InternationalJournalofOdonatology Vol.14,No.1,March2011,55–68 A synopsis of the genus Philosina with descriptions of the larvae of P. alba and P. buchi (Odonata: Megapodagrionidae) HaomiaoZhanga*,VincentJ.KalkmanbandXiaoliTonga aDepartmentofEntomology,CollegeofNaturalResourcesandEnvironment,SouthChinaAgricultural University,Guangzhou510642,PRChina;bNationalMuseumofNaturalHistory,POBox9517,2300RA Leiden,theNetherlands (Received29May2010;finalversionreceived31November2010) AsynopsisofthegenusPhilosinaisprovided.Larvaeofthetwoknownspecies,P.albaandP.buchiare describedforthefirsttime.Thedistributionofbothspeciesisdiscussedandinformationonbehaviour andhabitatissummarized.ThespecializedlarvaeofPhilosinashowastrongresemblancetothoseof Rhinagrion, suggesting that they are sister genera. The unique characters of the larva, especially the arrangementandstructureofthecaudallamellae,meanthatneithergenusfitsintoanyofthecurrently recognizedfamiliesofZygoptera.Itisnotedthatthesegeneracouldbeplacedintheirownfamily.However, cautionisexercisedpendingabetterunderstandingofthefamilyMegapodagrionidaebasedonDNAwork, andtheyarethereforeretainedinMegapodagrionidae. Keywords: Odonata;dragonfly;Zygoptera;Megapodagrionidae;Philosina;Rhinagrion;taxonomy;Asia Introduction The genus Philosina was established for Philosina buchi Ris, 1917, which was described in thesamepaperfromFujian,China.PhilosinawasplacedinMegapodagrionidaebasedonwing venation,butwasconsideredsufficientlydistinctfromothergenerainthisfamilytobeplacedinits ownsubfamilyPhilosininaebyKennedy(1925)andlaterauthors(e.g.Davies&Tobin,1984).In 1999asecondspeciesofthegenus,P.albaWilson,1999,wasdescribedfromGuangdong.Onlya handfulofrecordshavebeenpublishedforeitherspecies,informationontheirdistribution,habitat and behaviour is scarce, and no larva has been described for the genus. Based on resemblance oftheadultsandpreliminaryDNAresultsKalkman,Choong,OrrandSchütte(2010)suggested thatPhilosinaandRhinagrionCalvert,1913mightbecloselyrelated.ThelarvaofRhinagrionis structurallyuniqueamongtheZygopteraanditwassuggestedthatlarvalcharactersofPhilosina, oncediscovered,wouldeitherconfirmorfalsifythehypothesisofacloserelationshipbetween PhilosinaandRhinagrion.HZwasabletocollectlarvaeofbothPhilosinaalbaandP.buchiwhich areheredescribedforthefirsttime. *Correspondingauthor.Email:[email protected] ISSN1388-7890print/ISSN2159-6719online ©2011WorldwideDragonflyAssociation DOI:10.1080/13887890.2011.568189 http://www.informaworld.com 56 H.Zhangetal. Materialandmethods TerminologylargelyfollowsWatsonandO’Farrell(1991). Acronymsforcollectionsareasfollows: SCAU SouthChinaAgriculturalUniversity,Guangzhou,China CUMZ CambridgeUniversityMuseumofZoology,Cambridge,UK NMSF ForschungsinstitutundNaturmuseumSenckenberg,Frankfurta.M,Germany RMNH NationaalNatuurhistorischMuseumNaturalis,Leiden,theNetherlands RecognitionofPhilosina Adults of Philosina can be distinguished from all zygopteran genera except Rhinagrion by the combinationofthefollowingcharacters:(1)twoantenodalcrossveins;(2)Ptmorethantwiceas longasbroad;(3)R4andIR3originatingwellbeyondmidpointbetweenthearculusandthelevel ofthesubnodus;(4)mediancleftoflabiumdeep,abouthalfthelengthoftheprementumitself. PhilosinaalbaWilson,1999;(Figures1–4,5a–d) Specimensstudied Adults:2(cid:2),China,HainanProvince,LishuiCounty,Mt.Diaoluoshan,900m,24April2009,leg: HZ, SCAU. — 4 (cid:2), 1 (cid:3), China, Guangdong Province, Conghua City, Liuxihe National Forest Park(23◦43(cid:3)51(cid:3)(cid:3)N,113◦48(cid:3)24(cid:3)(cid:3)E),15May2009,leg.HZ,SCAU.—1(cid:3),China,HainanProvince, LishuiCounty,Mt.Diaoluoshan,23April2009,leg:HZ,RMNH.—1(cid:2),China,HainanProvince, LishuiCounty,Mt.Diaoluoshan,15May2009,leg:HZ,RMNH. Larvae(finalinstar):1(cid:2),1(cid:3),China,GuangdongProvince,ConghuaCity,LiuxiheNational ForestPark(23◦43(cid:3)51(cid:3)(cid:3)N,113◦48(cid:3)24(cid:3)(cid:3)E),12March2009,leg.HZ,SCAU.—1(cid:2),samelocation, 25November2008,RMNH. Differentialdiagnosisofadults Malesareeasilyrecognisedinthefieldbytheirhabitofperchingwiththeirwingsheldflat,their mediumsize(Hw28–31mm),moderatelyheavybuildandthewhitish-bluepruinositycovering allsegmentsoftheabdomen(Figure5a–c).ThethoraxdoesnotbecomeaspruinoseasinP.buchi Figure 1. DistributionofthespeciesofPhilosina. SynopsisofPhilosina 57 Figure 2. HabitusoffinalstadiumlarvaofPhilosinaalba. and the yellow antehumeral stripes are clearly visible even in fully mature males. Females are lessstrikingthanmalesbutarestilleasytoidentifybytheirmediumsizeandblackabdomenwith broadlateralyellowstripesandathinmedianstripeonS1-7(Figure5d).Theonlyspecieswith a very similar pattern is P. buchi, but in this species the antehumeral stripe runs the full length of the mesepimeron (only for four-fifths in P. alba), and there is an additional thin yellow line belowthemesopleuralsuturewhichisabsentoronlypresentalongtheposteriorhalfofthesuture inP.alba. Descriptionofthelarva A stout, brownish larva with depressed head and short abdomen (Figure 2), surface of body coveredwithminutepyramidalnodules. Head. Headflattenedabove,widestacrosstheeyes,rearofheadsquarebuthindcornerrounded andbearingirregularstoutspines,hindmarginofocciputconcave.Rowofsixshortspinesalong the eye-margin on the underside of the head. The labrum, clypeus and frons are entirely dark brown; anterior margin of anteclypeus with a row of fine setae. Ocellar triangle pale brown, ocelli large and distinct.Antenna with seven segments, long and filiform with fine setae, pale brown (Figure 3d).Antennal segments 1, 2, 4 and 5 each with a black subapical ring. Length of each antennal segment (mm): 0.40; 0.60; 0.68; 0.64; 0.40; 0.28; 0.20. Mandibles biramous, incisoroftherightmandiblewithsixteeth,identicaltotheleftone,themolarsurfacewithone tooth,lessdeveloped,hookshapedandnotbifidapically(Figure4a).Incisoroftheleftmandible with six teeth, the molar surface with one large tooth, apically bifid (Figure 4b). Maxilla with four long, stout dorsal hooks and three smaller ventral hooks, sharply pointed (Figures 4c, d). Basallywithlongsetae.Maxillarypalpcurvedandtaperingtoabluntpoint,bearingdense,long setae. Labium broad and flat (Figure 3a). Length to width ratio of prementum 1.25:1. Median lobestronglydeveloped,anteriormarginstronglyconvexwithregularshort,crenulationsandan 58 H.Zhangetal. Figure 3. LarvalstructureofPhilosinaalba:(a)labium,dorsalview;(b)ventralviewoffemaleabdomenshowing gonapophyses;(c)rightlateralpalpoflabium,dorsalview;(d)antenna(setaenotshown);(e)patternonmedianlamella, lateralview;(f)patternonlaterallamella,lateralview. apicallyclosedmediancleft(Figure3c).Outermarginofprementumwithstrong,sharp,forward pointingdenticles.Prementalsetaeabsent.Labialpalpnarrow,bearingthreeinner,subapicalsetae andapicallywiththreestrongandsharplypointedhooks,themedianonelargest;movablehook longandsharplypointed(Figure3c). Thorax. Thoraxbrown,sidesofprothoraxwithtwoprominences,onebeneathpronotumwiththe outermarginslightlyconcave,theloweronepyramid-shaped,approachingthefrontcoxa.Distal SynopsisofPhilosina 59 Figure 4. LarvalstructureofPhilosinaalba:(a)rightmandible,ventralview;(b)leftmandible,ventralview;(c)right maxilla,ventralview;(d)same,dorsalview. marginsoftheseprominenceswithpyramidalspines.Pronotumwelldevelopedandshield-shaped; brown with a pair of black spots near the hind margin.Anterior margin more or less straight, anterior corner forming an angle of 135◦, posterior corner rounded. Synthorax dark brown and robust, wing buds narrow, reaching the hind margin of abdominal S8. Legs with hind femora reachingS9,coveredwithfinesetaeandwithrowsofshortspines.Allfemoraandtibiaebrown withthreepalebands.Anteriormarginofeachfemurwitharowofshortpyramidalspines.Tarsi withthreesegments,clawssimple. Abdomen. Brownanddenselycoveredwithfinesetae.Cylindricalinshapeandslightlytapering backwards,withapalemiddorsallineoneachsegment.Lateralspinesanddorsalhooksabsent. Caudal lamellae arranged vertically, broad and oblong, largely black with marginal pale spots, themarginbearingwithlong,finesetae.Themedianlamella(Figure3e)isshorterthanthelateral ones(Figure3f)andhasonbothsidesaslightlyexpandedmidrib,whichisfreeofspines.The laterallamellaehavebasallyanexpandedmidribontheexterior,bearingdense,shortspines.The medianlamellaisthinnerthanthefleshy,undulatinglaterallamellae.Femalegonapophyseslong, almosttwiceaslongasS9-10(Figure3b). 60 H.Zhangetal. Figure 5. AdultsofPhilosinaspecies.P.alba:(a)malefromMt.Diaoluoshan;(b)pairinwheelfromDiaoluoshan; (c)malefromLiuxiheForestPark;(d)femalefromLiuxiheForestPark.P.buchi:(e)malefromLongsheng;(f)female fromLongsheng.PhotosbyHZ(a,b,c,d),andVJK(e,f). Measurements(mm). Bodylength(includinglamellae):14.1;maximumwidthofhead:4.4hind femur:5.0;hindwingbuds:4.8;medianlamella:3.8;laterallamellae:5.5. Distribution,habitatandbehaviour PhilosinaalbawasdescribedfromDinghushan,ZhaoqingCity,Guangdong,China(Wilson,1999; Wilson&Xu,2007)andhassincebeenfoundintheChineseprovinceofHainan(Wilson&Reels, 2001)andintheLakSaoareainLaos(Karube,2002)(Figure1). SynopsisofPhilosina 61 The habitats at Diaoluoshan and at Liuxihe National Forest Park are largely shaded, large, rockybrooksinforest(Figure10).Thespecieshasbeenrecordedfrom350–500ma.s.l.(Liuxihe NationalForestPark)andbetween850–1000ma.s.l.(Diaoluoshan). The earliest seasonal record is that of an immature male on 25 March 2008 at Diaoluoshan NationalNatureReserve,Hainan,observedbyHZandShanlianMo.InlateAprilofthatyearmore than10maturemaleswereobservedinthreedays.AtLiuxiheNationalForestPark(Guangdong Province)morethan50maleswereseenonasingledayinmidMay.Thespecieswasstillcommon thereinJunebutbymidJulyonlytwooldmalescouldbefound.Basedontheserecordstheflight periodextendsfromtheendofMarchwellintoJuly,withthehighestdensitiesfoundfromlate ApriltoJune. WilsonandReels(2001)statedthatmaleshold“territoryonhorizontalsurfacesoflogsandtree roots,whichoverhangsmallfastrunninghillstreams”.AtDiaoluoshanandatLiuxiheNational ForestParkmalesusuallyoccupiedterritoriesontheslowerflowingpartsofthestreamatplaces borderedbydensevegetation.Theyperchedontreesorbranchesalongtheshadystreamsatabout 0.5–2mheightandestablishedterritoriesbutseldompatrolled.Maleswithaterritorywereseento driveothermalesaway,duringwhichactivitytheywerehoveringfacetofacemidair.Following a chase the male was seen to return to his territory.When disturbed, they quickly moved away withasuddenandrapidflight.Nocourtshipwasobserved;asinglecopulationwasobservedin theafternoon. PhilosinabuchiRis,1917(Figures1,5e–f,6–9) Specimensstudied Adults:15(cid:2),1(cid:3)(syntypes),FujianProvince,1916,leg.P.A.Buch,NMSF(numberedas7192, 7193, 7194, 7200, 7205, 7210–7212, 7214–7217, 7219, 7220, 7222, 7223 (female)). — 17(cid:2), FujianProvince,1917,leg.P.A.Buch,NMSF(numberedas7191,7195–7199,7201–7204,7206– 7209,7213,7218,7221).—2(cid:2),2(cid:3),China,GuangxiProvince,GuilinCity,Longsheng,11–15 July2005,leg.VJK,RMNH.—1(cid:2),4(cid:3),China,GuangxiProvince,betweenLongshengandhot springs,13July2005,leg.VJK,RMNH.—2(cid:2),China,GuizhouProvince,LiboCounty,Maolan Figure 6. HabitusoffinalinstarlarvaofPhilosinabuchi. 62 H.Zhangetal. Figure 7. LarvalstructureofPhilosinabuchi:(a)labium,dorsalview;(b)ventralviewoffemaleabdomenshowing gonapophyses;(c)rightlateralpalpusoflabium,dorsalview;(d)mediancleft,dorsalview;(e)antenna(setaenotshown); (f)patternonmedianlamella,lateralview;(g)patternonlaterallamella,lateralview. NatureReserve,ZhangjiangRiver,7May2007,leg.HZ,SCAU.—1(cid:2),China,samelocality,27 July2008,leg.HZ,SCAU. Additional records: China, Guangxi Province, Guilin City, Longsheng, 11–15 July 2000, M.Wasscher(pictureofmalepublishedonwww.asia-dragonfly). Larva: 1 (cid:3), China, Guangdong Province,Yingde City, Qiaotou,Wengjiang River (24◦16(cid:3)36(cid:3)(cid:3) N,113◦46(cid:3)38(cid:3)(cid:3)E),27February2009,leg.YufengWang,SCAU. SynopsisofPhilosina 63 Figure 8. Larval structure of Philosina buchi: (a) right mandible, ventral view; (b) left mandible, ventral view; (c)leftmaxilla,ventralview;(d)same,dorsalview. Differentialdiagnosisofadults Malesareverystrikingandareeasilyrecognizedbytheirlargesize(Hw39–41mm),bluepru- inosityonhead,thorax,S1-6andS10andbrightredS7-9oftheabdomen(Figure5e).Withtheir sizeandstrongflighttheycanbemistakenforAnisoptera,especiallyonthewing,butnoChinese Anisoptera species has S7-9 red with the remainder of the abdomen blue.The genital ligula is illustrated in Figure 9a, b. Females are less striking than males but are still easy to identify by theirlargesizeandblackabdomenwithbroadlateralstripesandathinmedianstripefromS1-7. DifferencesfromthefemaleofP.albaaregiveninthediagnosisofthatspecies. Descriptionofthelarva A stout, brownish larva with depressed head and short abdomen (Figure 6), surface of body coveredwithminutepyramidalandroundednodules. Head. Headflattenedabove,widestacrosstheeyes,rearofheadsquarebuthindcornerrounded, withabroadblackstripeoneachside,andbearingstoutdenticles;hindmarginofocciputconcave. Six spines are present along the eye-margin on the underside of the head. Labrum and clypeus entirely dark brown. Frons largely brown with a pale central spot. Ocellar triangle pale brown; ocelli large and distinct.Antenna with seven segments, long and filiform with fine setae; pale 64 H.Zhangetal. Figure 9. SEMphotographsofthegenitalligulaof:(a)Philosinabuchi,China,Gaungxi,nearLongsheng,2005,ventral view;(b)samelateralview.Photos:DirkGassmann. brown, darkening toward the apex of each segment except the basal one (Figure 7e). Length of each antennal segment (mm): 0.52; 0.88; 0.72; 0.44; 0.56; 0.56; 0.48. Mandibles biramous, incisor of the right mandible with six teeth, similar to the left one, the molar surface with one, less developed, tooth (Figure 8a). Incisor of the left mandible with six teeth, the molar surface withonelargetooth,apicallybifidandpointed(Figure8b).Outermarginofeachmandiblewith two rows of well developed large spines. The ventral row contains six sharply pointed dentate spines.Maxillawithfourlong,stoutdorsalhooksandthreesmallerventralhooks,sharplypointed. Basallywithlongsetae.Maxillarypalpcurvedandtaperedtoabluntpoint,bearingdense,long setae.Labiumbroadandflat(Figure7a).Ratiooflengthtowidthofprementum1.3:1.Median lobestronglydeveloped,anteriormarginstronglyprojectingwithregularshortcrenulationsandan apicallyclosedmediancleft(Figure7d).Outermarginofprementumwithstrong,sharp,forward pointingdenticles.Prementalsetaeabsent.Labialpalpusnarrow,withthreesubapicalsetaeand threestrongandsharplypointedhooksapically,themedianonelargest;movablehooklongand sharplypointed(Figure7c). Thorax. Thoraxbrown,sidesofprothoraxwithtwoprominences,onebeneathpronotumbifid apically;theloweronepyramidal,approachingthefrontcoxa.Distalmarginsoftheseprominences withpyramidalspines.Pronotumshield-shaped,itsanteriormarginmoreorlessstraight,anterior corner forming an angle of 135◦, posterior corner rounded. Synthorax dark brown and robust, wingbudsnarrow,reachingthehindmarginofS7.Legswithhindfemorareachinghalfwayalong S8, covered with fine setae and rows of short spines. All femora and tibiae brown with three