Comparison of Ceriops Pseudodecanda Sp. Nov
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Botanical Studies (2010) 51: 237-248. SYSTEMATICS Comparison of Ceriops pseudodecanda sp. nov. (Rhizophoraceae), a new mangrove species in Australasia, with related species Chiou-RongSHEUE1,2,*,Ho-YihLIU3,Chi-ChuTSAI4,andYuen-PoYANG3,5 1Department of Biological Resources, National Chiayi University, 300 Syuefu Rd., Chiayi 600, Taiwan 2 Department of Life Sciences, National Chung Hsing University, 250, Kuo Kuang Rd., Taichung 402, Taiwan 3Department of Biological Sciences, National Sun Yat-sen University, 70 Lien-hai Rd., Kaohsiung 804, Taiwan 4Kaohsiung District Agricultural Improvement Station, 2-6 Dehe Rd., Changihih Township, Pingtung County 908, Taiwan 5Department of Bioresources, Dayeh University, 168 University Rd., Dacun, Changhua 515, Taiwan (ReceivedAugust28,2008;AcceptedSeptember24,2009) ABSTRACT. Thenotionthatthewidespreadmangrove genus Ceriops consistsofthreespecies,C. austra- lis (White)Ballment,Smith&Stoddart,C. decandra(Griff.)DingHouand C. tagal(Perr.)C.B.Rob.,is stillwidelyaccepted.However,ourrecentstudieshaveshownthatthepreviouslyrecognizedspeciesCeriops decandra canbeseparatedintothreespecies,C. decandra,C. zippelianaBlumeandCeriops pseudodecandra sp.nov. Sheue,Liu,Tsai&Yangbasedonmorphologicalandmolecularevidence.Thelastentityisnewly describedinthistreatment,whichhasbeenmisappliedwiththenameC. decandra forseveraldecadesinAus- tralia,PapuaNewGuineaandSeram,NewGuinea(IrianJaya)ofIndonesia.Morphologically,thenewspecies ismoresimilarto C. decandraandC. zippelianathantoC. australis andC. tagal.Furthermore,molecular phylogeneticanalysisalsosupportsthenotionthatC. pseudodecandra isdistinctlyanewspecies.Here,the botanicaldescriptionswithanillustrationandthedistributionrangesoftherelatedspeciesareprovided.Akey todifferentiatespeciesofCeriopsandacomparisonwithanemphasisonthesethreemorphologicallysimilar speciesaregivenaswell. Keywords:Australia;Ceriops decandra;Ceriops pseudodecandra;Ceriops zippeliana;Indonesia;Mangroves; PapuaNewGuinea. INTRODUCTION (Perr.)C.B.Rob.andC. decandra (Griff.)DingHou. Ceriops tagalisacommonandfamiliarconstituentof Mangroves are the characteristic intertidal plant mangroveswithawidedistributionrange,fromEast formations found along the sheltered tropical and AfricathroughoutMalaysiatoMicronesia,Melanesia subtropicalcoastlines.Themangrovefloraconsistsof and northernAustralia (Hou, 1958; Duke, 2006). around36generain26families(Saenger,2002).Among Ceriops decandraisthoughttorangefromIndiathrough themembers,thepan-tropicalfamilyRhizophoraceae, Thailand,VietnamandtheMalayPeninsula,Java,Borneo, whichcomprises16generaandabout120speciesof PhilippinestoNewGuinea(Hou,1958;Tomlinson,1986; evergreentreesandshrubs(Hou,1958),istherichest Field,1995;NaskarandMandal,1999;Duke,2006). mangrovefamilywithfourexclusivelymangrovegenera. Ceriops australis(White)Ballment,Smith&Stoddart However,adetailedmorphologicalandanatomicalstudy (1988)wasaddedasthethirdmemberofthegenuswhich ofthemangroveRhizophoraceaeshowedthattwospecies wasoriginallyrecognizedasavarietyofC. tagalbyWhite belongedtoKandeliaandCeriops,separately,andshould (1926).Theevidenceforitselevationtospecieslevelwas beaddedordescribed(Sheue,2003).ThespeciesKandelia basedonisozymecharactersaswellasthemorphological obovataSheue,LiuandYongwaslaterreportedasanew featureofhypocotyl(Ballmentetal.,1988).Arecent speciesfromEastAsia(Sheueetal.,2003)andreceiveda evaluationconcerningthetaxonomicstatusofthespecies strongsupportbythestudyusingmicrosatellitemarkers basedonmorphologicalandmolecularevidence(Sheueet (Giangetal.,2006). al.,2009a)supportsthistreatment. Thelastrevisionofthegenus Ceriopswasputforward Aspreviouslymentioned,anotheruncertainspecies byHou(1958),withtwospeciesrecognized:C. tagal wasnoticedinthegenusCeriopsArn.(Sheue,2003). ThisuncertainspeciesofCeriops hasbeenrevertedtoits *Correspondingauthor:E-mail:[email protected];Tel: originalnameC. zippelianafromitscurrentappliedname +886-5-22857395. 238 Botanical Studies, Vol. 51, 2010 C. decandra(Sheueetal.,2009b).Ceriops zippeliana tagalandC. australiswereincludedasoutgroupsfor ismorphologically andgeneticallydiscerniblefrom comparison(Table1).Threetofiveleavesweretaken C. decandrawhichoccursfromIndia,Burmaandthe fromeachindividualandstoredwithsilicagelinzip-lock westcoastofThailand;andC. zippelianashouldbethe plasticbagsuntilDNAisolation.Voucherspecimenswere correctnameforthepopulationsinsoutheasternAsiato depositedattheHerbariumofNationalChiayiUniversity whichthenameC. decandraismisapplied(Sheueetal., (CHIA). 2009b).Moreover,afurtheranalysisonthepopulations DNA extraction, PCR amplification and electrophoresis, ofthecurrent‘C. decandra’inNewGuineaandAustralia DNA recovery and sequencing.Thedetailedprocedures revealsthattheyaremorphologicallydifferentfromC. ofDNAextraction,conditionsofPCRandsequencingfor zippelianan andC. decandra.Thisraisesthequestion thisstudyweresimilartothosedescribedintheprevious aboutthetaxonomicstatusofthepopulationsandattracts report(Sheueetal.,2009a)andtheuniversalprimersfor ourinteresttoclarifytheirspeciesidentity. amplifyingthetrnLintronofchloroplastDNAwerethe Afteradetailedstudyusingherbariumspecimens, sameasdescribedbyTaberletetal.(1991). freshmaterials,andvisitstothehabitatsinAustraliaand Data analyses. DNA sequence alignment was Indonesiaoverfouryears,theauthorshavecometothe conductedusingtheprogramClustalWmultiplealignment conclusionthattheso-called‘C. decandra’ofAustralia, inBioEdit(Hall,1999).Geneticrelationshipswerethen NewGuineaandSeramactuallyisyetanundescribed determinedusingtheprogramMEGAversion4(Tamura speciesbasedonmorphologicalcharacters,palynological etal.,2007).Thegeneticdistancematrixwascalculatedby evidenceandmolecularevidenceobtainedinthisstudy. thetwo-parametermethodofKimura(1980)andthenused Here,thebotanicaldescriptionsandthedistributionranges toconstructthephylogenetictreesusingtheNeighbor ofthisnewspeciesandrelatedspeciesarepresented.Akey joining(NJ)method(SaitouandNei,1987).Maximum to distinguish the five taxa of Ceriopsisprovidedandthe parsimony(MP)analyses(Fitch,1971)weredoneusing interspecific morphological differences are discussed, with code modified from the Close-Neighbor-Interchange (CNI) anemphasisonthethreemorphologicallysimilarspecies algorithm(RzhetskyandNei,1992)inMEGAversion4 ofC. decandra,C. pseudodecandra and C. zippeliana. (Tamuraetal.,2007).Bootstrapping(1000replicates)was carriedouttoestimatethesupportforbothNJandMP MATERIALS AND METHODS topologies(Felsenstein,1985;HillisandBull,1993).The strictconsensusparsimonioustreewasthenconstructed Morphological and pollen characteristics usingtheprogramMEGAversion4(Tamuraetal.,2007). Freshsamplesandherbariummaterialswereexamined andphotographed.Threetotenbrancheswithflowers TAXONOMIC TREATMENT and viviparous seedlings were collected from each of five individualsofC. pseudodecandraalongtheriverbank Ceriops pseudodecandra Sheue,Liu,Tsai,andYang,sp. ofBlackmoreRiveratDarwinofNorthernTerritory, nov.—TYPE:Australia:DarwinHarbour:onbanksof CairnsofQueensland(October,2005)andCardwellof tidal Blackmore River (12°3′54.4″ S, 130°57′56.9″ E), Queensland in Australia (May, 2007). In addition, a field C.-R. Sheue M234,26thOctober2005(holotype:DNA; surveyfordistributionrangeofCeriopswasespecially isotypes:BM,BO,DNA,GH,HAST,K,L,MOand carriedoutinAmbon,Seram,Sulawesi,BaliandLombok SING).Figures1-3andTable2 ofIndonesia(July-August,2009).Voucherspecimens Ceriops decandraauct. non(Griff.)DingHou:Hou,Flora weredepositedintheHerbariumofNationalChiayi Malesiana,ser.1,vol.5:471,1958.pro part. University(CHIA).Thespecimensexaminedwerefrom Folia pseudodecandra vel obovato-elliptica, 6-12 cm herbariaBM,BO,CAL,DNA,GH,HAST,IBSC,K,L, longa, 2.5-5 cm lata, venis lateralibus primariis 8-12, MO,PPI,SING,SINUandTAI(HolmgrenandHolmgren, petiolis 1.2-3.5 cm longis. Pedunculis brevissimis. Petala 1998). 5, multi-fida, lobis apice 12-18 laciniatis, cilia in dimidio Pollen grains were smeared on the surface of a inferiore. microscopeslideandexaminedwithanOlympusBH-2 Smalltreesupto6mtall;barkgrayishbrownwith Light Microscope first to measure the size (N=30). Also, horizontalfissuresofmainstem,flakyandflanged pollengrainswereairdriedandscatteredonthesurface buttressesatbase.Leavesoblongtoelliptic-obovate, ofthestubcoveredwithdouble-sidetapeandcoated 6-12(-13)cm×2.5-5(-6)cm,apexobtuse,roundedto withgolddirectly.Thesesampleswereexaminedand emarginate,baseattenuate,lateralveins8-12pairs;petiole photographedwithaHitachS-2400ScanningElectron 1.2-3.5cmlong.Stipules2-3cmlongatextensionstage Microscope. beforedropping,with8-12layered,80-100colletersinside atadaxialbase.Inflorescencecompactbifurcatecyme- Molecular evidence like,usuallyborneattheuppernodesofabranch,axillary, Materials.SamplesofCeriops decandra, C. zippeliana (2-)3-20 flowered. Bracteole 2-lobed, disc-like, sessile, 2 and C. pseudodecandra were collected at different mm long, apex rounded with small fissures. Calyx lobes localitiesofIndia,SingaporeandAustralia.BothC. 5, erect while flowering and fruiting, ovate, acute, 3 mm SHEUE et al. — Ceriops pseudodecandra sp. nov., a new mangrove species 239 Table 1.Alistofmolecularstudyforthe20specimensofCeriops decandra (Griff.)DingHou, C. pseudodecandraSheue,Liu,Tsai andYang,C. zippeliana Blumeandtwooutgroupspeciesof thisgenus,andtheirdifferentgeographicaldistributionsandaccession numbers. No. Taxon Locations Accessionno. Rh-26 C. decandra Pichavarum,India(IN) EF118952 Rh-28 C. decandra WestSundarbans,India(IN) EF118953 Rh-29 C. decandra WestSundarbans,India(IN) EF118954 Rh-30 C. decandra WestSundarbans,India(IN) EF118955