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Forest damselflies of the Philippines, their evolution and present status, with the description of Drepanosticta moorei spec. nov. from Luzon (Zygoptera: Platystictidae) PDF

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Preview Forest damselflies of the Philippines, their evolution and present status, with the description of Drepanosticta moorei spec. nov. from Luzon (Zygoptera: Platystictidae)

Odonatologica32(1):39-45 February24, 2003 Forest damselfliesof thePhilippines, theirevolutionandpresent status, withthedescription of Drepanostictamooreispec. nov. fromLuzon (Zygoptera: Platystictidae) J.Van To¹l andR.A.Mülle²r 'National Museum ofNatural HistoryNaturalis,P.O.Box9517,NL-2300 RALeiden,The Netherlands e-mail: [email protected] 2Rehetobelstrasse99,CH-9016 St.Gallen,Switzerland Received June 26,2002/AcceptedAugust30,2002 D.mooreisp.n.(holotype6:Philippines,Luzon,NuevaViscaya,StaFe,AtboRiver, 550-800m, 10-VI-1991,inRMNH)isdescribed,andillustrated.ItiscloselyrelatedtoD. belyshevi Hamalainen from the Philippines.Some generalremarks onthe historical biogeographyandthepresentstatusofthefamilyaremade.Thecurrentdistribution ofthe family(SEAsia,MiddleandnorthernSouthAmerica)presumablydatesbacktotheUpper Cretaceous. INTRODUCTION The familyPlatystictidae seems tobeawell-establishedmonophyletic taxon (e.g. JARZEMBOWSKI etal„ 1998),basedonthepresenceof a“basal post-cubital vein” (FRASER, 1957). BECHLY(1995)doubtsthe homology ofthe latterwiththe post- -cubitalveininthe‘ancientzygopteroid stock’ (e.g. Kennedya Tillyard),andconsiders itthegenuineCuP.Thepresenceofthisveinis acharacterthateasily distinguishes this family fromotherZygoptera, andis anapomorphy ratherthan aplesiomorphy inthe Platystictidae (JARZEMBOWSKIetal., 1998).The family canalsobeidentifiedby someotherstructuralfeatures, suchas the groundplanoftheanalappendages andthe structure ofthe so-calledgomphine-shaped laterallobes ofthe labium.However,the diagnostic charactersof, andtherelationships betweenthe generaare atpresent still unclear. Inthe field, almostall SEAsian species are inconspicuous, brownand brownish black, confinedtoseepagesandsmallerstreams. Mostspecies move very littleduring 40 J.vanTol&R.A.Muller the day.Withtheircrypticcolorationtheyare difficulttofindamongdensevegetation. The males ofsome species have abrightblue spot atthe tipof theabdomen,which perfectly mimicsthewater dropsatthetipofthinbranchesintheforest,thevery sites whereDrepanosticta LaidlawandProtostictaSelys restduring the day. Thefamily occurs insoutheasternAsia,fromSri Lankaover India, southernChina andMalaysia, intotheIndonesiauptotheSolomonIslands.Drepanosticta palauensis Lieftinckis known from Palau(LIEFTINCK, 1962). Most species of SE Asia are assignedtothePlatystictinae, butrecently MATSUKI& SAITO(1996)andWILSON &REELS(2001)foundtwo species forwhichthesubfamilySinostictinaewaserected by WILSON (1997). Another subfamily, thePalaemnematinae,occurs in Middle Americaandinthe northwesternpartofSouthAmerica.AllNewWorld species are assigned tothegenusPalaemnemaSelys. Severaloftheseare remarkably similarto thoseofSEAsia,bothingeneral appearanceand instructural details,such as theanal appendages. Although thehistoricalbiogeography of thefamily shouldbeinvestigated inmore detail,basedonphylogenetic andpreferably alsomolecularstudies, our hypothesisis thatitspresent geographic patterngoes backtothe LateCretaceous (ca65Ma). The familywasatthetimepresumably distributedoverLaurasiaandnorthernNorthAmerica. Theclimatewas tropical, and Europe andAmericawere stillconnected.Aftertheir separation,theclimatebecamelessfavourablefortropicalbiota.Apparently,ancestors ofthe present Platystictidae movedsouthward,following the tropical conditions, in bothNorthAmericaandLaurasia.ThepresenceofPalaemnemainSouthAmericais presumably ratherrecentingeological terms,following theemergenceof theIsthmus ofPanama,theconnectionofNorthandSouthAmericaabout3Ma(COATES, 1999). MostofthePhilippineDrepanosticta species are structurally moresimilartothose oftheMoluccas(undescribed species) andNewGuineathantothemainlandspecies. Thecharacteristicstructures includelargeprocesses posteriorly on theposteriorlobe ofthepronotumandrelatively simple maleanalappendages. Theyarealsoparticularly dullincoloration.Wehypothesize thatthispatternis duetothepresenceofanisland arc along the Pacific plate during the Oligocène andMiocene (20-30 Ma) (HALL, 1998). DREPANOSTICTA OF THEPHILIPPINES DIVERSITY. — Drepanosticta isa very largegenus, withpresently 87 validspecies describedandmorethan20awaitingdescriptioninthecollectionoftheLeidenMuseum alone,mostly fromthePhilippines, theMoluccasandNew Guinea.Mostspecies have smalldistributionalranges.Thesituationmayprove evenmorecomplex, since many species showsignificant geographical variation(cf.e.g.VANTOL, 1999,forSulawesi Protosticta). Although themorphological differencesbetweensuch populations may be rather small, it is not unlikely that thesepopulations are separated formillionsof years.Molecularstudiesby thefirstauthortoaddress thisproblem are inpreparation. Drepanostictamoorei sp.n. 41 Asignificant numberoftheundescribedspecies fromthePhilippineswere collected bythesecondauthorandhiscollaboratorsinthe1980’sand1990’s.Generalinformation ofthis collection, whichis nowhousedinthe NationalMuseumof Natural History Naturalis, at Leiden,was published by HAmALAINEN& MULLER (1997),where alsothehistoryofthestudy ofOdonatainthePhilippines is adequatelyoutlined.The firstDrepanostictaspecies were describedby BRAUER (1868),viz.D. halterataand D. lestoides.It isremarkablethat Semper also collectedsuch inconspicuous insects, whichsuggeststhatthesemusthavebeenverycommon.Thefirstspeciesisstillcommon indeedintheareastudiedby Semper, butD.lestoidesisnot commonatallnow. See alsoTableIforalistofthe 13nominaltaxa, representing 12 species, describedfrom the Philippines up to date. D. septima Needham & Gyger was considered by HAMALAINEN& MULLER(1997) apossible synonymofD. mylettaCowley. Apart from the species described hereunder,more than ten new species will be describedinarevisionofthePhilippine Platystictidae (VANTOL,2003). CONSERVATION. — Likeallbiotaconfinedtotropical forestsandcharacterizedby smalldistributionalranges,mostPlatystictidae areunderseriousthreat.ThePhilippine forestsrankhigh amongthetropical forestsofsoutheastern Asia,andthey are under severepressureofdegradationanddestruction.Thetotalsurfaceofthe7100islandsof thePhilippines amountstojust over 300,000km2.Thetotalforestcoveris ca20%,of whichless than 3%primary forest.AnFAO reportsuggeststhatless than 1million hectaresof“oldgrowth”dipterocarp forestremain. The humanpopulation amounts to ca 62 million, while the total surface ofthe archipelago is similartoItalyinEurope, ortoNewMexicointheUSA. Accordingto arough estimate, nearly500haofsecondary forest disappear daily. Atpresent more than90% ofthe forestis already destroyed; on someislandshardlyany forestis left (e.g.lessthan3%onthe islandof Negros). The Table 1 bestforestremnants can Nominal taxa ofDrepanosticta fromthe Philippines(in chronological be found on Palawan, order) whichis inhabitedby a SSppeecciieess--ggrroouupp OOrriiggiinnaallggeennuuss AAuutthhoorr((ss)) completely different nnaammee floraandfaunaand,bio- geographically doesnot hhaalllteerraaltaa PPllaattyyssttiiccttaa BBRRAAUUEERR., 11886688::555511 form a part of the lleessttooiiddeess PPllaattyyssttiiccttaa BBRRAAUUEERR,, 11886688::555522 Philippines. aannnnuullaattaa PPllaattyyssttiiccttaa SSEELLYYSS,, 11888866:: 115566 mmyylleettttaa DDrreeppaannoossttiiccttaa CCOOWWLLEEYY,, 11993366:: 116600 Conservation llyymmeettttaa DDrreeppaannoossttiiccttaa CCOOWWLLEEYY,, 11993366::116611 Internationaldescribes mmeeggaammeettttaa DDrreeppaannoossttiiccttaa CCOOWWLLEEYY,, 11993366:: 116633 the Philippines as “the ttrriimmaaccuulalatata DDrreeppaannoossttiiccttaa LLIIEEFFTTIINNCCKK,,11993399::114499 hottestof hotspots”. aarriieess DDrreeppaannoossttiiccttaa NNEEEEDDHHAAMM&&GGYYGGEERR,, 11994411;; 114444 Approximately 50%of ttaauurruuss DDrreeppaannoossttiiccttaa NNEEEEDDHHAAMM&&GGYYGGEERR,, 11994411::114455 pphhiilliippppaa DDrreeppaannoossttiiccttaa LLIIEEFFTTIINNCCKK,, 11996611:: 113322 the 8000 native plant cceerraattoopphhoorraa DDrreeppaannoossttiiccttaa LLIIEEFFTTIINNCCKK,, 11997744:: 111177 species are endemic. bbeellyysshheevvii DDrreeppaannoossttiiccttaa HHAÄMMAÄLLAÄIINNEENN,, 11999911::6655 42 J.vanTol&R.A.Muller Thenumberofnameddragonflyspecies ofthePhilippines isca250(HAMALAINEN &MULLER, 1997),ofwhichmorethan50%areendemic,includingallDrepanosticta species. ItisfortunatethatnoneoftheDrepanosticta species isundersuchanimmediate threatas itis forexample theplatycnemidid RisiocnemisseidenschwarziHamalainen (HAmALAINEN, 2000),confinedtoastretchof30mofstreamintheTabunanforest of Cebu. On thisisland, withatotal surface areaof 1707squaremi(4400km2),not morethan 145haofprimary forestare left(pers. comm.Dr Hamalainen,2002). We arehonoured tobe able tocontribute this short paper tothe Festschrift onthe occasion ofDr NormanW. Moore's80thbirthday. ThroughDr Moore’s efforts,Odonata arehighon the international agendaofinvertebrate conservation andprotection(e.g.MOORE, 1997).NamingoneofthePhilippine forestdamselflies afterhimis morethanappropriate. DREPANOSTICTA MOOREI SP. NOV. Figures1-5 Platystictaannulata SELYS, 1886: 156 Drepanostictasp.,LIEFTINCK, 1961: 136(specimen“588”inSelyscollection,IRSN Brussels,aparalectotypeofDrepanostictaannulata[Selys],notconspecificwithlectotype). Drepanostictasp.n.note7; HAMALAINEN& MULLER, 1997:257,276-277, Material. — Holotype3,Philippines,Luzon,NuevaViscaya, StaFe,AtboRiver, 550-800m, 10- -VI-1991 /R.A.Mullerleg./coll. R.A.Muller[JvT22225],coll.RMNH. - Paratypes(allexcoll.R.A. Muller,nowinRMNH):Luzon,NuevaViscaya, DaltonPass,850-900m,8/9-VI-1991,R.A.Muller, 1<J; - samesiteasholotype, 11cj 109; - Luzon,Nueva Ecija,Caranglan,BatchingRiver,700-850 m, 11- -VI-1991,R.A.Mullerleg.,2S\ —Luzon, Auroraprov.,Dilalongan,SitioBiyak,Dibaraybay,3/15-VII- -1996,300-500m,CelsoM,Nazareno leg.; - Luzon, ZambalesProv.,Masinloc, MtGoto,Tal-tal,400- -700m,April1/8-IV-1997,C.M.Nazareno leg. Etymology. —NamedafterDrNormanW. Moore, forhiscontributions totheconservation of Odonata. Aremarkableandunmistakablespecies by thepronotumstructure;thecolorationof the synthorax isalso differentfromthatinmostotherPhilippine congeners. MALE[JvT 22225,Luzon,NuevaViscaya]. - Head. - Labrumandanteclypeus very clearblue,anteriorborderof labrumwithnarrow black line; mandiblesbrown, but innerbasal comerblue; rest of headbronze-black; parorbital carina distinct; postoccipital carinaindistinctandlacking conspicuous extremities. Thorax. — Pronotum(Fig. 3) with anteriorand medianlobe yellowish, lateral lobebrown,posteriorlobedirtyyellow;anteriorlobeapproximately halfway eachside withvery long subcylindrical process, touching medianlobe, andreaching virtually hindmarginofcentralportion ofposteriorlobe; medianandlaterallobessimple; hind lobelaterally withenormous collar, half-round, nearly twiceas long asbasal partof posterior lobe; lateralmargin with long andthin setae. Synthorax bronze-black, but metepistemum nearly fullygreenishyellow,and dorso-posteriorthirdofmetepimeron greenishyellow.Legs dirtyyellow.Wingswith 13Pxinforewing, 13Pxinhindwing; originofR4+5just distaltonodus,R3 arises6 cellsdistaltonodusinforewing,5 cells Drepanostictamooreisp.n. 43 Figs 1-5.Drepanostictamooreisp. n.:(1)maleanal appendages,dorsal view; — (2) same,left lateral view; — (3)male pronotum,obliqueview; — (4)pterostigmarighthind wing,male; — (5)femalelast abdominal segments. inhindwing; Arculusjust distaltoAx2, distinctly stalked; Acand Abforming aY, hardly stalked; numberofcellsbetweendistalendofquadrangle andplacewhereCuP meetshindmarginofwinginforewing5,inhindwing7;pterostigma(Fig. 4)brown, withanarrow palelineagainst veins; ca 1.6 timeslongerthanhigh, theproximal side oblique, thedistalsiderounded;veinsdistadtopterostigma undivided. Abdomen. — Brown, butbasal one-tenthof segment3, basal one-sixth of segments4-6,basalone-thirdofsegment7dirtyyellow; segments8-10tergitesbrown. Appendages (Figs 1-2)greyish brown,topofinferiorscastaneous;superiors withbasal one-fourthstout,restnarrowindorsalview,actually strongly flatteneddorso-ventrally, andbroadlytriangularinlateralview;inferiorsinventralviewsubparallel, thetopbent outward,theninwardinasemicircle, inlateralviewthetopbendsin90°. Measurements (inmm). - Abdomenincludingappendagesca37,hind wing22. FEMALE. — Very similartomale, including coloration,andstructure ofpronotum; segments 8-10brown,anal appendage lightbrown, genital valvessurpassing apexof 44 J.vanTol &R.A.Muller appendages (Fig. 5). DIFFERENTIALdiagnosis. — ThisspeciesdiffersfromotherspeciesofthePhilippine islands by the long processeson the anteriorlobeof the pronotum.This characteris presentinbothmalesandfemales.TheonlyotherPhilippine specieswithlongprocesses ontheanteriorlobeofthepronotumis D. belyshevi Hämäläinen(typelocality Panaon Is.), characterizedby thestrongly curved,slenderprocessesoftheposteriorlobeofthe pronotum,andpalemarkingsintheposteriorthirdsofmetepistemumandmetepimeron (in somespecimens alsoofmesepimeron). distribution. — CentralLuzon(provinces NuevaViscaya, NuevaEcija, Aurora and Zambales). Apparently itoccurs inlow densities over arelatively large area. In larger numbersfoundatthetypelocality only. RELATIONSHIPS. — Basedonthehighlycharacteristicprocesses oftheanteriorlobe ofthepronotum. D.mooreisp. n. seems to bemost closely related toD. belyshevi Hamalainen.Althougha directlinkofthesetwo species tomostoftheotherPhilippine congenersis not apparent,a closerelationship tothemainlandspecies-groups is also unlikely. ACKNOWLEDGEMENTS We aregratefultoDr MATTI HAMALAINEN (Espoo, Finland),who sorted outtheR.A. Muller collection fromthe Philippines,and washelpfulwithexpert advise.Figures I and2 weremadebyB. BLANKEVOORTandFigure3by E.J.BOSCH(bothMuseum Naturalis,Leiden). REFERENCES BECHLY, G.H.P., 1995. MorphologischeUntersuchungenam FlügelgeäderderrezentenLibellen und derenStammgruppenvertreter(Insecta; Pterygota;Odonata) unter besonderer Berücksichtigung derphylogenetischenSystematikund des Grundplanesder Odonata. Petalura (SpecialVol.) 1: 1-341, BRAUER, F., 1868.DritterBerichtüberdievonHerrnG.Sempermitgetheilten,vondessen Bruderauf denPhilippinengesammeltenNeuropterenundBeschreibungeinerneuenLibellen-Gattung.Verh. zool.-bot.Ges. Wien 18:541-558. COATES,A.G., 1999.LithostratigraphyoftheNeogenestrata oftheCaribbeancoastfromLimon,Costa Rica,toColon,Panama.Bull.Am.Paleont. 357: 17-40, COWLEY, J„ 1936.Descriptions ofthree new speciesofDrepanosticta(Odonata)fromthe Philippine Islands. Trans. R. ent.Soc. Land. 85: 157-168. FRASER, F.C., 1957.A reclassificationofthe orderOdonata.R. zool.Soc.NSW,Sydney. JARZEMBOWSBI, E.A., X. MARTINEZ-DECLÔS,G. BECHLY, A. NEE,R. CORAM & F. ESCU1LLIE,1998. The Mesozoic non-calopterygoidZygoptera: descriptionofnew generaand speciesfromthelowerCretaceous ofEnglandandBrazil and theirphylogeneticsignifficance (Odonata,Zygoptera,Coenagrionoidea,Hemiphlebioidea,Lestoidea).Cretaceous Res. 19: 403- -444. HALL, R„ 1998.TheplatetectonicsofCenozoicSEAsiaandthe distributionoflandandsee.In:R.Hall &J.D.Holloway,[Eds],BiogeographyandgeologicalevolutionofSEAsia,pp.99-131.Backhuys, Leiden. Drepanostictamooreisp.n. 45 HAmAlAINEN,M., 1991. Drepanostictabelyshevi spec.nov. tromthe Philippines(Zygoptera: Platystictidae).Odonatologica20:65-68. HÄMÄLAINEN,M,2000.Risiocnemis seidenschwarzi spec, nov.,anendangereddamselflyfromTabunan forestinCebu, thePhilippines(Odonata,Platycnemididae).Ent. Ber.,Amst.60(3):46-49. HÄMÄLÄINEN,M.& R.A.MÜLLER, 1997. Synopsisofthe PhilippineOdonata, with lists ofspecies from forty islands. Odonatologica26:249-315. LIEFTINCK,M.A., 1939. Surquatreespèces nouvelles dePlatystictidaedel’Indeorientale(odonates). Revuefr.Ent.6: 144-154. LIEFTINCK,M.A., 1961. PhilippineZoologicalExpedition 1946-1947. New and interestingOdonata from the Philippines.Fieldiana (Zool.) 42(10): 119-149. LIEFTINCK,M.A., 1962.Odonata. InsectsofMicronesia 5(1): 1-95. MATSUKI, K. & Y. SAITO, 1996. A newspecies ofDrepanostictafrom Hong Kong (Odonata, Platystictidae).Nature andInsects 31(3):39-43. MOORE,N.W., 1997.Dragonflies.Statussurveyandconservation actionplan.IUCN, Gland. NEEDHAM,J.G.&M.K.GYGER, 1937.TheOdonataofthe Philippines,Philipp.J.Sci. 63:21-101,pis 1-10excl. NEEDHAM,J.G.&M.K.GYGER, 1939.TheOdonataofthePhilippines,2,Philipp.J.Sci.70:239-314, pis 11-22excl. NEEDHAM, J.G.& M.K.GYGER, 1941, More Odonatafrom thePhilippines.Philipp.J. Sci. 74: 141- -151,pi. 1excl. SELYSLONGCHAMPS,E. de, 1882. OdonatesdePhilippines.An.Socesp. Hist. nal. 11:[sep.] 1-32, 1 pi. excl. SELYSLONGCHAMPS, E.de,1886.RévisiondusynopsisdesAgrionines.Premièrepartie,comprenant les légionsPseudostigma,Podagrion,PlatycnemisetProtoneura. Mem.cour.Acad.r. Belg.38: [sep.] 1-233. SELYSLONGCHAMPS, E.de,1891.AdditionsauxodonatesdesPhilippines.An.Soc. esp. Hist.nat.20: 209-218. VAN TOL, J., 1999. The Odonata ofSulawesi and adjacentislands,5: The genus Protosticta Selys (Platystictidae).Tijdschr. Ent. 143: 221-266, VAN TOL, J., 2003. Arevision ofthe genusDrepanostictaLaidlawofthePhilippines.Zool. Meded. Leiden — [inpress] WILSON,K.D.P., 1997. The PlatystictidaeofHongKongand Guangdongwith descriptionsofa new genusandtwonewspecies(Zygoptera).Odonatologica26:53-63. WILSON,K.D.P.&G.T.REELS, 2001.OdonataofHainan,China. Odonatologica30: 145-208.

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