E:h,l,・a,'fok、o,25(2):349-417,Novemberl5,1997 ARevisionoftheJapaneseSpeciesoftheGenera E1odesandSacodes(Coleoptera,Scirtidae) HiroyukiYosHIToMI ZoologicalLaboratory,MeijoUniversity, Tempaku-ku,Nagoya,468Japan* Abstract The Japanese speciesof the genera E/odes LATREILLE, 1796 and SacodesLECoNTE, l853arerevised.SacodesisaseniorsynonymofFlavo1/e1odesKLAus- NITzER,1980.Fivespeciesof thegenusE1otiesandsixofthegenusSacot/esarerecog- nized. 0fthese,E/odeselegansandSacode.・l tsils/11,nenslsarenew toscience.TheJapa- nesespeciesofE/odesaredividedintotwospecies-groups:themat-gl,lataspecies-group comprisingE. ino1・nata L三1wis, E.、l・11so111PIcandE.sapu/a1・IsLEWIS,andthe,11imlta species-groupcomprisingE elega11ssp nov and E koJ1111a1NAKANE.E1odesohba、,ash11 M. SATo and Flavo/1e1odes t11grata KLAUsNITzER are junior synonymsof E kojimal NAKANE.Sal'aband1ls111ontlcolaNAKANEisajuniorsynonymofE. _1son1Plc.Fournew combinationsareproposedinthegenusSa(・odes.vl,.,naka11el,a,11amiens1s.,,nln1,naand 加_、. Descriptionsofthetwogeneraaregivenwithkeystospecies.Alltheadultsheredealt ,. withandtheimmaturestagesof 加etna'a,9.ll,fisc川,1.人,o/加al,S.IlaXane1 S pl-o'ecf . andSdu.、-aredescribedorredescribedwithfigures. Introduction E1odesLATREILLE,1796isthethirdlargestgenusofthe familyScirtidaeandis representedbyabout100speciesfromallovertheworld.Flavohe1odesKLAUsNITzER, 1980waserectedforthefavlco11isspecies-groupofthegenusE1odesandcontains17 speciesrecorded fromtheHolarcticandtheOrientalRegions.Thesegeneraarecon- sideredcloselyrelatedtoeachotherandshowsister-grouprelationshipwithotherscir- tidgeneraon thebasis of phylogenicstudies of adults and larvae (KLAusNITzER, 1974a; HANNAPPEL &PAULUS,1987). Unfortunately, Flavohe1odes isajuniorsyn- onymofSacodesLECoNTE,1853describedonthreeNorthAmericanspecies. ThefirststudyontheJapanesespeciesofthegenusE1odesanditsrelativeswas madebyKIEsENwETTER(1874)whorecordedHe1odesflavlco11isKIESENwETTERfrom Nagasaki.Sincethen,manytaxonomicandfaunalstudieswerepublishedbythe fol- lowingentomologists: HAROLD (1881); LEWIS (1895); PIC (1914, 1918); NAKANE *Presentaddress:BioindicatorCo., Ltd.,Takada3-l6-4,Toshima-ku,Tokyo,171Japan 350 HiroyukiYosHIToMI (1963); SAT0 (1966, 1985a, b, l989), SAT0 & CHUJ0 (1972); SAToU (1975) and KLAUsNITzER(1973,1974b,1980a,1995). Twelvespecieshavehithertobeenrecog- nized fromJapanunder the following threegenera:E1odes, Flavohe1odesandSa1-a- bandusLEECH,1955,butnocomprehensivestudyaboutthesehasyetbeenmade.Be- sides, therehasbeennostudyabout theimmaturestageandbiology, except HAYAsHl (l957). Inthispaper,IamgoingtorevisethegeneraE1odesandSacodesofJapan,with biological notesanddescriptions of theimmaturestages ofsixspecies,viz., three E1odesandthreeSacodes. MaterialsandMethods Thisstudy ismainlybasedondried(adults)andalcoholic(larvae, pupaeand adults;preserved in70%ethanol)specimensinmycollection. Inaddition, I wasable toborrow andexaminemanyspecimenspreserved in the following institutionsand personal collections: EUM-Entomo1ogica1 Laboratory, Ehime University, Matsu- yama;KMNH- KurashikiMuseumofNaturalHistory;NWU-Bio1ogica1Laboratory, Nagoya Women's University; TMNH-Toyohashi Museum of Natural History; TPM -Tochigi Prefectural Museum; TN-T. NAKANEs' private collection; SO-S. OHMOM0s' privatecOllection;SI-S. IMASAKAs' privatecOllection;SS-S. SAKURAI's privatecollection;YH-Y.HIRANos' privatecollection. Theho1otypesandsomeparatypesofthetwospeciestobedescribedhereinwill bepreservedintheNationalScienceMuseum(Nat.Hist),Tokyo,andotherparatypes areintheBiologicalLaboratory,NagoyaWomen'sUniversityandinmypersonal col- lection. Generalobservationanddissectionweremadeunderastereoscopicmicroscope. Microstructureswereobservedunderamicroscopeondissectedpartmountedonhol- lowslideswithpureglycerine(mainlyusedfortegmenandpenis)orCanadaBalsam (used forotherparts).Somemicrostructuresofadultsandlarvaewereobservedwith SEM (HitachiS-2050). TheterminologyrefersgenerallytoNYHoLM(1972,1984) for thegenitalorgan, andmainlytoHANNAPPELandPAULUS(1991) for themouthpartsofthelarva. The termsofsomestructuresoflarvalmouthpartsareshowninFig.1. Theabbreviations formeasurementsused inthepresent paper areas follows: HW- widthofhead;PL-lengthofpronotum;PW-widthofpronotum;EL-lengthof elytra; EW-widthofelytra;TL-total length (PL plusEL inadult); TW-maximal width(betweenleftandrighthindlegsinpupa).Totallengthismeasuredfromanterior marginofclypeustoapex ofabdominalsegment in fullyextended larva,and from apexofpronota1spinetoapexofabdominalsegmentinpupa.Arithmeticmeanofthe measurement isgiveninparenthesesaftertherange. Ifpossible,twentyspecimensare pickedup formeasurement inbothsexes. E1odesandSacodesofJapan 351 Tb A Fig. 1. LarvaofE1odesino・,nata,showingtermsoflarvalmouthparts. - A,Labruminventralaspect (epipharynx);B,hypopharynx.C:Cushion;Ca:clawapparatus;Cd:chitindent;Cp:combplate;Ct: clawteeth;Et:epipharyngea1teeth; Ks:keel-sclerite;Tb:tooth-bristles;VI:ventrallobe;Vs:ventral seta. GenusElodesLATREILLE [Japanesename:MaruhananomiZoku] E1odes LATREILLE, 1796, Prec. Caract. Gen., 44. - PIc, 1914, Coleopt.Cat., (58): 21 [world list]. - PoPE,1976,Entom mon.Mag.,111:186.Typespecies:La''lpyt'is川' nutaLINNAEUS, l767(sub- sequentdesignationbyLATREILLE,1810). He1oales[incorrectspelling]:AGASSIZ,1847,Nom.Zoo1. lndexUniv.,393. - KLAusNITzER,1974,Zool. Jb.Syst.,101:479. - SAT0,1985,Coleopt.JapanCol.,Osaka,2:420[key]. Sa,abandus:SAT0,1985,Coleopt.JapanCol.,Osaka、2:420[key]. ecLEECH,l955]. Adult. Bodymoderateinsizeforscirtidbeetles,TL2.8-6.0mm,ovaloroblong, lightly or moderately convex above,shining,closelycoveredwithhairysetae. Col- orationyellowtoblack,variablewithspecies.Headrathersmall,slightlyconvex,com- pletelycoveredbypronotumandinvisibleindorsalview.Antennaefiliform,covered withshortsetae,moreor lesslonger inmale;scapeovate,broadest;pedicel ovate, small;3rdthesmallest,withdistalmargindiagonal;4ththelongest inmostspecies; l ithoblong. Pronotumsemicircular, covered with setaewhichareeasily removed. Scutellumtriangular,moderateinsize,visibleindorsal view.Elytraovalor oblong, with rather longsetaewhichareeasily removed. Abdominal sternitescoveredwith shortsetae. Malegenital organ. Eighth tergitewellsclerotized, with apair of rather long apodemes;8thsternitewellsclerotized;9thtergitemoderatelysclerotized,withapair ofapodemes;9thsterniteweaklysclerotized,coveredwith longsetae. Tegmenwell sclerotizecLrathersmaller ofthesamesizeaspenis,withdistinctparameres.Penis wellsclerotized,stick-like,bifidinapicalpart(parameroids). Femalegenitalorgan. Eighthtergitewithapairof rather longapodemes; 8th 352 HiroyukiYoslln'oMI s1erniteovaltooblong.Ovipositorwithratherlongbaculus. Lan,a. Bodywellsclerotize(i,campodeiform.Headvisibleindorsalview.An- tennaefjliform,rathershort,reachingthorax;scapecurvedposteriorly,flagellumabout 30_50segmented. Labrumtransverse,withalmoststraightanteriormargin;epiphar- ynxwithlongandsimplesetaeinanteriormarginofventrallobe,withapairofventral setaewhosesocketshavesimplesurroundingparts. Mandiblespointedat terminal tooth,with featheredbristlesininnerareas.Maxillarypalpiwithsomeshortsetaein ventro-1atera1areaof1stsegment.Hypopharynxwithseparatedkeel-scleriteandsock- etsoftooth-bristles,tri-ormulticornuteatapicalmarginsoftooth-bristles,withapair of longsetaeonkeel-sclerite.Thoracesdistinctlybroader thanabdomen.Ninthab- dominaltergitebicornuteatposteriormargin. Pupa. Bodyverysoft,withtwopairsoflongspinesondorsalsurfaceofprono- tum,withsimpleabdominalsegments. Rema,・ks. Cistelapa11idaFABRIcIUs,1775wasdesignatedby LATREILLE(1810) asthetypespeciesof thegenusE1odesLATREILLE,1796. However, thisspecieshas been regardedasajuniorsynonymofE1odeslmntlta (LINNAEUS, 1767) which isa commonandwellknownspeciesinEurope. Thegenus is divided into two species-groups, namely ,na,-g1,lata and mlnuta species-groups(KLAUsNITzER,1970;NYHOLM,1972). Biologicalnotes. Thehabitatsof thelarvaeso far reportedaresmall streams andclearponds(e.g.,LESAGE,1991;KLAUSNITZER,1996). KeytotheJapaneseSpeciesoftheGenusElodes 1. Antennaandlegsmoderateinlength;bodyoval;(,na1glnataspecies-group) _ _2. - Antennaeandlegslong;bodyoblong,almostparallel-sided;(,11inutaspecies-group) 4 2.Pronotumwithshallowanddistinctconcavities,seeminglydistinctlypunctate E. ino1'nata LEWIS - Pronotumfinelypunctate 3. Fronsandclypeusyellowinmale;7thsterniteshallowlyconcave...E.wiison1PIc - Fronsandclypeusbrownishblack;7thsternitedeeplyconcave E.scapularisLEWIS*) 4. Pronotumwithblackmarking;8thtergiteofmaletrapezoidal ! .A' 濶チa NAKAN *) Femalehasnotbeendeterminedyet Fig. 2. HabitusofE1odesspp. - A, E. inc,・11ataLEWIS,male; B, E.、l,fisc,11Pfc, female(holotypeof Sa'abandusmonticolaNAKANE,1963);C, typelabelsofSa''abandus'i1onticola;D, E scapula,・is LEWIS,male;E,Ee/eganssp nov.,holotype,male;Fditto,paratype, female;G,E koJima1'NAKANE, holotype,female;H,ditto,labels;I,ditto, female(1、olotypeofE.o/lba、,as/111M.SAT0,1985). E1odesandSacodesofJapan 353 354 HiroyukiYosHIToM :,11・ .-,.-,・・ - : , I 益 - -ま f 9 '、 .・ 1 Fig.3. TypespecimensofE1odesspp.preservedintheBritishMuseum,London. - A, E. inc・,nata LEWIS;B,E.scapula''IsLEWIS.PhotobyM.SAT0. - Pronotumyellowishorange;8thtergiteofmaleformedbyrod-likehemitergites Eeleganssp nov. GroupofElodesmarginata Thisspecies-groupischaracterizedbythefollowingfeatures:bodyoval;antennae and legsrathershort; parameresdistinctlyserrate; apicesofpenis ratherround;ab- dominaltergitesmorestronglysclerotizedthanthoseoftheminutaspecies-group. ElodesinornataLEWIS,comb rev. [Japanesename:Ko-kuro-maruhananomi] (Figs 2A,3A,4A,5A,6-12,20A) He1odesinornatusLEWSI,1895,Ann.Magnat.Hist.,(6),16:107(Type:inBritishMuseum,London,not examine Fig3A). E1odesinornata:PIc,1914,Coleopt.Cat.,(58):23. Sa'a'ha'tdusinc''natus:NAKANE,1963,Icon.Ins.Japon.Col nat ed.,2:140,pl 70,fig. 18. _ SAT0, 1985,Coleopt.JapanCol.,Osaka,2:424,pl 77,fig31. Adult. Body oval,moderately convex, closely covered with yellowish white E1odesandSacodesofJapan 355 A M B C -M (=l(::) D E _ Fig 4. AntennaeofE1odesspp.(above,male;below,female). - A,E. inc・,nataLEWIS;B,E.、、,ilso,ll Pfc;C,1.scapll/a,-isLEWIS;D, .e/ega1lsspnov;E, .A'oJ1'italNAKANE.(Scale:0.5mm) hairs.Headblack; labrumyellowishbrown;maxillaryandlabialpalpiandmandibles yellow,buttheapicesofmaxillaryandlabialpalpiaresomewhatdarker;1stto3rdan- tennalsegmentsyellow;4thantennalsegmentbrownishyellow, itsapicalendsome- whatdarker;5thto11thantennalsegmentsbrownishblack,but5thand6thsegments paler; pronotumblack, butanterior and lateral margins paler;scutellumandelytra brownishblack;ventralsurfaceofbodyblack; forelegsyellow,except forduskytarsi; midcoxaeyellowtobrown,variableincolor;midfemorayellow,except forblackish brownapexandposteriormargin;midtibiaeandtarsiblack;hindlegsblack,buthind tarsipaler. Headfinelyandsparselypunctate.Eyesmoderateinsize,prominent;thedistance betweeneyesabout1.6timesaslongasthediameterofaneyeinmale,1.8timesinfe- male.Frontmarginofclypeusstraight.Antennaemoderateinlength,reachingabout proximal1/5ofelytra,broadestatapicalendof4thsegment;approximateratioofan- tennal segments as 2.5:1.6:1.0:3.6:2.8:2.8:2.7:2.7:2.5:2.1:3.1 inmale (n=2, mean)and2.6:1.7:1.0:3.5:3.l:2.9:2.9:2.7:2.6:2.3:3.4in female(n=2,mean). pronotumsemicircular,sparsely andfinely punctate,closely coveredwithlargeand shallowconcavitieswhichhaveasetaateachcenter;thedistancebetweenconcavities of thesame lengthas adiameter ofconcavity; PW/PL 15-2.0 (1.7) in male and 1.6-1.9(1.7)in female.Scutellumfinelypunctate.Elytraoval,broadestatthemiddle, withadeepanddistinctpuncturesituatedonposteriorendofeachsetalsocket,butthe punctureisshallowandindistinctinproximalarea;EL/EW14-1.5(1.4)inmaleand 356 HiroyukiYosHIToMI c Fig.5. Sixthto7thabdominalsternitesofE1odesspp.(above,male;below,female). - A,E.inornata LEWIS; B, 1. wl'fso川 PIc;C, 1.scapll/a''IsLEWIS;D, 1.e/eganssp nov;E, 1.A,of加fatNAKANE. (Scales:0.5mm) 1.3-1.5 (1.4) in female;EL/PL2.8-3.7 (3.1) inmaleand2.9-3.7 (3.2) in female; EW/PW l2-1.4(1.3) inmaleand13-1.4(1.3) in female; TL/EW l8-1.9(1.9) in male and 18-2.0(1.8) in female.Ventral surfaceofbodycloselycoveredwithfine punctures.Legsmoderateinlength. Male. Apicalmarginof7thabdominalsterniteconcave.Eighthtergitesemicir- cular,coveredwithminutespinesinapicalarea;8thsternitegingko-leafshaped,cov- ered with minute spines on apical margin; 9th tergite moderately sclerotize(i, but broadlymembranous inapicalandlateralareas,withapair ofstoutapodemes;9th sterniteoblong,withlongsetaeatapicalmargin. Tegmenbifid inaboutapical 1/3 (parameres),withroundedapices; lateralmarginsofparameresserrateinapical1/5 and3/5tobase.Penisabout1.0mminlength,sparselycoveredwithpuncturesinapi- cal 2/5,bifidinaboutapical1/5(parameroids),withroundedapices,withapairof smallhooksataboutapical2/3oflateralmarginsofparameroids;dorso-proxima1mar- gindeeplyconcave;ventro-proxima1margindeeplynotched. Female. Apicalmarginof7thabdominalstemitearcuate.Eighthtergitemod- eratelysclerotized,coveredwithfinepuncturesandminutespinesatapicalmargin;8th sterniteweaklysclerotized,coveredwithfinepunctures,withdeeplynotchedapical marginwhichiscoveredwithminutespines. 0vipositor moderatein length;stylus E/odesandSacodesofJapan 357 with twosetaeatapicalend;coxiteandbaculuslong, finelypunctate;approximate ratioofthelengthsofstylus,coxiteandbaculusas follows(n=2,mean): - 1.0:4.1: 139. . Measurements of adult. Male (n=20). TL: 2.8-3.8 (3.3)mm; EW: 1.5-2.0 Fig 6. E1odes lnornata LEWIS; A,head; B,metasternum;C,pronotum; D,ditto; E,caudalareaof elytron;F,ditto,seta. 358 HiroyukiYosHIToM F Fig. 7(onp 358). .aodesinornataLEWSI. - A-F:Malegenitalia;A,8thtergite;B,8thsternite;C, 9thtergite;D,9thsternite;E, tegmen;F,penis,dorsalaspect. - G-I:Femalegenitalia;G,8thter- gite;H,8thsternite;1,ovipositor.(Scales:0.5mm) (l.8)mm;PL:0.6-1.0(0.8)mm;EL:2.2-2.8(2.5)mm;PW:1.2-1.5(1.3)mm.Female (n=20). TL: 3.1-4.2 (3.7)mm; EW: 1.7-2.3 (2.0)mm;PL: 0.7-1.0 (0.9)mm; EL: 2.4-3.3(2.8)mm;PW:1.3-1.6(1.5)mm. M,ature larva. Dorsalsurfaceofbodycoveredwith veryminutesetae; lateral marginsofheacしthoraxandabdomenwithspinousshortsetae,butthesetaesituatedat Fig. 8 (onp. 359). E1odes ino・,nataLEWIS,mature larva. - A, Dorsal aspect (scale: 1.0mm). - B-F:Mouthparts(scale:0.lmm);B,labrum,ventralaspect;C leftmandible,ventralaspect;D,1eft maxillarypalpus,dorsalaspect; E,ditto,ventralaspect;F,hypopharynx. - G-J: Abdominalseg- ments(scale:0.5mm);G,8thtergite;H,8thsternite;I,9thtergite;J,9thsternite.