonsandFeer1997;Nowak1999).Theserodentscanattainrelatively high densities in some tropical forests (Jorge and Peres 2005) and are important seed predators and seed dispersers (Rankin 1978; Smythe 1986;ForgetandMilleron1991;Forget1996;SilviusandFragoso2003). In this way, agoutis can affect composition and spatial distribution of theirfoodtreesintheseforests(Asquithetal.1999;Terborghetal.2008; Galettietal.2010).Additionally,thisspeciesisanimportantpreyspecies for terrestrial carnivores such as the threatened ocelot Leopardus pardalis (AliagaRossel et al. 2006), which is the largest terrestrial carnivoreontheislandofTrinidad. In spite of this rodents importance as a keystone species in the tropicalforestsonTrinidad,anditshighvalueasagamemammal,there havebeenrelativelyfewattemptstoestimatetheagoutisabundancein theremainingforestsontheisland.Previousstudieshavesuggestedthat densities of this game mammal are well below theoretical carrying capacityexpectedfortropicalforestsonTrinidad,andthatthespeciesis beingexploitedatrateshigherthanwhatcouldbeconsideredsustainable (Nelson1996). In 2007, the Wildlife Conservation Committee (WLCC), an advisory committee constituted under Trinidad and Tobagos Conservation of Wildlife Act, commissioned a survey of the Central Range Wildlife Sanctuary (CRWS), a wildlife conservation area on Trinidad (Bacon and ffrench 1972). The objective of this study was to assess the status of game species within the CRWS and refine survey methodologies for a future national survey. The density estimates of D. leporina within the CRWS based on this survey are reported here and compared with abundance data from other South American localities. The implications forsustainableharvestingagoutisinTrinidadarealsoconsidered.

124

StudyAreaandMethods The Central Range Wildlife Sanctuary lies at the headwaters of the Tumpuna and Talparo watersheds in the central part (UTM Zone 20N 692069N,1155165E)oftheislandofTrinidad(Baconandffrench1972). This wildlife sanctuary consists of 2,153 hectares of undulating topography that is mostly below 200m in elevation. The climate at this site is seasonal and determined by variation in precipitation, with a dry seasonthatextendsfromJanuarytoMay,andawetseasonfromJuneto December,whilethemeanannualprecipitationiscaapproximately230 cm. The native vegetation of this area consists of tropical moist forests (Nelson2004),whileedaphicconditionswithinthesanctuaryproducesa vegetation mosaic at the standlevel, which fall into three forest associations Crappocarat, Crappococorite and Acurelfiguer (Beard 1946).Sincetheearly1950s,alittleunderonethirdoftheSanctuaryhas been converted to a teak, Tectona grandis, plantation monoculture (Figure1). Figure 1 Location of Central Range Wildlife Sanctuary and extent of naturalforestsinTrinidad. Surveys of agoutis were conducted along thirty systematic 1km transects.Theinitiallocationofthefirstofthesetransectswasselected at random using ArcGIS 9 (ESRI 2005), and each subsequent transect placedtoensureregularspacing(usingeitheranorthsouthoreastwest alignment) of the remaining 29 transects. Distance surveys (Buckland et al.2004;Bucklandetal.2001)wereconductedduringthemonthofApril 2007,inthemiddleofthedryseason,witheachlinetransectsampledat leastonceduringthemorningandagainduringtheafternoon.Distance to animals and sighting angles were estimated using electronic rangefinders and handheld compasses, and density of agouties

125

estimated using Kings estimator (Glanz 1990; Wright, Gompper, and DeLeon 1994) and Distance 6.0 software (Thomas et al. 2009). Kings estimator calculates the density of a population (D) using the formula D=n/(2lR), where N is the number of observed individuals, l is the total transect length and R is the mean distance of the animal from the observer.ForanalysesconductedinDistance6.0,theAkaikeInformation Criterion (AIC) (Burnham and Anderson 2002) was used to identify the mostsuitabledetectionfunctionmodelforourdata,specificallyadopting athresholdvalueofdeltaAIClessthan6(Richards2008). Results Atotalof60kmoflinetransectsweresurveyedwithintheCentralRange WildlifeSanctuaryduringthisstudy,with20agoutisdetectedalongthese linetransects. Using Kings estimator, the density of D. leporina within the CRWS protected areas was 10.48 individuals per km2, with a 95% confidenceintervalof9.6811.18individualsperkm2. Analyses of the line transect survey results using Distance 6.0 suggested that the most conservative model that fitted the data was a negative exponential model with a simple polynomial expansion. This modelgavethelowestdeltaAIC,andaKologmorovSmirnofgoodnessof fit value that was not significantly different from the observed data (p=0.77). Using this model Distance 6.0 predicted D. leporina density at theCRWSat8.38individualsperkm2,witha95%confidenceintervalof 4.3816.05individualsperkm2.. These densities from this study site were then compared with reported densities for D. leporina from three other Latin American sites wherethisspeciesisknowntooccur.ThesedataarepresentedatTable 1. Discussion This study represents the first time that a density estimate of the red rumped agouti populations at the Central Range Wildlife Sanctuary is beingreported,andoneofthefewattemptstoestimatethedensityof game species from the island of Trinidad, through direct surveys. When comparedwithdatafromotherNeotropicalsites,thisfieldstudysuggests that the density at the CRWS is at least onehalf to one quarter the densityofotherlocationswhereD.leporinaoccurs.

126

Table1DensityestimatesofD.leporinainNeotropicalsites. Location Density Hunting ForestType (individuals Intensity /km2) CentralRange 10.43*, Low Moist Wildlife 8.38 moderate tropical Sanctuary, (?) Trinidad PinkaitiResearch 31 NoneLow Semi Station,Brazil deciduous tropical IlhadeMaraca, 40 NoneLow Semi Roraima,Brazil evergreen seasonal Manaus,Brazil 1026 unknown Tropical rainforest Trinity Hills 2.10 High Moist Wildlife tropical Sanctuary/ VictoriaMayaro Reserve,Trinidad *Kingsestimator,Distance6.0modelestimate

Source

Thisstudy

Jorgeand Peres2005 Silviusand Fragoso 2003 Jorge2008 Nelson1996

The limits of the data collected in this study are recognised, particularly with regard to the use of Distance 6.0 to analyse agouti detections. Density estimates using this programme for fewer than 60 observations may not be robust (Buckland et al. 2001). This in part accounts for the wide confidence intervals reported for D. leporina densityestimatebasedonthismodel.However,itisinstructivethatthe densityestimateusingKingsestimatorisveryclosetothatpredictedby Distance 6.0, and well within the confidence limits for that model. This suggeststhatthesevaluesprovideareliableindexofagoutidensityatthe CRWS. Withagoutibeingheavilyhuntedontheisland,theCRWSmay beperforminganimportantmetapopulationrole,byprovidingarescue effecttosurroundingforestswhereintensehuntingcouldbedepleting densitiesofthisrodent.Asaresult,thefateofagoutipopulationsatthe CRWSmayhavesignificantimpactsonhuntingsuccessinthesurrounding Central Range Forest Reserve. It should be noted that at the Victoria Mayaro Reserve, another forest site in Trinidad with intense hunting pressure, agouti populations have been suggested to be as low as 2.1 individualsperkm2(Nelson1996).

127

Additionally, this study points to a potential impact on longterm forest structure and composition, should agouti populations at this site remaindepressed.Specifically,otherauthorshavedemonstratedthatthe loss of species such as agoutis can have a significant impact in the long term on forest structure and composition, by affecting seed dispersal (Asquithetal.1999;Terborghetal.2008;Galettietal.2010). In this context, there is a need to understand the reasons for the comparatively low densities of D. leporina within the CRWS, given its status as a protected area. The development of a longterm population dataset on the status of this rodent at the CRWS will provide a clearer understanding of the metapopulation dynamics and its contribution to sustainabilityofhuntinginsurroundingforests. Acknowledgments WewishtothankthestaffoftheWildlifeSection,ForestryDivision,for their logistical support, and to the field staff of the Divisions South Central Conservancy who assisted with the collection of the field data. We would also like to recognize the support of Dr. David Rampersad of The University of West Indies Business Development Office and the Wildlife Conservation Committee of the Ministry of Housing and the Environment. This study was supported by a grant from the Wildlife SectionsPublicSectorInvestmentProgrammesAssessmentoftheStatus ofWildlifeProject. References
AliagaRossel,Enzo,RicardoS.Moreno,RolandW.Kays,andJacalynGiacalone.2006.Ocelot (Leopardus pardalis) Predation on Agouti (Dasyprocta punctata). Biotropica 38, no. 5 (September):691694. Asibey,E.O.A.1984.EconomicroleofwildlifeinTrinidadandTobago.Rome.doi:FAOFO DP/TRI/79/001. Asquith,NigelM.,JohnTerborgh,a.ElizabethArnold,andC.MailnRiveros.1999.Thefruits the agouti ate: Hymenaea courbaril seed fate when its disperser is absent. Journal of TropicalEcology15,no.2(March):229235. Bacon,P.R.,andR.P.Ffrench.1972.ThewildlifesanctuariesofTrinidadandTobago.Portof Spain:WildlifeConservationCommittee,MinistryofAgriculture,LandsandFisheries. Beard,J.S.1946.TheNaturalVegetationofTrinidad.OxfordForestryMemoire20. Buckland,S.T.,D.R.Anderson,K.P.Burnham,J.L.Laake,D.L.Borchers,andL.Thomas.2001. IntroductiontoDistanceSampling.London:OxfordUniversityPress. .2004.AdvancedDistanceSampling.London:OxfordUniversityPress. Burnham, K. P., and D. R. Anderson. 2002. Model Selection and Multimodel Inference: A PracticalInformationTheoreticApproach.2ndeditionSpringerVerlag,NewYork. Eisenberg,J.F.1989.MammalsoftheNeotropics,vol.1.:Thenorthernneotropics.Chicago: UniversityofChicagoPress. Emmons, L.H., and F. Feer. 1997. Neotropical rainforest mammals: a field guide. 2nd ed. Chicago:UniversityofChicagoPress.

128

ESRI.2005.ArcGIS.Redlands,CA:EnvironmentalSystemsResearchInstitute. Forget,P.M.1996.RemovalofseedsofCarapaprocera(Meliaceae)byrodentsandtheirfate in rainforest in French Guiana. Journal of Tropical Ecology 12, no. 06: 751761. http://journals.cambridge.org/abstract_S0266467400009998. Forget,P.M.,andTarekMilleron.1991.Evidenceforsecondaryseeddispersalbyrodentsin Panama.Oecologia87:596599. Galetti, Mauro, Camila I. Donatti, Carla Steffler, Julieta Genini, Ricardo S. Bovendorp, and Marina Fleury. 2010. The role of seed mass on the caching decision by agoutis, Dasyproctaleporina(Rodentia:Agoutidae).Zoologia(Curitiba,Impresso)27,no.3(June): 472476. Glanz,W.E.1990.Neotropicalmammaldensities:HowunusualisthecommunityonBarro ColoradoIsland,Panama.InFourneotropicalrainforests,ed.A.H.Gentry,287313.New Haven:YaleUniversityPress. Jorge, MSP. 2008. Effects of forest fragmentation on two sister genera of Amazonian rodents(MyoproctaacouchyandDasyproctaleporina).BiologicalConservation141,no. 3:617623. Jorge, MSP, and Carlos a. Peres. 2005. Population Density and Home Range Size of Red RumpedAgoutis(Dasyproctaleporina)WithinandOutsideaNaturalBrazilNutStandin SoutheasternAmazonia1.Biotropica37,no.2(June):317321. Nelson,H.P.1996.EcologicalStudiesofForestMammalsintheWestIndieswithaFocuson Trinidad.TheUniversityoftheWestIndiesSt.Augustine. .2004.TropicalforestecosystemsofTrinidad:ecologicalpatternsandpublicperceptions. UniversityofWisconsinMadison. Nowak,R.M.1999.WalkersMammalsoftheWorld.6thed.Baltimore:TheJohnsHopkins UniversityPress. Rankin, J.M. 1978. The influence of seed predation and plant competition on tree species abundancesintwoadjacenttropicalrainforests.UniversityofMichigan. Redford,K.H.,andJ.G.Robinson.1991.SubsistenceandcommercialusesofwildlifeinLatin America. In Neotropical wildlife use and conservation., ed. J. G. Robinson and K. H. Redford,623.Chicago:UniversityofChicagoPress. Richards, S. A. 2008. Dealing with overdispersed count data in applied ecology. Journal of AppliedEcology45,no.1(February):218227. Silvius, K.M., andJ. Fragoso. 2003. Redrumped Agouti (Dasyprocta leporina) Home Range UseinanAmazonianForest:ImplicationsfortheAggregatedDistributionofForestTrees. Biotropica35,no.1:7483. Smythe, N. 1986. Competition and Resource Partitioning in the Guild of Neotropical Terrestrial Frugivorous Mammals. Annual Review of Ecology and Systematics 17, no. 1 (November):169188. Terborgh, John, Gabriela NuezIturri, Nigel C a Pitman, Fernando H Cornejo Valverde, Patricia Alvarez, Varun Swamy, Elizabeth G Pringle, and C E Timothy Paine. 2008. Tree recruitmentinanemptyforest.Ecology89,no.6(June):175768. Thomas,L.,J.L.Laake,E.Rexstad,S.Strindberg,FFCMarques,S.T.Buckland,D.L.Borchers, etal.2009.Distance6.0.ResearchUnitforWildlifePopulationAssessment,Universityof St.Andrews,UK. Wright,S.Joseph,MatthewE.Gompper,andBonifacioDeLeon.1994.AreLargePredators KeystoneSpeciesinNeotropicalForests?TheEvidencefromBarroColoradoIsland.Oikos 71,no.2(November):279.

129

Abstracts

130

NaturalHistoryandConservationoftheTrinidadPipingGuan KerrieNaranjit
DepartmentofLifeSciences,TheUniversityoftheWestIndies,St.Augustine,Trinidad,West Indies.Email:kerriemonster@gmail.com

Abstract

The Trinidad Pipingguan or Pawi, Pipile pipile, is a critically endangered NewWorldcracid,endemictoTrinidad.Duetohabitatlossandhunting, it is considered the second most threatened cracid worldwide and an immediate conservation priority by the IUCN/Birdlife/WPA Cracid Specialist Group. This paper presents the findings of a twoyear field studythatassessedthenaturalhistoryofthePawi.TheTrinidadPiping guanwasobservedfor26%ofthe762fieldhoursandduring73%ofthe 304fieldsessions.Thesebirdsweremostactiveduringtheearlymorning when they displayed foraging, preening, aggressive and courtship interaction behaviours. The Pawi appears to be primarily arboreal, with lessthan1%ofobservedactivitiesoccurringontheground.Dietconsists mainly of fruits, leaves, flowers and seeds. Group size ranged from 18 individuals with an average group size of 1.3 birds. The phenology of behavioursuggeststhatthePawihaveanextendedbreedingseasonwith chicks being first observed in February and mating and courtship behaviour observed as late as July. Significant differences in wing drummingandlongpipingvocalisationswerefoundbetweenthedryand rainy seasons, suggesting that these behaviours are related to reproduction. Attempts were made to mistnet birds for radio tracking; however, these attempts were unsuccessful. The information gained fromthisstudywillbeusedtopreparerecoveryandmonitoringplansfor thespecies. Keywords:pipingguan,Trinidad,Pipile,behaviour,diet,groupsize

131

BiofoulingonrecreationalvesselsinTrinidadandTobago JudithF.Gobin1,AlanaJute1andAnuradhaSingh2
1

DepartmentofLifeSciences,TheUniversityoftheWestIndies,St.Augustine,Trinidad, WestIndies.Email:Judith.Gobin@sta.uwi.edu 2 DepartmentofBiology,COSTAAT,PortofSpain,Trinidad,WestIndies.

Abstract
The twinisland state of Trinidad and Tobago has seen a significant growthintheyachtingindustryoverthelast30yearswiththenumbers of boats having increased 10fold between 1980 and 2000. Being geographically outside of the hurricane belt, the islands offer a prime hurricane shelter in the Caribbean with excellent boating and repair facilities. The Chaguaramas coastline (Trinidad) has seven anchorages, oneofwhichtheChaguaramasBaywastheselectedstudysite.Thirty tworecreationalvesselsweresampledforbiofoulers.Overall,hullfoulers belonged to six major phyla: Plantae (macroalgae), Bryozoa, Porifera, Mollusca, Crustacea, and Annelida. To date, 50 species have been identified: 25 annelids, 14 barnacles and 6 bivalves (including Perna viridis, a known introduced species). Preliminary identifications suggest thatatleastonespecies(Megabalanuszebra)mayhavebeenintroduced toTrinidadswatersviathispathway.Thisresearchhasprovidedthefirst specieslistofmacrofaunalfoulersofrecreationalboatsforTrinidadand Tobago.Additionally,thestudyhascontributedtoourknowledgeoflocal biodiversity data, while attempting to address the issue of introduced speciestoourwaters.

Keywords biofouling, Chaguaramas, introduced species, Tobago, Trinidad, Perna viridis,Megabalanuszebra

132

PatternsofbiodiversityinTrinidadianspiders JoanneSewlal
DepartmentofLifeSciences,TheUniversityoftheWestIndies,St.Augustine,Trinidad,West Indies.Email:joannesewlal@gmail.com

Abstract
Trinidad is a continental island, which was isolated from South America about 10,000 years ago. Habitat types on Trinidad and the fauna they contain are, therefore, representative of the northeastern part of the neighbouring continent. Trinidads small size makes studies of its biodiversitymoremanageable,whileatthesametimeprovidinginsights about South American biodiversity, which is littleknown for most invertebratetaxonomicgroups. In this study, the biodiversity of three orbweaving spider families, Araneidae, Nephilidae and Tetragnathidae, were examined in natural habitats. This study also looked at how factors such as habitat classification and geographic location affected spider biodiversity. According to Beard (1946), the natural vegetation of Trinidad can be classified into six formations containing 16 habitat types. Data was collected using the visual search and sweepnetting methods at 46 localities throughout the island. Biodiversity was determined by examining the observed and estimated species richness, species distribution, distribution models and diversity indices between the formationsandhabitattypes. It was found that there was a high proportion of rare species, and that species of intermediate abundance were also found frequently throughout the study sites. Spider communities found in Trinidad were simple in nature and their ecology appeared correlated with a single factor,canopycover,whichseemedtohaveaneffectonbiodiversitywith respect to species richness, diversity, evenness and dominance. Formation type had influence on observed species richness, species diversity and dominance but neither formation nor habitat influenced evenness.Geographicfactorssuchaslatitude,longitudeandaltitudedid not influence diversity. The likelihood that climate change will affect abundance, species diversity, and species composition of spider communitiesinTrinidadisdiscussed.

Keywords habitat classification, orbweaving spider, species richness, species diversity,Trinidad

133

Aninitialinvestigationintothethirdrecordedmassbleaching eventinTobago JahsonAlehmu


EnvironmentalResearchProgramme,InstituteofMarineAffairs,HilltopLane,Chaguaramas, Trinidad,WestIndies,Email:jalemu@ima.gov.tt

Abstract
Masscoralbleachingisaphenomenonthataffectsmultiplespeciesover a large area. Historically, Tobagos reefs have experienced extensive bleaching events during 1998 and2005. Instances of milder, small scale bleachingwerealsoreportedin2000,2001and2003,affectingveryfew species.InSeptember/October2010,Tobagobeganexperiencingitsthird recordedmassbleachingevent.Thisinvestigationintocoralbleachingin Tobago was initiated in August 2010, to determine the extent of the event,speciesaffected,thedegreeofmortality,andtoidentifysiteswith good recovery, using qualitative and quantitative coral surveys. At the onset(August2010),bleachingwasmoreprevalentonshallowreefzones (~5m)comparedtothereefslopes.However,itlaterspreaddowntothe lowerslope(33m),andwasmostsevereat1015m.Corals,(includingthe reef building genera Montastrea, Siderastrea and Diploria), sponges, octocoralsandalgaehaveallbeenaffected.Themostsevereincidences of bleaching were at Speyside (northeastern Tobago) where up to 90% coral bleaching was noted, with widespread mortality of the Caribbean barrelsponge,Xestospongiamuta,andextensivepartialmortalityofthe yellow tube sponge, Aplysina fistularis. A steady rise in sea surface temperatures (SSTs) to levels well beyond the bleaching threshold of corals for this region (29.5C), has been attributed as the cause to the widespread bleaching. The highest SSTs were noted at Speyside with temperatures up to 32 C. Similarly, in situ temperature recorders registered water temperatures exceeding bleaching thresholds on the reef, with the highest temperatures also recorded at Speyside (31 C). Investigations continue to assess the impact of the bleaching and recoveryofthereefs.However,climatechangepredictionssuggestthat widespreadmassbleachingeventsmaybecomemorefrequent,andthat greater resilience measures should be incorporated into reef managementpolicy.

Keywords Aplysina,Diploria,coralbleaching,Montastrea,seasurfacetemperature, Siderastrea,Tobago,Xestospongia

134

SpatialDistributionandextentofmangrovesinTrinidad. RahannaJuman1,DeaneshRamsewak2
1

EnvironmentalResearchProgramme,InstituteofMarineAffairsHilltopLane,Chaguaramas, Trinidad,WestIndies 2 GeomaticsUnit,InstituteofMarineAffairs,HilltopLane,Chaguaramas,Trinidad,West Indies,Email:rjuman@ima.gov.tt,dramsewak@ima.gov.tt

Abstract Mangroveforestsprovidearangeofprovisioning,regulating,culturaland supporting services, yet they are the most threatened ecosystem worldwide. In Trinidad, mangrove dominated wetlands were, and continuetobeimpactedbyreclamationforurbanization,industrialization and tourism in coastal areas. These wetlands are further threatened by humaninducedclimatechange.Thegoalofthisprojectwastoassessthe statusandtrendofmangroveforestsinTrinidadandestablishabaseline forresponsetoclimatechangeimpacts.Usinga2007baseline,thisstudy developed mangrove forest maps based on highresolution IKONOS imagery,GIStechnologyandextensivefieldsurveyswithGPS. Estimated mangrove cover was higher than predicted, perhaps because of inaccuracies in historical data and in some cases because of regrowth following past disturbances. In Trinidad, the majority of mangrove forests were found on the west coast. This coastline is occupied by more than 70% of the countrys population and has experienced the most intense development activities within the past three decades. The current mangrove coverage was estimated at 7,532 haonthewestcoastcomparedto1,132.8haontheeastcoast,481.3ha on the south coast and 0.3 ha on the north coast. The Caroni Swamp accountsfor56%ofallmangrovesinTrinidadandTobago. Whilemangroveforestswereimpactedbylandusechangesand erosion, there were instances where the forests have expanded at the expense of freshwater wetland communities. As landward migration continues,coastalsqueezefrombuiltdevelopmentonthelandwardedge of these wetlands can limit their movement. Sitespecific mangrove vulnerability assessment is recommended since the ecosystem is responding to other human threats besides climate change, and its responseisdeterminedbysitephysiography,hydrologyandecology. Keywords mangrove,climatechange,landuse,satelliteimagery,Tobago,Trinidad

135

Developingpublicawarenessandeducationtoolstopromotean understandingandappreciationofbiodiversityinthecoastaland marineenvironment LoriLeeLum


InformationCentre,InstituteofMarineAffairsHilltopLane,Chaguaramas,Trinidad,West Indies Email:lleelum@ima.gov.tt

Abstract

Among the general public there is a lack of awareness of the role and functions that coastal and marine biodiversity have on the health of oceans and humans. In order to promote a greater level of public understanding of marine biodiversity, the Institute of Marine Affairs (IMA) has been engaged in school outreach and education as well as publicawarenessprogrammes.Acombinationofmethodshasbeenused. The general public is reached mainly through IMA participation in environmentallythemed public exhibitions and through openaccess to itsspecializedlibrary.However,muchofthefocushasbeenonschools. TheMarineEducationCentreattheIMAhostsstudentsatboththe Primary and Secondary School level. In 2003, an outreach programme with four neighbouring schools was started. This programme was expanded in 2006, to schools in other areas throughout Trinidad and Tobago.Towidenthereachofthisprogramme,theteachingmodulesof theProgrammearebeingdevelopedintoaTeachingManualonCD. EngagementisbothattheschoolaswellasthroughvisitstotheIMA. Visits to the IMA often include mentoring with researchers at research laboratories. Several methods are used in presentations including multimedia, and the use of posters and brochures developed with local examplesofecosystems.Muchofthismaterialisdistributedtoschoolsas teachingmaterials.

Keywords awareness,education,exhibitions,marinebiodiversity,schools,Trinidad

136

Aninnovativeapproachformonitoringabioticfactorsinfluencing mangroveforestbiodiversityinanestuarineecosystem
1

M.Atwell1*,M.Wuddivira1,J.Gobin2andD.Robinson3

DepartmentofFoodProduction,UniversityoftheWestIndies,St.Augustine, 2 DepartmentofLifeSciences,UniversityoftheWestIndies,St.Augustine, 3 CentreforEcologyandHydrology,EnvironmentCentre,Wales,UK *Correspondence:emailmelissa.atwell@sta.uwi.edu

Abstract
Mangroveforestbiodiversityisdecliningworldwideduetoencroachment and other human impacts. These ecosystems are very important since they protect vulnerable coastal and marine ecosystems from rising sea levels and storm tides. In Small Island Developing States (SIDs) such as Trinidad and Tobago, mangrove forest biodiversity faces unprecedented degradationduetoanthropogeniclanduseandclimatevariability.With increasing urbanization and flow management via sluice gates, the hydrology and surface water flows in the Godineau swamp of Trinidad has been greatly altered. As a result, saline water has intruded into the swamp, altering soil salinity and changing the mangrove zonation patterns. This study investigates the water quality factors influencing mangrovezonationalongtheSouthOropoucheRiver,Godineauswamp. AerialphotoswereanalysedusingGIStodeterminemangroveboundary change. A novel geophysical approach (electromagnetic induction) was employed to assess apparent electrical conductivity (ECa) along two channels of the river (a 6 km forest and agricultural runoff dominated channel and a 2km wetland runoff dominated channel) bimonthly. A Horiba water quality checker U10 was used to determine the spatial distribution of water quality parameters along these two channels simultaneously with the ECa. Mangrove species were identified at 5m intervals along two sides of both channels. Results show that ECa was higherinthe2kmchannel(2711to3178mS/m)thaninthe6kmchannel (1776 to 2711 mS/m) while pH and DO levels were lower in the 2 km channel than in the 6 km channel. This suggests that salts accumulate while higher levels of decomposition take place in the more stagnant shorter channel. Red mangrove (Rhizophora mangle L), the hardiest of thethreespecies,wasdominantunderadversewaterqualityconditions in the 2 km channel and at the sea ward edge of the river. The red mangroveisindeedbettersuitedforwetlandrestorationefforts. Keywords electromagnetic induction, mangrove, Rhizophora mangle, SouthOropouche,Trinidad,zonation

137

Monitoringandmanagementofmarineinvasivealienspeciesin TrinidadandTobago. RosemarieKishore1,3,FrancisWeekes2andKhamaPhilip1

InstituteofMarineAffairs,P.O.Box3160,CarenagePostOffice,Carenage,Trinidadand Tobago. 2 MaritimeServicesDivision,MinistryofWorkandTransport,ANSAHouse,2ndFloor, CornerQueenandHenryStreets,PortofSpain,TrinidadandTobago. 3 CorrespondingAuthor:rkishore@ima.gov.tt

Abstract
Theintroductionofmarineinvasivealienspecies(IAS)hasbeenreported to be one of the greatest threats to marine biological diversity. In addition, marine IAS causes economic harm to marine industries and related infrastructure as well as impacts on human health. While the greenmussel,PernaviridisistheonlyknownmarineIASinTrinidad,little isknownofitsecologicalandeconomicimpacts.Additionally,nosystem exists to monitor the introduction of marine IAS. As part of a larger regionalIASproject,anecologicalassessmentofP.viridisisunderwayto determine its current distribution, the community structure of habitats associated with P. viridis, including pier pilings, mangrove prop roots, intertidal mud flats and artificial water channels. An economic assessment will focus on valuing the cost of P. viridis as a fouling organism since its introduction, as well as identify potential benefits. Findings from these assessments will inform efforts to mitigate the spreadofP.viridisinTrinidad.UndertheGloballastPartnershipProject, biological surveys for ports in Trinidad and Tobago are currently being plannedtoestablishedbaselineconditionsagainstwhichintroductionsof marine IAS via ships ballast water and associated sediments can be monitored.Biologicalsamplingstrategiesforbothprojectswillfollowthe CSIROCentreforResearchonMarineIntroducedPest(CRIMP)Protocol. These port biological baseline surveys will also assist in testing the efficacyofproposedlegal,policyandinstitutionalreformsrelatedtothe introductionofmarineIASfromballastwaterandassociatedsediments. AlreadydraftamendmentshavebeenmadetotheMarinePollutionBill 2008 and Shipping Bill 2008, which seeks to harmonise national legislations with that of the international Ballast Water Management Convention(2004). Keywords Perna viridis, port biological baseline survey, marine IAS, GloBallast

138

TheextentoftheseaturtlefisheryinTobago,WestIndies MichelleCazabonMannette
DepartmentofLifeSciences,TheUniversityoftheWestIndies,St.Augustine,Trinidad,West Indies.Email:mcazabon@gmail.com

Abstract
TrinidadandTobagocurrentlyprovidesonlyincompleteprotectiontosea turtles, through the Protection of Turtles and Turtle Eggs Regulations, 1975oftheFisheriesAct,(Ch.67:51).OnTobago,asmallartisanalfishery still exists, principally involving green (Chelonia mydas) and hawksbill (Eretmochelys imbricata) turtles. The current extent of this fishery is unknown but such information is necessary to inform the ongoing nationaldebateonthereviewoftheexistingRegulations. Thisstudysprimaryobjectivewastoestimatethenumberoffishers inTobagoengagedinthefishery,theirdistributionaroundtheisland,the income they derived from this activity, and the numbers of turtles harvested. The knowledge and opinions of fishers on sea turtle biology andlifehistory,andthelocalRegulations,wasalsoinvestigated. Two hundred and fifteen fishermen at 31 landing sites around Tobagowereinterviewedduring2007.Twentytwofishersat12landing sites confirmed that they targeted turtles, and fourteen considered turtles an important source of income. Twelve fishers reported turtle fishing year round while six reported limiting their fishing to the closed season.Onlyoneturtlefisherdemonstratedcompleteknowledgeofthe law,whileninehadincompleteknowledgeandelevendemonstratedno knowledge. The turtle fishery in Tobago remains active and widespread; however, relatively few fishers there obtained significant income from this activity. It is evident that enforcement of the current regulations is inadequate. This study supports the call for a moratorium on the turtle harvest, until the legislation can be revised to reflect the current understandingofturtlebiology,andtheFisheriesDivisionhasdeveloped thecapabilitiestoeffectivelyenforcethelegislation,closelymonitorany legalfisheryandtheimpactontheturtlepopulation. Keywords Cheloniamydas,Eretmochelysimbricata,fishery,harvest,turtle,Tobago

139

Mitigatingathreatofinvasivealienspeciesintheinsular CaribbeanATrinidadandTobagoPerspective. VeldaFergusonDewsbury


MinistryofFoodProduction,LandandMarineAffairs,St.ClairPortofSpain,Trinidad,West Indies.email:veldafergusondewsbury@yahoo.com

Abstract
This project aims to strengthen the regional efforts to address invasive alien species in the Caribbean. It sets out to accomplish this task by strengthening existing national capacity and management approaches, andbydevelopingCaribbeanwidecooperationframeworksonIAS.Such frameworks will facilitate the development of coordinated regional strategies for managing alien invasive species. The countries involved in the project are Bahamas, Dominican Republic, Jamaica, St. Lucia and TrinidadandTobago. Through this regional project, Trinidad and Tobago is engaged in three pilot projects, which include protecting the native biodiversity of the Nariva Swamp: protecting the local cocoa industry and germplasm from Frosty Pod Rot, and mitigating the spread of the marine invasive Perna viridis. These pilot projects strongly emphasise the need for capacitybuildingandincreasedawarenessofInvasiveAlienSpecies(IAS) issues among stakeholder groups and are designed to enable their findings and lessons to be readily transferred to other sites in the Caribbean. Keywords invasive alien species, Trinidad, Tobago, Perna viridis, frosty pod rot, NarivaSwamp

140

Lifeanddeathinthesavannasastudyoftherareterrestrial orchidCyrtopodiumparviflorum. HowardP.Nelson1,4,SharonLaurent,CarlysleMcMillanand EleanorDevenishNelson.


1

DepartmentofLifeSciences,TheUniversityoftheWestIndies,St.Augustine,Trinidad, WestIndies.Email:howien@hotmail.com 2 TrinidadandTobagoOrchidSociety,P.O.Box1128PortofSpain,Trinidad,WestIndies 38StAnnsRoad,PortofSpain,Trinidad,WestIndies 4 CorrespondingAuthor

Abstract
The terrestrial orchid Cyrtopodium parviflorum Lindl. (1843) is an extremely rangerestricted species in Trinidad, with extant populations only known from the Aripo Savannas. Although widely distributed from northernSouth America to Brazil, its limited distribution in Trinidad makes it a conservation concern locally. In the opensavanna habitat at Aripo,abioticconditionsaredominatedbyfloodedconditionsinthewet season and xeric conditions in the dry season, and the soil is extremely nutrient poor and acidic. One hundred and eighty nine individuals of C. parviflorum were marked and monitored over a 3year period (2008 2010) within Savanna 5 of the Aripo Savannas, and monthly data were collectedonfloweringfrequency,phenology,fruitset,andmortalityrates within the population. Over three years, average annual flowering rates forC.parviflorumwere16.37%,matureseedsetwas2.56%andnatural mortality was 2.5%. Anthropogenic disturbance, in the form of fire and illegalcollectionofplants,contributed32.8%tomortalityintheseplants over a oneyear period. This study suggests that C. parviflorum exhibits very low rates of reproductive effort, and points to an extremely K selectedlifehistorystrategybythisorchid.Thesedataalsoindicatethat persistent anthropogenic disturbance such as that observed during this studycouldleadtoadeterministicdeclineinthepopulationofthisorchid attheAripoSavannas. Keywords AripoSavannas,flowering,fruitset,phenology,mortality,Trinidad,

141

Index
abiotic ............................60,61,68,73,88,91,93,94,95,137,141 . agouti........................................................... 23,124,126,127,128 1 Annelida...........................................................................61,70,132 anthropogenicdisturbance.........................................................141 Aplysina.......................................................................................134 aragonite.............................................................................116,117 Araneidae....................................................................................133 AripoSavannas...............................99,100,101,105,106,108,141 Asclepiadoideae................................................................. 8,89,98 8 awareness........................................................................v,136,140 Bahamas......................................................................................140 Beachprofiles ...............................................................................42 . behaviour.............................................................15,17,26,28,131 benthicmacrofauna..................................................... 1,55,68,70 3 biofouling....................................................................................132 Biofouling....................................................................................132 biogeography...................................................................6,110,114 biomonitoring.............................................................................111 Biorock........................................................................................116 brucite.................................................................................116,117 Bryozoa.......................................................................................132 buoyantweighing ...............................................................116,122 . butterfly.............................30,32,33,34,35,36,37,38,39,40,41 Caligominor..................................................................................30 CaroniSwamp........................................8,9,10,11,12,13,16,135 Cebusapella...........................................................19,20,25,26,28 CentralRange.........................................20,123,124,125,126,127 Chaguaramas..............19,20,21,23,24,25,26,27,28,132,134, 135,136 Cheloniamydas.....................................................................54,139 climatechange.............................vii,68,70,71,122,133,134,135 cocoa............................................30,32,34,35,36,37,38,39,140 Cookstreeboa........................................8,9,10,11,12,13,14,16 coralbleaching............................................................................134 142

coralreef..............................................................................116,117 Corallusruschenbergerii..............................................................8,9 cracid............................................................................................ 31 1 Crustacea..................................................................................... 32 1 crustosemicrolichens.................................................................. 13 1 Cyrtopodiumparviflorum............................................................. 41 1 Dasyproctaleporina.....................................................123,124,129 Demography..................................................................................19 diet.............................................................................25,28,70,131 Diploria ........................................................................................ 34 . 1 directcurrent...............................................116,117,118,120,121 distancesampling........8,10,11,12,13,14,16,17,18,19,22,123 distribution..................6,14,16,21,31,37,41,53,54,55,56,60, 63, 67, 70, 88, 89, 91, 92, 93, 96, 97, 110, 111, 124, 133, 138, 139,141 disturbance..............30, 31, 32, 33, 34, 35, 36, 39, 40, 41, 42, 84, 99,110,141 diversity....................v,2,6,7,22,28,30,31,32,33,34,35,36,39, 40,41,55,63,66,68,72,73,74,76,77,78,79,81,82,84,85, 86,110,113,114,133,138 DominicanRepublic..................................................................... 40 1 edaphic...........................................................99,100,106,107,125 education..................................................................................... 36 1 electrolyticmineralaccretion...................................................... 16 1 electromagneticinduction........................................................... 37 1 endemic.................15,19,20,21,27,88,89,98,99,100,106,131 Eretmochelysimbricata...................................................53,54,139 Euptchiahermes............................................................................30 Euptchiapenelope....................................................................30,35 exhibitions ................................................................................... 36 . 1 fires................................99,100,101,102,103,105,106,107,108 fishery.......................................................................................... 39 1 flowering................................................3,73,97,99,105,106,141 forest....................7,17,20,21,22,25,28,30,32,34,35,36,37, 38, 40, 41, 72, 75, 81, 85, 86, 87, 95, 100, 102, 107, 110, 113, 114,124,125,127,128,129,135,137 frostypodrot............................................................................... 40 1 143

fruitset........................................................................................141 gamemammal....................................................................123,124 Geneticdiversity.....................................................................72,77 germplasm..................................................................................140 GloBallast....................................................................................138 Godineauswamp........................................................................137 grainsize..............................................55,59,60,63,64,65,67,68 GrandeRiviere.....................................32,33,35,36,38,39,42,53 groupsize..............................................................................25,131 habitat.............8, 9, 14, 15, 20, 22, 25, 28, 30, 31, 34, 35, 40, 42, 68,70,87,88,91,95,96,97,100,106,131,133,141 habitatclassification...................................................................133 habitatpreference................................................................42,106 harvest..........................................................................20,124,139 hunting...............................19,20,99,101,105,124,127,128,131 indicator.................................................................30,33,38,40,96 intertidal.....................55,56,59,62,63,64,65,66,67,68,70,138 introducedspecies................................................................19,132 invasivealienspecies..........................................................138,140 Invertebrate....................................................................................2 Jamaica..................................................................................85,140 Kingsestimator..........................................................123,126,127 landuse.................................................................................32,135 lichens..................................................................110,111,112,114 linetransect...............................8,11,12,17,18,91,123,125,126 macrolichen ................................................................................110 . managementplan.......................................... 9,100,101,105,108 9 mangrove...............................8,9,10,11,14,58,85,135,137,138 marinebiodiversity...............................................................69,136 Maxent.......................................................88,91,92,94,95,96,98 Megabalanuszebra ....................................................................132 . Metastelmafreemani........................................................ 8,89,92 8 microlichens................................................................................110 Milleporaalcicornis.....................................................116,119,121 Mollusca......................................................................................132 Montastrea.........................................................................121,134 Moraexcelsa................................................................ 2,73,79,86 7 144

mortality....................................17,73,81,100,105,106,134,141 NarivaSwamp..................................................................20,28,140 Nephilidae.................................................................................... 33 1 Nereis...........................................................................55,64,66,67 Opensavannas...............................................................................99 orbweavingspider...................................................................... 33 1 orchid........................................................................................... 41 1 particlesize........................................................................42,68,70 percentagepolymorphism.................................................72,76,77 Pernaviridis.................................................................132,138,140 pH...................................55,58,60,63,68,75,88,91,93,120,137 phenology....................................................................108,131,141 Pipilepipile................................................................................... 31 1 pipingguan.................................................................................. 31 1 populationdensity.................................8,10,11,12,14,16,19,21 populationestimate ..............................................................15,123 . Porifera..................................................................................62,132 Rhizophoramangle................................................................10,137 salinity..................................................................55,58,60,65,137 sandybeaches............................................55,56,57,61,66,69,70 satelliteimagery.......................................................................... 35 1 schools......................................................................................... 36 1 seaturtles..................................................................42,53,54,139 seasurfacetemperature............................................................. 34 1 sedimentcomposition...................................................................42 Shannondiversity..........................................................................81 Siderastrea................................................................................... 34 1 SouthOropouche......................................................................... 37 1 speciescomposition ..............................99,100,102,104,107,133 . speciesdiversity.............................................................65,102,133 speciesrichness...................2,31,34,36,37,40,89,113,115,133 squatting........................................................................................99 St.Lucia........................................................................................ 40 1 Tetragnathidae ............................................................................ 33 . 1 tropicalcoastlines....................................................................55,56 tropicaltree.............................................................................72,86 Tuftedcapuchin.............................................................................19 145

turtle................................................................................42,53,139 TurtleBeach..................................................................................42 unbiasedheterozygosity.........................................................72,76 Venezuela.....................................................9,28,72,73,74,75,79 vertebrate............................................................................ ,4,105 2 voucherspecimens.....................................................................110 Xestospongia...............................................................................134 zonation.......................................55,56,59,64,66,67,68,70,137

146

You might also like