全 文 ：广 西 植 物 Guihaia Aug．２０１５,３５(４):４４７－４６３ http://journal．gxzw．gxib．cn
DOI:１０．１１９３１/guihaia．gxzw２０１５０６０１５
唐光大,张国强,洪文君,等．基于ITS和matK序列的兰科沼兰族分子系统研究及二新种[J]．广西植物,２０１５,３５(４):４４７－４６３
TangGD,ZhangGQ,HongWJ,etal．PhylogeneticanalysisofMalaxideae(Orchidaceae:Epidendroideae):twonewspeciesbasedonthecombined
nrDNAITSandchloroplastmatKsequences[J]．Guihaia,２０１５,３５(４):４４７－４６３
PhylogeneticanalysisofMalaxideae(Orchidaceae:
Epidendroideae):twonewspeciesbasedonthecombined
nrDNAITSandchloroplastmatKsequences
TANGGuangＧDa１,２,ZHANGGuoＧQiang２,HONGWenＧJun１,
LIUZhongＧJian１,２,ZHUANGXueＧYing１∗
(１．CollegeofForestryandLandscapeArchitecture,SouthChinaAgriculturalUniversity,Guangzhou５１０６４２,China;
２．ShenzhenKeyLaboratoryforOrchidConservationandUtilization,TheNationalOrchidConservationCenter
ofChinaandTheOrchidConservationandResearchCenterofShenzhen,Shenzhen５１８１１４,China)
Abstract:Malaxideaeisalargercosmopolitanorchidtribeconsistingofnearly２０００speciesandisdistributedworldＧ
wide,excludingthepolaranddesertregions．Itisabundantinthetropicalregions,particularlyinAustralia,andthetropiＧ
calAsia,SoutheastAsia,thetropicalAmericasandAfrica．Molecularandmorphologicalanalyseshavebeenperformedto
establishthephylogeneticrelationshipswithinthistribe．However,thesubtribeandintergenericrelationshipsremainunＧ
clear,andthegenusdefinitionofthistribeisconsiderablycontroversial．Themaximumparsimony,maximumlikelihood
andBayesianinferenceanalyseswereperformedbasedonthecombinedsequencesofthenuclearITSandchloroplast
matKgenesof１３３taxa(１０otheraliedspeciesasoutgroups),whichrepresentnearlyalofthemajorgeneraoftheMalＧ
axideae．TheseresultssuggestedthatMalaxideaecouldbedividedintothreeclades,theepiphyticOberoninae,theterresＧ
trialLiparidinae,andaterrestrialCrepidiumclade．TheOberoninaeandLiparidinaecouldbedividedintosixgeneraand
fivegenerarespectively,andCrepidiumcladecamposesoffourgenera．ThesubtribeYpsilorchidinaeshouldbeatributed
toOberoninae．DisticholiparisshouldberemovedbecauseitsharedthesametypewithStichorkis．ThephylogeneticreＧ
lationshipofCrepidiumandDieniaismonophyletic,althoughtheyhavediferentlipstructures．Inaddition,twonew
speciesfoundinSouthwestChinaandnorthernVietnam,PlatystyliparismalipoensisG．D．Tang,X．Y．Zhuang&Z．J．
LiuandCestichispingtaoiG．D．Tang,X．Y．Zhuang&Z．J．Liu,weredescribedbasedonthemolecularanalysisand
morphologicalobservation．
Keywords:Malaxideae;nrDNAITS;matK;phylogeny;Paraphyletic
CLCnumber:Q９４３．２ Documentcode:A ArticleID:１０００Ｇ３１４２(２０１５)０４Ｇ０４４７Ｇ１７
基于ITS和matK序列的兰科沼兰族分子系统研究及二新种
唐光大１,２,张国强２,洪文君１,刘仲健１,２,庄雪影１∗
(１．华南农业大学 林学与风景园林学院,广州５１０６４２;２．深圳市兰科植物保护与利用重点实验室,
国家兰科植物保护中心,深圳市兰科植物保护研究中心,广东 深圳５１８１１４)
摘 要:沼兰族是兰科植物的大族之一,约２０００种,除了极地和沙漠地区,全球均有分布.该族植物主要分
收稿日期:２０１５Ｇ０６Ｇ１３ 修回日期:２０１５Ｇ０７Ｇ０６
基金项目:国家林业发展计划项目(２０１１Ｇ１００);广东省自然科学基金(２０１４A０３０３１０４６５);深圳市科技计划项目 (JC２０１００５３１０６８９A).
作者简介:唐光大(１９８２Ｇ),男,贵州普安人,博士,讲师,研究方向为植物系统与进化,(EＧmail)gdtang＠scau．edu．cn.
∗通讯作者:庄雪影,博士,教授,研究方向为植物系统与进化,(EＧmail)xyzhuang＠scau．edu．cn.
布在热带地区,尤其在东南亚、热带美洲、非洲以及澳大利亚等地区种类非常丰富.目前,已有关于该族植物
形态和分子系统的研究,但有关该族亚族和属间的系统关系尚不清楚,属的界定争议也较大.该文基于核基
因片段ITS和叶绿体基因片段matK序列,采用最大简约法、最大似然法贝叶斯推理分析法,对现有沼兰族主
要属的１２３种植物和１０个外类群植物进行了分子系统学研究.结果表明:沼兰族主要分为３个亚族分支,包
括附生的鸢尾兰亚族(Oberoninae)、地生的羊耳蒜亚族(Liparidinae)和沼兰亚族分支(Crepidiumclade).鸢
尾兰亚族包括６个属、羊耳蒜亚族分支包括５个属、沼兰亚族分支包括４个属;丫瓣兰亚族(Ypsilorchidinae)
应归并为鸢尾兰亚族;Disticholiparis属与Stichorkis属的模式标本相同,应并入Stichorkis属;沼兰属(CreＧ
pidium)和无耳沼兰属(Dienia)的唇瓣结构差异较大,但二者均为单系类群.此外,在收集野外实验材料过程
中,发现了２种产自中国西南部和越南北部的沼兰族新种,分别命名为麻栗坡羊耳蒜(PlatystyliparismaliＧ
poensisG．D．Tang,X．Y．Zhuang&Z．J．Liu)和秉滔羊耳蒜(CestichispingtaoiG．D．Tang,X．Y．Zhuang&
Z．J．Liu).
关键词:沼兰族;核基因ITS;matK;系统学;并系类群
Malaxideaeisoneoftheoldestorchidtribesand
encompassesnearly２０００speciesthatarenativeto
bothtemperateandtropicalareasworldwide(Chenet
al．,２００９)．Lindley(１８２６)firstclassifiedandrevised
２１generainto Malaxideae．SixofthesegenerareＧ
mainedinthistribe,includingMalaxis,Microstylis
(attributedtoCrepidiumnow),Liparis,Dienia,EmＧ
pusa,andPedilea,butother１５generawereattributed
toothertribes．Lindley(１８３０)redefinedthistribeand
emphasizedthatthe mostimportantfeatureswere:
polencoheringinmassesofafirmwaxytexture,withＧ
outanyofthecelularsubstance,andremainingunder
theformofadistinctglandlyinguponthestigma．
Pfitzer(１８８７)redefinedthistribeagainas“plantwith
leavesduplicative,stem gracileorincrassateleaved
pseudobulb;leaflamina exceptforCestichis and
Oberonia,withcontinentbasalsheath;petalsandlaＧ
belummoreconspicuousthanexternalsepals,column
apodalorformingashortspurwiththelabelum,anＧ
thererectorincumbent,polinia４,waxy,glabrous;
dispersed”．Schlechter(１９１１)revisedtheLiparidinae
(Malaxideae)ofNewGuineaandleftMalaxis,MicroＧ
stylis,Orestias,Liparis,Cestichis,OberoniaandHipＧ
peophylluminthissubtribe．Dressler(１９８１)emphaＧ
sizedthatthemaincharactersofMalaxideaewerenot
onlythenakedpolinia,butacolumnlackingafootat
thegynostemiumbase,andstatedthatpolinialacking
adistinctcaudiclewerealsoequalcharactersfordistinＧ
guishingthistribe．Szlachetko(１９９５)recognized１６
generainthesubtribeMalaxidinae,sevenwerenewly
describedbyhim．HedividedMalaxideaeintotwosubＧ
tribesofterrestrialMalaxidinaeandOberoninae,but
thelatterwasnamedbyAveryanov(１９９１)．Szlachetko
andMargońska(２００２)provideddetaileddrawingsof
thecolumnstructureofrepresentativesof１５genera．
Pridgeonetal．(２００５)stated１４generainthetribeof
Malaxideae．However,thedescriptionof Malaxideae
wasquiteaugmented,especialyintermsofleafstrucＧ
ture,floralarrangementwithintheinflorescences,tepal
formandovary．
MalaxideaeisalsooneofthefewgroupsoforＧ
chidsthatcontainlargenumbersofbothobligateterＧ
restrialandepiphyticspecies(Atwood,１９８６)．There
areobviousdiferencesbetweenterrestrialandepiphytＧ
ictaxainvegetativeorganscharacters．Anexampleis
thattheterrestrialgenera,suchasCrepidium and
Diteilis,havesignificantlyplicateleaves,buttheepiＧ
phyticgeneraareconduplicateexceptforOberoniaand
afewLiparisspecies．However,theirflowermorpholＧ
ogiesareratherhomogenous,whicharetypicalyvery
smal,bornonaterminalinflorescence,containinga
single,terminal,incumbentanther,usualywithfour
nakedpolinia (Cameron,２００５),onlyafewspecies
havetwopolinia(Lietal．,２０１３)．Manyspeciesof
Malaxideaearepolinatedbyfungusgnats,andonlya
fewarepolinatedbyantsandichneumonflies(Tang
etal．unpublisheddata)．
Inrecentyears,thereareconsiderablecontinuing
controversyaboutthesizeofgenerawithintheMalＧ
axideaeduetotheirwidedistributionanddiversemorＧ
phology,andthemaindebatesfocusonthegenericdefＧ
initionofthistribe．InthestrictestandmostconservaＧ
８４４ 广 西 植 物 ３５卷
tivesenses,somescholarsconsideredthatthetribe
shouldbedividedintoatleastthreegenera:Oberonia,
Liparis,andMalaxis,butstil manygenerawereesＧ
tablishedorseparated(Cameron,２００５)．Forexample,
Dressler(１９９３)recordedthatMalaxideaeconsistedof
sixgenera:Hippeophyllum,Liparis,Malaxis,OresＧ
tias,OberoniaandRisleya,butnowRisleyainColaＧ
bieae(Xiangetal．,２０１４)．Szlachetko(１９９５)suggesＧ
tedthereweremorethan１８generainthistribe．FurＧ
thermore,MargońＧskaandSzlachetkodescribedsome
newgenerasuchasAlatiliparis (Margońskaetal．,
２００１),Disticholiparis(Margońskaetal．,２００４),SeiＧ
denforchis (Margońska,２００６),Platystyliparis
(Margońska,２００７),andCrossoliparis (Margońska,
２００９)．Margońskaetal．(２０１２)describedthat２５genＧ
erabelongedtothistribeanddividedthemintothree
subtribes:Oberoninae(containingtwogenera),MalＧ
axidinae(containing１２genera),andLiparidinae(conＧ
taining１１genera)．
Althoughsome molecularphylogeneticstudies
haveexaminedthephylogeneticandevolutionaryrelaＧ
tionshipsofMalaxideae(Cameron,２００５;Margońskaet
al．,２０１２),someresultsindicatedthatLiparisand
Malaxiswerepolyphyleticaliances．Cameron(２００５)
describedandidentifiedfour habitＧleafＧmorphology
syndromesasindicativeofphylogeneticrelationshipsin
Malaxideae,suchasepiphyteswithseveralelongate
conduplicateleaves(Liparis)andwithunifacial,laterＧ
alycompressed,equitantleaves(Oberonia),terrestrial
withtworoundedconduplicateleaves(Malaxis)and
terrestrialwithplicateleaves(Crepidium)．However,
thegenericdefinitionofMalaxideaeremainsunclear．
Morecomprehensivestudiesareneededtoexplorethe
exactevolutionaryrelationshipsofMalaxideaeatthe
genericlevel．MalaxideaeshouldbereＧclassified,espeＧ
cialygiventheaggregatedstudyresults,combined
withthemolecularphylogeneticsandtheirhabits,vegＧ
etation,andmorphologies．
Inthisstudy,thenewsequencesofnuclearITS
andchloroplastmatKfrom３５specieslocatedinChina
werecombinedwiththesequencesquotedfromGenＧ
Bank,andassessedforthephylogeneticrelationshipsaＧ
mongMalaxideae,incombinationwiththeplanthabＧ
its,pseudobulbsandleaves．Inaddition,weprovidea
supportedphylogeneticresolutionforMalaxideaeand
comprehensiveresultsregardingthistribe．
１ MaterialsandMethods
１．１Materials
Atotalof１２１speciesofMalaxideaewereanaＧ
lyzed,consistingofmaingeneraofMalaxideae,incluＧ
dingLiparis,Oberonia,Malaxis,Crepidium,Dienia,
Empusa,Oberonioides,Stichorkis,Cestichis,and
Platystyliparis．Tendiferentaliedspecies,AcantheＧ
phippium mantinianum,AncistrochilusrothschildiaＧ
nus,Bulbophyllumodoratissimum,CalantheargenＧ
teostriata,C．sinica,Collabiumsimplex,Dendrobium
crumenatum,Listerasmalli,Phaiustancarvilleae,
andRidleyellapaniculata,wereselectedasoutgroups
accordingtopreviousstudies(VandenBergetal．
２００５)．Thereare３５sequencesofspeciesweregeneraＧ
tednewforthisstudyfromYunnan,Guangdongetc,
and９８sequencesofspeciesweredownloadedfromexＧ
istingsequencesinGenBank．
１．２Amplificationandsequencing
TotalDNAwasextractedfromfreshmaterialor
silicagelＧdriedplanttissueusinga MultisourceGeＧ
nomicDNA MiniprepKit(AxygenBiosciences)acＧ
cordingtothemanufacturer’sinstructions．TheampliＧ
ficationreactionincludedtotalDNA,primers,ExＧTaq
buferandExＧTaq DNApolymerase (Takara Bio)．
Thepolymerasechainreaction(PCR)profileconsisted
ofaninitial５minpreＧmeltingstageat９５℃,folowed
by３０cyclesof３０sat９５℃ (denaturation),３０sat５０
℃to５５ ℃ (annealingtemperaturewasdetermined
basedontheprimerrequirement),１minto３minat７２
℃ (extensiontimewasdeterminedbasedonthelength
ofthetargetDNAregion)andafinal１０minextension
at７２℃．
AmplificationoftheITSregionwasperformeduＧ
singtheprimerpairsITSAandITSB(Mikeetal．,
１９９９)．FormatKsequences,amplification wasperＧ
formedusingtheprimermatKＧ７３１F (Liuetal．,
２０１１)．Table１containsthedetailedinformation．
ThePCRproductswererunon１．５％agarosegels
９４４５期 唐光大等:基于ITS和matK序列的兰科沼兰族分子系统研究及二新种
Table１ Primersusedinthisstudy
Primer Sequence(５′to３′) Origin
ITSA GGAAGGAGAAGTCGTAACAAGG Mikeetal．,
１９９９
ITSB CTTTTCCTCCGCTTATTGATATG Mikeetal．,
１９９９
matKＧ７３１F AAGAAAAGATTCTTTTGGTTCC Liuetal．,
２０１１
toassessthequalityoftheamplifiedDNA．Thegels
withthetargetproductswereexcisedpurifiedusing
DNAGelExtractionKits(AxygenBiosciences)and
thensequencedbyLifeTechnologiesCorporation．
１．３Sequenceeditingandassembling
Bothforwardandreversesequences,aswelas
electropherograms,wereeditedandassembledusing
DNASTAR (http://www．dnastar．com/)．DNAseＧ
quenceswerealignedtothemusclemodelandmanualＧ
lyadjustedusingMEGA５．０５(Tamuraetal．,２０１１)．
ThealignedsequencesareavailablefromthecorreＧ
spondingauthorsuponrequest．
１．４Dataanalyses
ThedatasetsofanuclearITS,plastidDNAmatK
andtheircombinationwereusedforthesubtribeand
genuslevelanalysis．Insertions,deletionsandsomeunＧ
availablesequencesweretreatedasmissing．PhylogeＧ
neticanalyseswereperformedusingBayesianinference
(BI),maximumparsimony(MP)andmaximumlikeliＧ
hoodanalyses．ThebestＧfitmodelforeachdatasetwas
selectedusingModeltest３．７undertheAkaikeInforＧ
mationCriterion(Posadaetal．,１９９８)．ThehomogeＧ
neitiesbetweennrDNAITSdata,andmatKdataset
weretestedusingtheincongruencelengthdiference
(ILD)test(Farrisetal．,１９９５),asimplementedin
PAUP∗ version４．０b１０ (Swoford,２００２)．TBR
branchwasswappingandkeepingnomorethan１００
treesperrandomadditionalreplicate．AspreviouslydeＧ
scribedbyCunningham (１９９７),asignificancelevelof
P＝０．０１wasadoptedforthistest．
MPanalyseswereperformedusingthePAUP∗
version４．０b１０(Swoford,２００２)．AlcharacterswereeＧ
qualyweighedandunordered．Thesettingsincluded
１０００randomadditionalsequencesandaheuristicsearch
withtreebisectionＧreconnectionbranchswapping．Table
２listthetreelength(TL),consistencyindex(CI)and
retentionindex(RI)．MLanalysiswasperformedusing
RA×MLwith１００bootstrapreplicatesandsettingsas
describedin(Stamatakisetal．,２００８)．BIanalysiswas
performedusingMrBayes３．１．２(Ronquistetal．,２００３)．
Thefolowingsettingswereapplied:samplingfrequency
＝１０００,temp＝０．１,burnＧin＝１００００andnumberof
Markovchain MonteCarlogenerations＝４０００００００．
Thefirst１００００treeswerediscardedasburnＧintoenＧ
surethatthechainsreachedstationarity．AmajorityＧrule
consensustreewasconstructedonthesetreessampled
aftergeneration４０００００００．
Table２ Parsimonystatisticsfromphylogenetic
analysesofthevariousdatasets
Data Taxa Alignedlength
Information
site TL CI RI
ITS １２１ ８４７ ４７５(５６．１％) ２９４６ ０．３７７８０．８０４５
matK １２０ １４０５ ３０８(２１．９％) １０３０ ０．６２１４０．８６４１
Combined １３９ ２２５２ ７８３(３４．８％) ４１０７ ０．４３６４０．８１５１
２ ResultsandAnalysis
ITSanalysissupportedthatMalaxideaecouldbe
dividedintothreeclades,includingtwoterrestrial
cladesandoneepiphyticclade(PP＝１００％,BPMP＝
９４％,BPML＝９９％,Fig．１)．Thefirstterrestrialclade
includedfivegenera:Malaxis (containing１０taxa),
Liparis(１２taxa),Oberonioides (２taxa),andtwo
unnamedgenera(CladeB,C;Fig．１)．ThesecondterＧ
restrialcladeincludedfivegenera:Crepidium (１５taxＧ
a),Dienia(４taxa),Diteilis(７taxa),Empusa(４taxＧ
a)andanunnamedgenus(CladeD;Fig．１)．Among
theseclades,LitseaauriculatawasattributedtoMalＧ
axis,L．maingayiandL．purpureoviridisaresisters
toOberonioides．Thethirdepiphyticcladeconsistedof
sixgenera:Oberonia (１５taxa),Platystyliparis (７
taxa),Cestichis (１２taxa),Stichorkis (５taxa),and
twounnamedgenera(CladeE,F;Fig．１)．Oberonia
wasmonophyleticinthistreeandasisterofLipariss．
l．(Fig．１)．
TheresultsoftheBI,MPandMLanalysisofmatK
alsoshowedthatMalaxideaecouldbedividedintothree
clades(PP＝１００％,BPMP＝９６％,BPML＝９７％,Fig．
２)．However,theposteriorprobabilitiesofthematK
０５４ 广 西 植 物 ３５卷
Fig．１ PhylogramobtainedfromBayesianinferenceanalysisofthenrDNAITSdataofMalaxideae Numbersatthenodesare
bootstrappercentagesandBayesianposteriorprobabilities,respectively．“－”indicatesthatthevaluesofPPorBPMPorBPMParelowerthan５０％inthe
threeanalysis．PPistheleftvalueatthenodes;BPMPisthemiddlevalue,andBPMListherightvalue．
treewerelowerthanthoseoftheITStreeatthegenus
level．ThissequencemightbenotoptimalfortheperＧ
formanceofmolecularphylogeneticsofMalaxideaebeＧ
lowthesubtribeandgenus．Theseresultshavealso
beendescribedinapreviousstudy(Cameron,２００５)．
ThedatasetsfortheITSandmatKsequences
werecombinedintoasingledataset．Theconsensus
treessupportedthedivisionofMalaxideaeintothree
１５４５期 唐光大等:基于ITS和matK序列的兰科沼兰族分子系统研究及二新种
Fig．２ PhylogramobtainedfromBayesianinferenceanalysisofthechloroplastmatKdataofMalaxideae Numbersatthenodes
arebootstrappercentagesandBayesianposteriorprobabilities,respectively．“－”indicatesthatthevaluesofPPorBPMPorBPMParelowerthan５０％．
BIistheleftvalueatthenodes;BPMPisthemiddle,andBPMListheright．
mainclades,similartotheITSandmatKtrees,butthe
posteriorprobabilitiesandbootstrappercentageswere
higherthanthesetwosequences(PP＝１００％,BPMP＝
９９％,BPML＝１００％;Fig．３)．Thefirstterrestrialclade
２５４ 广 西 植 物 ３５卷
Fig．３ PhylogramobtainedfromBayesianinferenceanalysisofthecombineddataofthenrDNAITSandchloroplastmatK
ofMalaxideae NumbersatthenodesarebootstrappercentagesandBayesianposteriorprobabilities,respectively．“－”indicatesthatthevaluesof
PPorBPMPorBPMParelowerthan５０％．BIistheleftvalueatthenodes;BPMPisthemiddlevalue,andBPMListherightvalue．
wasstronglysupportedbythepolyphyleticgroups(PP
＝９８％,BPMP＝１００％,BPML＝１００％;Fig．３),consisＧ
tingofMalaxis,Liparis,Oberonioides,andthreeunＧ
namedgenus(CladeA,B,C;Fig．３)．Surprisingly,their
３５４５期 唐光大等:基于ITS和matK序列的兰科沼兰族分子系统研究及二新种
Fig．４ LivingplantsofPlatystyliparismalipoensisandP．assamica AＧB．P．malipoensis
A．Floweringplant;B．Flower,frontview CＧD．P．assamica C．Floweringplant;D．Flower,frontview．
morphologicalcharactersweresimilar,withthesame
shortpseudobulbsandconduplicateleaves．ThisterresＧ
trialcladeshouldbenamedLiparidinaebecauseitconＧ
tainsL．loeseli (L．)Rich．,thetypespeciesofLiparis
Rich．Thesecondterrestrialclade,includingCrepidiＧ
um,Dienia,Diteilis,Empusa,andanunnamedgeneＧ
ra(CladeD;Fig．３),wasalsostronglysupportedbased
onthepolyphyleticgroups(PP,BPMP,BPML＝１００％;
Fig．３)．Thespeciesinthiscladeareunitedbyalonger
pseudobulbandplicateleaves．AlthoughsomesubＧ
tribeswerenamedinMalaxideae,suchasMalaxidinae
(Malaxeae)byBentham (１８８３),thetypespeciesof
Malaxis(M．spicata)isamongtheLiparidinaesubＧ
tribe．Thus,weintroducedthenewnameCrepidinae
forthissubtribetentativelyaccordingtotheclassified
historyofMalaxideae(Rasmussen,１９７９;Margońskaet
al．,２０１２)．Thissubtribeconsistsofmainlyterrestrial
plicateleafspecies．ThethirdepiphyticcladeOberoniＧ
inaeconsistsofStichorkis,Platystyliparis,Cestichis,
Oberoniaandtwounnamedgenera(CladeE,F;PP＝
４５４ 广 西 植 物 ３５卷
Fig．５ PlatystyliparismalipoensisG．D．Tang,X．Y．ZhuangetZ．J．Liu １．Floweringplant;２．Flower,sideview;３．Flower,
frontview;４．Dorsalsepal;５．Lateralsepal;６．Petal;７．Lip;８．Poliniaandanthercap(DrawnbyX．Y．MafromtypeZ．J．Liu４４１６)．
９９％,BPMP＝９９％,BPML＝１００％;Fig．３)．
３ Discussion
３．１SystematicstatusofMalaxideae
Malaxideaehavebeendificulttoplace within
Epidendroideaebecausetheylacktypicalappendages
ontheirpolinia,althoughtheyappeartohavesmal
viscidiainmostcases,therearestilsomeexceptional
anduniquespecies(Lietal．,２０１３;Suetal．,２０１４)．
Columnmorphology,particularlythestructureofthe
poliniaandassociatedappendages,hasalwaysbeen
givenpreＧeminenceinorchidclassification．Therefore,
proposalsastotherelationshipswithinthistribehave
beenlimited．Dressler(１９９３)hypothesizedthatalＧ
thoughDendrobieaeappeartobesecondarilynaked,
５５４５期 唐光大等:基于ITS和matK序列的兰科沼兰族分子系统研究及二新种
Fig．６ LivingplantsofCestichispingtaoiandC．manni A,C,D．C．Pingtaoi A．Floweringplant;C．Inflorescence;
D．Flower,frontview B,E,F．C．mannii B．Floweringplant;E．Inflorescence;F．Flower,frontview．
Malaxideaemaybeprimitivelynaked．InthemorphoＧ
logicalanalysesaboutMalaxideae,twospeciesofLiＧ
pariswereamongalargenumberofEpidendroideaeon
apolytomy(Freudensteinetal．,１９９９),whereasanaＧ
nalysisbyCameronetal．(１９９９)onrbcLdatareＧ
vealedthatLiparisandMalaxisweresisterstoDenＧ
drobieaebutwithoutbootstrapsupportgreaterthan
５０％．TheadditionofplastidpsaBormatKsequences
totherbcLdatasetdidnotachieveagreaterresolution
forthe position of Malaxideae (Cameron,２００４;
Freudensteinetal．,２００４)．VandenBergetal．(２００５)
placedDendrobieaeandMalaxideaeastwosuccessive
sistercladestotherestofEpidendroideae．ThesereＧ
sultsrevealedthat Malaxideae wasa monophyletic
cladeandcloselyrelatedtothetribeDendrobinae,alＧ
thoughthewelＧsupportedconsensustreescannotbe
acquired．
InMalaxideaetribe,thebasictaxonomicproblem
６５４ 广 西 植 物 ３５卷
Fig．７ CestichispingtaoiG．D．Tang,X．Y．ZhuangetZ．J．Liu １．Floweringplant;２．Flower,frontview;３．Flower,sideview;
４．Dorsalsepal;５．Lateralsepal;６．Petal;７．Lip;８．PoliniaandAnthercap(DrawnbyX．Y．MafromtypeZ．J．Liu５８１４)．
isthelackofunequivocalcriteriafordistinguishingthe
genera．ManygenerasuggestedinpreviousstudiesapＧ
pearartificialorexhibitpolymorphism (Margońskaet
al．,２０１２)．Eventhelocation,accesstotypespecimens,
andoriginalreferencesareunavailablebecauseoftheir
dispersal,ageandstateofpreservation(Szlachetkoet
７５４５期 唐光大等:基于ITS和matK序列的兰科沼兰族分子系统研究及二新种
al．,２００８)．Mosttaxonomistsagreethatthetraditional
classificationof Malaxideaedividedintothree main
generaisunsatisfactoryandunnatural,butnoclear
strategyhasbeenoferedtoorganizethetribepreviＧ
ously(Cameron,２００５)．Inthisstudy,anupdatedphyＧ
logenyofMalaxideaeisproposedwithamoresampling
oftaxaandcombinedchloroplastandnucleardatathan
previousmolecularphylogenies．AndtheresultssugＧ
gestthatthetribeshouldbesupportedwiththreemain
subtribes,ratherthantwo(Averyanov,１９９１;SzlachetＧ
ko,１９９５)．
３．２MoleculardataanalysisofMalaxideae
IntheMalaxideaetribe,twomaingeneraofLiＧ
parisandMalaxiswerepolyphyletic．ManynewgeneＧ
raareestablishedinrecentyears,butthesegeneraare
controversial．Inthisstudy,weanalyzed１１established
generaand１２１speciesofMalaxideae,andthesetaxa
included mostoftherepresentativegenerainthis
tribe,althoughthesequencesfromspecieswithinsome
generawerenotexamined,suchasAllatiliparis,HipＧ
peophyllum,Seidenforchis andCrossoliparis．The
phylogeneticanalysesofthese１２１specieswereperＧ
formedusingBayesianinference(BI),maximumparsiＧ
mony(MP)andmaximumlikelihood(ML)foreach
dataset．TheBI,MPandMLtreesofeachdatasethad
highersimilartopologicalstructures(Figs．１,２,３),inＧ
dicatingthattheanalysishadthehighcongruenceand
thattheexperimentaldatawerereliable．ThediferＧ
encesbetweentheBI,MPandMLtreeswerethevalＧ
uesofthebootstrappercentagesorposteriorprobabiliＧ
tiesinsomenodes．
ThisstudyrevealedthatthesequenceofITSwas
betterthanmatKinresolvingtheintergenericandinＧ
terspecificrelationshipswithin Malaxideae．TheITS
andcombineddataoftwosequencesproducedapproxiＧ
matelyequivalenttreetopologies(Figs．１,３)．TheconＧ
sensustreeofmatKwasweaklysupportedforthegeＧ
nusＧlevelclassificationofMalaxideae(Fig．２),which
maybeduetoinsuficientvariationsinthesequenceof
matK(Table２)．Manystudieshavedemonstratedthat
atlowertaxonomiclevels(tribesandbelow),theITS
nuclearribosomalDNAspacercanyieldbetterresults
thanproteinＧcodinggenes(Marcinetal．,２０１０)．ConＧ
versely,atthefamilylevel,plastidproteinＧcodinggenes
havebecomethe primaryfocuses,includingrbcL
(Chaseetal．,１９９４,Freudensteinetal．,２００４)and
matK(Freudensteinetal．,２００４)．Thisstudyfocused
onlowertaxonomiclevels(tribesandbelow)．Thus,it
wasreasonablethattheconsensustreesofmatKproＧ
ducedweaklysupportedresults．However,whenthe
ITSandmatKdatasetwerecombinedtoanalyzethese
relationships,thebesttreetopologywasobtainedand
wasconsistentwiththedelimitationbasedontheir
morphologicalcharacters．UsingthiscombineddataＧ
set,mostofthecladeswerebetterdelimitedandmore
stronglysupported．Inthepresentstudy,thematK
fragmentswerestilusefulinresolvingtherelationＧ
shipsbelowsubtribeandthegenericrankwhencomＧ
binedwithITSsequences．
３．３SubtribeＧlevelanalyses
Althoughsomesubtribeshavebeenpreviously
describedbybotanists,theywereconfusinganddidnot
haveconsistentmorphologicalcharacteristics．Bentham
(１８８１)divided“Epidendreae”intoninesubtribes,inＧ
cluding“Microstyleae”(Microstylidinae)whichcomＧ
binedwiththealreadyexisting“Liparideae”(LiparidiＧ
naeLindl．exMiq．)．However,hisdescriptionof“MiＧ
crostyleae”wasveryshort,“anthererectordecumbent
oftenpersistent”,anddidnotlistanyofitsgenera．
Soonthereafter,BenthamandHooker(１８８３)changed
thename“Microstyleae”to“Malaxieae”(MalaxidiＧ
nae)andseparatelydescribed“Liparieae”(Liparidinae
Lindl．exMiq．)．Boththeirsubtribescontainedplants
withapseudobulbousshootandterminalinfloresＧ
cences．Inaddition,thetypicalcharactersof“MalaxeＧ
ae”weredefinedashavingonetothreeleavesinthe
shoots,smaloftenminuteflowers,andanthersthatare
erectordecumbentoftenpersistent．TheyplacedMalＧ
axisandMicrostylisinthesubtribe．Incontrast,“LiＧ
parieae”wascharacterizedasplantswith “１Ｇmany
leavesoraphylusshoot;４rarely８,subequal,gathered
oroftenfree,inappendiculatapolinia．”“Liparieae”
embracedthesegeneraofLiparis (thetypeＧgenus),
Oberonia,Gastroglotis,Microstylis,Chrysoglossum,
Orysicera,andDendrochilum．OberoninaewasdesigＧ
natedbyAveryanov(１９９１),itcontainsonlytwogeneＧ
８５４ 广 西 植 物 ３５卷
ra,Oberonia (thetypeＧgenus)andHippeophyllum．
ManycommoncharacteristicsexistinthesetwogeneＧ
ra,suchasaslendershoot,duplicatefleshy,lateraly
compressedleaves,erectgynostemiumcolumn．Thus,
thelatterwasattributedtoOberonia (Chenetal．,
２００９)．However,the phylogeneticrelationship of
OberoniawasveryclosewithsomespeciesofLiparis
andformedamainsubtribeclade．Therefore,thissubＧ
tribeshouldcontainmoregenerainadditiontoOberoＧ
nia．Inaddition,Liuetal．(２００８)proposedasubtribe
YpsilorchidinaebasedonasingleLiparisspecies‘LiＧ
parisfissipetala’．Theyrecorded “２granularＧwaxy
poliniaeach withasomewhatelasticcaudicleand
deeplybilobedpetals”,buttheresultsofthemolecular
phylogeneticanalysisdidnotsupportthatthesubtribe
Ypsilorchidinaewasasinglesubtribe(Figs．１,２,３)．
Ourresultssuggestthatthetribe Malaxideae
shouldbedividedintothreesubtribes．Thetypegenus
(thetypeisLiparisloeseli)ofLiparidinaeandthe
typegenus(thetypeisMalaxisspicata)ofMalaxidiＧ
naewereattributedtoonesubtribeinthisphylogenetic
analyses．LiparidinaeLindleyetal．(１８５５),hasprecedＧ
enceoverMalaxidinaeBenthetal．(１８８３)．Thus,we
proposethatMalaxidinaeshouldbeattributedtothe
Liparidinae．Moreover,anewsubtribeshouldbeestabＧ
lishedaccordingtotheresultsofthemolecularphyloＧ
geneticanalyses,whichwenamedCrepidinaetentaＧ
tively．Nearlyalofthedatasetssupportedthedivision
ofLiparisinto１１generaandMalaxisdividedintofive
generainthisstudy．Oberoniaisasinglemonophyletic
genusasthetypegenusofsubtribeofOberoninae．
３．３．１SubtribeLiparidinaeLindl．exMiq． Liparidinae
wasestablishedbyLindleyetal．(１８５５),accordingto
thecharacteristicsofcolumnparterect,basalyelongaＧ
ted,andthissubtribecontainedthegeneraLiparis
(thetypeＧgenus),Oberonia,Gastroglotis,Microstylis,
Chrysoglossum,Orysicera,andDendrochilum．HowＧ
ever,GastroglotisisasynonymofDienia,whichwas
establishedbyLindley(１８２４)．MicrostylisisasynoＧ
nymofMalaxis,andChrysoglossumisattributedto
CollabiumbySchlechter(１９１１)．Inaddition,noother
informationisacquiredonOrysicera．Dendrochilumis
replacedinthetribeCoelogyninae(Pridgeonetal．,
２００５)．Recently,１１generawereincludedinthissubＧ
tribe,includingStichorkis,Liparis,Orestias,CrossoＧ
glossa,Kornasia,Lisowskia,Oberonioides,AlatiliparＧ
is,Disticholiparis,PlatystyliparisandCrossoliparis
(Margońskaetal．,２０１２)．However,thesegenera
cladescontainedterrestrialandepiphytichabitstaxa
andlackedconsistentmorphologicalcharacteristics．
Accordingtothismolecularanalysis,thissubtribe
consistsofsixgenera．Malaxiswerepolygeneticand
dividedintoMalaxiss．s．,Oberonioides,Lipariss．s．,
anunnamedcladecontainingM．monophyllos,andtwo
Liparisspecies(Figs．１,３)．Thesegenerasharethe
sameterrestrialvegetativehabits,mainlygrowingin
grasslands,distributingoveraloftheEuropean,Asian
and American temperate grasslands,except for
Oberonioides,whichgrowsontheuniqueDanxiarocks
locatedinsouthChina．Inaddition,theirsimilarconduＧ
plicateleaveswitherinwinter,reservingtheirshort
andfleshypseudobulbstoovercomethecoldwinter．A
highlevelofdiferentiationappearsintheirfloralmorＧ
phology,suchasinflorescence,lipsandcolumn,andthe
lengthoftheovaryinthissubtribe．Ourphylogenetic
resultssuggestthatthesubtribeＧlevelclassificationof
Malaxideaeshouldemploythecharacteristicsoftheir
habits,pseudobulbsandconduplicateleaves．Thus,we
suggestedthatnearlyalofthegrasslandspeciesinthe
tribeMalaxideaeshouldbeincludedinthesubtribeLiＧ
paridinae．
３．３．２SubtribeOberoninaeAver ThiscladewasdeＧ
scribed by Averyanov (１９９１) and revised by
Margońskaetal．(２０１２)Margońskaetal．(２０１２)reＧ
visedthissubtribe．Theyconsideredthemorphological
characteristicsofOberoninaeas“astemＧlike,slender
shoot;duplicatefleshy,lateralycompressedleaves;a
shortand massive,erectgynostemium columnand
suberectanther”．Mostepiphyticgeneraandtaxa
formedasinglesubtribecladeaccordingtothisphyloＧ
geneticanalysis(Figs．１,２,３)．Thus,wesuggested
thatOberoninaeshouldconsistofmoregenerainaddiＧ
tiontoOberonia,suchasCestichis,Platystyliparis,
Stichorkisandtwounnamedgenera (Figs．１,３)．
ThesegeneradistributefromCentralAfricatoFrench
Polynesia,andfromIndiatoNew Guinea．Themain
９５４５期 唐光大等:基于ITS和matK序列的兰科沼兰族分子系统研究及二新种
similarmorphologicalcharacteristicsofthissubtribe
areepiphytichabits,leafduplicateorcompressedunifaＧ
cialorbifacialfeatures．Accordingtotheresultsof
phylogeneticrelationshipof Malaxideae,wesuggest
thatthesubtribeOberoninaeshouldincludemostof
theepiphyticspeciesinthetribeMalaxideae．
In Oberoninae,Resmussen (１９７９)established
thegenusStichorkisThouarsaccordingtothedrawing
ofMalaxisdistichaThours(１８０９),whichwasdesigＧ
natedasiconolectotypes．Disticholiparis Marg．et
Szlach．isanobjectivejuniorsynonymofStichorkis
duetotheidenticaltypespecies(Thouars１８０９,１８２２;
Margońskaetal．,２００４)．Liuetal．(２００８)proposeda
subtribeYpsilorchidinaeonthebasisoftwogranularＧ
waxypolinia,eachwitharelativelyelasticcaudicleand
deepbilobedpetals．OnlythemonotoypicYpislorchis
wasincluded．Itsspecies,Y．fissipetalawasnotrecovＧ
eredasadistinctlineageinthisphylogenticstudy,nor
domorphologicalattributessupportitsdistinctnessat
thesubtribelevel(Figs．１,２,３)．Moredetailedstudies
arerequiredforthisclade．
３．３．３Crepidinaeclade Inthisstudy,weproposeda
newsubtribeCrepidinaetentativelyanddesignatedthe
genusCrepidiumasitstypeＧgenus,becausethespecies
ofCrepidiumaremoreabundantthanothergenerain
thisclade．AlthoughthetypespeciesofC．rheedi
BlumewasdesignatedasthelectotypespeciesofCrepＧ
idium (Seidenfaden,１９７８;Alrichetal．,２０１１),butit
hasattributedtoEmpusa (Figs．１,２,３)．Thenew
subtribeformsastablecladeonthisphylogenetictrees．
ThegenusDieniaisclosewithCrepidium (Figs．１,
３)．ThesubtribeCrepidinaeconsistsoffivegenera
subclades:Crepidium,Diteilis,Dienia,Empusaand
anunnamedgenus(Figs．１,３)．Thesegenerashare
somecommonmorphologicalcharacteristics:theyown
thesameterrestrialvegetativehabit,mainlygrowingin
thedeepforests,anddistributinginthetropicaland
subtropicalislandsofSoutheastAsia．TheyareparticＧ
ularlyabundantontheislandsofMalaysia,Java,IndoＧ
nesiaandNewCaledonia．OnlyonespeciesofDiteilis
nervosa (Liparisnervosa)distributsfromthePacific
islandstoCentralandSouth America．Theirlonger
pseudobulbsandevergreenplicateleavesenablethem
toadapttoforesthabitats．TwoLiparistaxaofL．
laxaandL．chalandeiformamonophylum(PP,BPMP
andBPML＝１００％;Fig．１)asthebasegroupofthis
clade,andthesetwoLiparisspeciesdistributontheisＧ
landofNew Caledonia．Thus,moredetailedstudies
shouldbeperformedonthesespecies．Accordingtothe
mainsimilarmorphologicalcharacteristics,wesuggest
thatthiscladeshouldincludetheterrestrialandplicate
leafspeciesinthetribeMalaxideae,becausetheyshare
samiliarforesthabitatsandlongerpseudobulbs．
４ Conclusions
Inthepresentstudy,weinvestigatedthephylogeＧ
neticrelationshipsofMalaxideaebasedoncombined
molecularevidence．Oberoniaismonophyletic,butLiＧ
parisandMalaxisarenotmonophyletic．Theycould
bedividedintothreemajorclades,whicharethesubＧ
tribesLiparidinae,Crepidinae (Wenamedhere,but
moredetaileddataandnewsequenceshouldbeusedto
confirmitsphylogeny),andOberoninae．TheterresＧ
trial Liparidinae comprises Malaxis,Liparis and
Oberonioidesandthreeunnamedgenera．Thenewly
establishedsubtribeCrepidinaecomprisesCrepidium,
Dienia,DiteilisEmpusaandanunnamedgenus．The
epiphyticOberoninaecomprisessixgenera,Oberonia,
Stichorkis,Cestichis,Platystyliparis andtwo unＧ
namedgenera．TheseunnamedgenerashouldbestudＧ
iedmorecarefulyinthefuturebeforetheirphylogeＧ
neticrelationshipsarerevised．Finaly,wedescribeda
newsubtribeandtwonewspeciesbasedonmolecular
andmorphologicalanalysis．
５ Taxonomictreatment
５．１Newsubtribe
CrepidinaeG．D．Tang,Z．J．Liuet X．Y．
Zhuang,nov．subtribe．NewsubtribeisdistantlyrelatＧ
edtothesubtribeLiparidinae,fromwhichitdifersby
havinglongerpseudobulbs,evergreenandplicateleavＧ
es．Itconsistsofthreeknowngeneraandoneunnamed
genera．TheyareCrepidium,Diteilis,Empusaandan
unnamedgenus．Typusgenus:Crepidium Blume,BiＧ
０６４ 广 西 植 物 ３５卷
jdr．Fl．Ned．Ind．,８:３８７,t．６３(１８２５)．LectotypespeＧ
cies:C．rheedi BlumedesignatedbySeidenfaden,
DanskBot．Arkiv,３３(１):４３(１９７８)．
５．２Newspecies
(１)Platystyliparismalipoensissp．nov．G．D．
Tang,X．Y．Zhuang&Z．J．Liu．(Figs．４,５)
Type:China,Yunnan,Malipo,alt．１４００ m,２７
Feb．２００９,Z．J．Liu４４１６(type,NOCC!)
Diagnosis:ThenewspeciesissimilartoPlatyＧ
styliparisassamica,fromwhichitdifersbyhavingoＧ
voidpseudobulbsandlipbaseconspicuouswith２auＧ
ricularlobesonbothsides,apexcaved．Columnbroad
andlarge．
Description:Epiphyticherbs．PseudobulbsaggreＧ
gate,oblongＧfusiform,１．５－２．５cm×４－６mm,apicalＧ
lywith２－３leaves．LeaveselipticoroblongＧobovate,
１．２－２．６cm ×５－７mm,apexacute,basecontracted
intoashortstalk,articulated．Scape５－１０cmlong,
nearlywingless,nearlowerhalfofracemewith５－８
sterilebracts;raceme３－６cmlong,７－２４Ｇflowered;
bractsovateＧlanceolate,３－４mmlong;pedicelandoＧ
vary３．５－４．５mmlong;flowersyelowishＧgreenwhen
young,laterbecameorangeＧyelow;dorsalsepalnarＧ
rowlyovateＧoblong,５－６×１．５－１．７mm,apexobtuse,
marginsbentforward;lateralsepalssuboblong,５－６×
２－２．２mm,apexobtuse,marginsbentforward;petals
narrowlylinear,５－６×０．３mm,apexacute;labelum
ovateＧoblong,３．７－４mmlong,apextruncateandmuＧ
cronate,１．５mmabovebaseconspicuouscrispedand
flexuose,lookedlike２auricularmaterials,middlewith
athickcalusconcavecentraly;columnerect,３－３．５
mmlong,frontapicalywith２wings,relativelybroad,
belowmiddlewithapairofwingsonbothsides．PolＧ
linia４,２pairs,narrowlyovate．Fl．DecemberＧMarch．
Distributionandhabitat:PlatystyliparismaliＧ
poensisisepiphytic,formingscatteredcolonieson
shadyandwetareasonthetrunksatelevationsof
１２００－１６００minevergreenbroadＧleavedforestsof
Malipo,YunnanProvince,China;HaGiangVietnam．
Conservationstatus:Thenewspecieshasbeen
foundinfourpopulations,eachofwhichhasnomore
than５０individuals．
Polinators:Weobservedthatthisnewspecies
waspolinatedbytheants,butmoredetailedresearch
shouldbetakeninfuturefordeeplyunderstandingthis
uniquespecies．
(２)Cestichispingtaoisp．nov．G．D．Tang,X．Y．
Zhuang&Z．J．Liu．(Figs．６,７)
Type:China,Yunnan,Malipo,alt．１４００ m,１４
Nov．２０１１,Z．J．Liu５８１４(type,NOCC!)
Diagnosis:ThenewspeciesissimilartoCestichis
manni,fromwhichitdifersbyhavingalongerpediＧ
celandovary,flowerlargebutthenumberarelesser,
labelum３－４mmligule,longand２－３mmbroad,
apicalmarginsentire．
Description:Epiphyticherbs．PseudobulbsobＧ
longＧovoid,ovoidandoblong,１．５－２cm ×５－９mm,
nearlyflat,apicalywith１leaves．Leavesnarrowlinear
andlanceolate,９－１７cm×８－１８mm,papery,apexaＧ
cuminate,basecontractedintoashortstalk,articulated．
Inflorescence２１－２４cmlong,petioleflat,verynarＧ
rowlywinged,nearlowerhalfofracemewith１－８
sterilebracts;raceme１３－１６cmlong,１０－２０flowＧ
ered;bractsnarrowlanceolate,４－５mmlong;pedicel
andovary１－１．４cmlong;flowerslightgreenand
white;sepalnarrowlyovateＧoblong,４－５×０．７mm;
petalsnarrowlylinear,５－６×０．４mm;labelumoblong
ligule,３－４×２－３mm,nearly３Ｇlobed,laterallobeseＧ
rect,obovoid,obtuse,midＧlobeoblong,２－２．５×１．８－２
mm,baseecalose,apicalmarginsentire;column１．８－
２mmlong,slightlyarcuateforward,basedilatedand
thick．Polinia４,２pairs,oblongovate．Flowering,OcＧ
toberＧDecember．
Distributionandhabitat:CestichispingtaoiisepiＧ
phytic,formingscatteredcoloniesonthewetareason
thetrunksatelevationsof１３００－１６００mineverＧ
greenbroadＧleavedforestsofMalipo,YunnanProvＧ
ince,China．
Conservationstatus:Thenewspecieshasbeen
foundintwopopulations,eachofwhichhasnomore
than３０individuals．
Polinators:Weobservedthatthisnewspecies
waspolinatedbythefungusgnats．MoredetailedreＧ
searchshouldbeundertakeninfuture．
Acknowledgments We would liketo thank
CHENXuＧHui,GUOXiＧBin,CHENLiＧJunfortheir
１６４５期 唐光大等:基于ITS和matK序列的兰科沼兰族分子系统研究及二新种
helpinthefieldwork,MAXueＧYongforthedrawing
of new species;Philipp Bayer of University of
Queensland,Australia,forhelpinginthemanuscript
preparation．
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３６４５期 唐光大等:基于ITS和matK序列的兰科沼兰族分子系统研究及二新种