©EntomologicaFennica.29April2004 Redescription and biology of Limonia badia (Walker) (Diptera: Limoniidae) JaroslavStarý&JukkaSalmela Starý, J. & Salmela, J. 2004: Redescription and biology of Limonia badia (Walker)(Diptera:Limoniidae).—Entomol.Fennica15:41–47. Limoniabadia(Walker,1848),previouslyknownfromtheNearcticRegiononly, isredescribedfromFinland.Itssystematicpositionandbiologyarecurrentlyde- bated. Limonia indigenoides (Alexander, 1920), another Nearctic species, is synonymizedwithL.badia.Thespeciesisdistinguishedbyseveraldistinctive characters, such as the wing pattern and the structure of the male terminalia. Thesecharactersareillustratedforthefirsttime.Immaturestagesareprobably associatedwithdecayingwoodofaspen(PopulustremulaL.).Alistofspeciesof theTipuloideaassociatedwithL.badiainFinland–includingGnophomyiaach- eronAlexander,1950,aspeciesnewtoEurope–ispresented. JaroslavStarý,DepartmentofZoologyandAnthropology,FacultyofScience, Palacký University, tr. Svobody 26, CZ-77146 Olomouc, Czech Republic; E- mail:[email protected] JukkaSalmela,DepartmentofBio-andEnvironmentalSciences,P.O.Box35, FIN-40014UniversityofJyväskylä,Finland;E-mail:[email protected] Received18August2003,accepted21January2004 1.Introduction 2.Materialandmethods The genus Limonia Meigen, 1803 is relatively The material was collected with trunk-window richinspecies:atotalof65taxa(speciesandsub- traps in two localities, Sb (Savonia borealis): species)areknownfromthePalaearcticRegion Savonranta, central Finland, and Obb (Ostro- (Savchenkoetal.1992).Thelarvaeofthegenus bothnia):Rovaniemimunicipality,northernFin- inhabit a wide array of habitats, such as aqua- land, in 1996 (Martikainen 2001). Solution of tic/semiaquaticandterrestrialsubstrates(Brink- water,saltanddetergentwasusedasapreserva- mann1991), leafdetritus(Tjeder 1958), decay- tive in the traps, and later the material obtained ing wood (Beling 1886) and fungi (Lindner waspreservedinethanol. Typespecimenswere 1958). borrowed from The Natural History Museum, Duringaninvestigationofinsectmaterialcol- Department of Entomology, London, UK lectedbytrunk-windowtrapsinFinlandin1996, (BMNH)andOhioStateUniversity,Department some highly interesting species of Tipuloidea ofEntomology,MuseumofBiologicalDiversity, werediscovered(seesection4.2andAppendix). Columbus,USA(OSU). The material included Limonia badia (Walker, The morphological terminology follows 1848), a species new to the Palaearctic fauna. McAlpine (1981), except for notation of the Here,weredescribethespeciesandillustrateits veins, which essentially is in accordance with diagnosticfeatures. Hennig (1954). The outer radial field in the 42 Starý&Salmela (cid:127) ENTOMOL.FENNICAVol.15 Fig.1.Wingof Limoniabadia (Walker)(Fin- land,Savon- ranta).Photo: T.Nieminen. Limoniinae is interpreted as proposed by Starý 14-segmented, moderatein length, not reaching (1992),andtheveinsandtheirpartsparticipating tobaseofwing.Scapeandpedicelslightlydarker intheso-calledreticulationpatternarenamedac- than flagellomeres. Scape cylindrical, about 1.5 cordingly. length of pedicel, the latter short-oval to nearly spherical.Flagellomeressubovaltoelongate,be- comingpaler,narrowerandlongertowardsapex 3.Limoniabadia(Walker,1848) of antenna. Terminal segment longest, slightly constrictedatmidlength,orbeyondit,withlittle- LimnobiabadiaWalker,1848:46(description). distinct transversal suture at that place, so that Limonia (Limonia) badia: Alexander 1943: feebly separated segment 15 often distinguish- 300(diagnosis,note,key)(reprinted1966);Al- able.Longestverticilssomewhatmorethantwice exander1965:43(Nearcticcatalogue). lengthofrespectivesegments.Wholesurfaceof LimnobiaindigenoidesAlexander,1920:193 flagellum with dense, short, semierect pubes- (description).syn.nov. cence. Limonia(Limonia)indigenoides:Alexander, Thorax generally brown, somewhat shiny, 1962: 13 (faun. record); Alexander 1965: 43 partlygreyishpruinose,andvariouslypatterned (Nearctic catalogue, tentative synonymy with with paler or darker hues. Pronotum essentially badia). palebrown,withsparsesetaeinanteriorhalfme- Note:Otherrecordsunderbadiaprobablyre- dially,andwithanothergroupofsetaeatposterior fer to Dicranomyia (Dicranomyia) humidicola margin.Prescutumdividedincolourintothreear- OstenSacken,1860. eas by two diffused, longitudinal, yellowish Diagnosis.Generalcolorationbrown,some- brown streaks, marked with rows of setae. Me- what shiny, greyish pruinose in parts, patterned dianarea,representingbroadmedianstripe,gen- withpaleranddarkerhues.Antennamoderatein erallybrown,darkeranteriorly.Palecapillaryline length,verticilsonflagellomeresovertwicethe splitting median stripe sometimes apparent in lengthofrespectivesegments.Wingwithdistinct specimens preserved in ethanol (not so in those stigma,andbroadseamsalongmostoftheveins. dried from ethanol). Lateral areas (stripes) ap- VenationofgeneraltypeasinotherLimonia,Rs pearingalittlepaler,providedwithfairlydistinct angulated and spurred, discal cell widened dis- darkbrownpatchatlateralmarginofprescutum, tally.Femorawithnarrowringbeforetip.Tarsal just before wing base. Scutum distinctly darker claws slender, with only two teeth near base. onlobesthanmostofprescutum,dirtyyellowish Gonocoxite of male terminalia with broad ven- brown in middle. Both scutellum and post- tromesal lobe. Gonostylus comparatively long. scutellumbrown,darkestinmiddle.Pleuronpat- Paramerebroad,darklypigmented.Bodylength terned with various hues of brown, somewhat 8.5–14mm,winglength8–10mm. shinyandmoregreyishpruinosethanprescutum, Redescription (male). Head dark brown, with anepisternum, katepisternum and posterior mostlydull.Palpusandantennabrown,thelatter partofso-calledsubspiracularsclerite(astermed ENTOMOL.FENNICAVol.15 (cid:127) RedescriptionandbiologyofL.badia 43 Fig.2.Limoniabadia Walker(Finland, Savonranta).–a. Dis- talpartofmalehind tarsus,lateralview.– b.Maleterminalia,dor- salview.c.–Aedeagal complex,dorsalview. so by Starý 1992) representing darkest areas. terior margins. Proximal sternites with yellow Groupofsetaeonlaterotergitedorsally. colourmoreextended. Wing(Fig.1)tingedwithyellowish,withdis- Legsratherlong,generallyyellowtoyellow- tinct dark brown stigma and broad seams along ish brown. Coxae and trochanters pale yellow, mostofveins,mostpronouncedsoalongRs,Cu foreandmidcoxaedarkenedatbase.Femorayel- and on cross-veins. Insidesofcellswithoutany lowish,withnarrow,notespeciallydark,ringjust markings. End of Sc darkened. Seamalong Rs beforetip.Tibiaeyellowish,narrowlytippedwith 1 markedly diffused into larger clouds on both darker. Tarsi dark except for base of basitarsus. ends;distalcloudfusedwithstigma.Venationof The latter long, about three times as long as generaltypeasinotherLimonia,yetdifferingin tarsomere2,thisabouttwicelengththeoftarso- manydetailsfrommanyotherEuropeanspecies. mere3.Bothlasttarsomeressubequalinlength, Sc ratherlong,endingataboutmidlengthofRs. abouthalfthelengthoftarsomere3.Tarsalclaws 1 R (mostly termed cross-vein r or R) slightly (Fig. 2a) rather long and very slender, slightly 2+3 2 lesssoorabouttwiceaslongasR (distalsection shorterthanlasttarsomere,mostlysmooth,with 3 beyondattachmentofcross-veinrofwhatprevi- onlytwoslenderteethnearbase,alongerone,sit- ouslywastermedR throughitsentirecourse;or uatedmoredistally andalittleshorter thanhalf 1 R ,ordistalsectionofSc invariouspapersby lengthofclawbeyondit,andaminuteproximal 1+2 2 Alexander).Rsangulatedanddistinctly spurred one. proximally.Discalcellessentiallytrapezoidorir- Maleterminalia(Fig.2b–c):Tergite9gener- regularly pentagonal (in former case often with ally transversally elliptic, with shallow median anotherspurdirectedinsidediscalcellatdeflec- emargination at posterior margin enclosed by tionofM),wideneddistally,withbasalsection palebar.Gonocoxiteshortandstout,withbroad, 3 ofM (withindiscalcell)twiceaslongasm-m,or slightly darkened ventromesal lobe. Gonostylus 3 more,andoftenarched,sothatlowerdistalmar- comparativelylong,swollenatbase,thendrawn ginofdiscalcellobliquelyconcave.Halterdirty out into long beak, the latter stout and almost yellowish, comparatively short, slightly dark- straight proximally, gradually narrowed into enedatmidlengthofstem. slender obtuse tip curved caudally. Gonostylus Abdomenbrown,withtergitesyellowatpos- mostlydarklypigmented,exceptatbaseandtip, 44 Starý&Salmela (cid:127) ENTOMOL.FENNICAVol.15 and provided with very long setae in proximal fromethanolandmountedonpoints. half.Aedeagusratherbroad.Paramerebroadand Comparative material: Limonia indigena: comparatively short, somewhat sclerotized and USA, Pennsylvania, Westmoreland Co., 10 km conspicuously darkened distally, with subacute SLigonier,PowdermillNatureReserve(400m), tipdirectedoutwardly.Otherdetailsareevident 27.IX.1986,1(cid:2),1(cid:1)(C.W.Youngleg.etdet.);in fromFig.2b–c. coll.J.Starý,Olomouc. Description(female).Resemblesmaleingen- Distribution. Holarctic: Canada: Nova Sco- eral appearance and structure, but antenna tia; USA: Alaska, Maine; Finland. First record slightlyshorterthanthoseofmale.Wingvenation fromthePalaearcticRegion. and wing pattern identical in both sexes. Tarsal clawsmuchshorterthaninmale,aboutonethird length of last tarsomere, with two teeth shorter 4.Discussion thaninmale,otherwise,however,similarinposi- tionandrelativelength.Incontrasttomale,last 4.1.DistinctionbetweenL.badia tarsomere longer than penultimate, about 1.5 andotherLimoniaspecies lengthofthelatter.Femaleterminalia,notexam- inedindetail,withcercusgenerallyslender,sub- Limoniabadiawasredescribedfromalargese- equalinlengthtofemaletergite10. riesofspecimens,allteneralinvariousdegrees, Type material examined. Limonia badia: preservedinethanol,ofwhichafewweresubse- Holotype(cid:1):[Canada:NovaScotia];inBMNH. quentlydriedfromethanolandmountedoncard- Labels:“R”(hand-written,agreenrhomboidla- boardpoints.Bothdry-mountedtypespecimens bel) “Limnobia/ badia/ Walker./ (type).” (hand- examinedappearsomewhatdarker,moregreyish written)“Type”(printed,agreen-marginedcircu- pruinose,otherwise,however,they areidentical larlabel),“Identifiedasthetype/byE.A.Water- withtheFinnishmaterialinallexternalcharacters house.”(printed)“HOLOTYPE”(printed,ared- andinthestructureofthemaleterminalia(incase marginedcircularlabel),“HOLOTYPE/Limno- of indigenoides). As usual for Limonia, female bia/badia/FrancisWalker/1848(G.B.1960)/G. terminaliadonotprovidesignificantfeatures,yet W.Byers”(partlyhand-written,red).Thespeci- identityoftheothermaterialwiththetypeofL. menpinned;tipoftherightwingmissing,alllegs badiaisbeyondanydoubt. and both antennae broken off. Limonia indi- Byitswingpattern,shapeofthetarsalclaws genoides: Holotype (cid:2) [USA], Alaska, Katmai, anddetailsinthestructureofthemaleterminalia August 1917 (J.S Hine leg.); in OSU. Labels: (Figs. 1–2), L. badia takes up a somewhat iso- “Katmai/Alaska/Aug’17”(printed)“JasSHine/ latedsystematicpositionandcannotbeconfused Collector” (printed) “Holotype/ Limnobia indi- with any of its Palaearctic and Nearctic conge- genoides/C.P.Alexander”(speciesnamehand- ners.Itisdistinctiveinthatitswingpatternisex- written,red)(inOSU).Thespecimenpinned;left clusively associated with venation, leaving in- wing missing; left mid and hind legs and right sides of cells without any markings, and being forelegbrokenoff.Tipofabdomenwascleared represented, except for thestigma, by well-pro- in KOH by the present authors and placed in a nouncedseamsalongveinsonly.Thetarsalclaws microvialwithglycerine,pinnedwiththespeci- bearonlytwoteeth.Theseexternalcharactersare men. quitesufficientfordistinguishingthespecies. Other material examined (22(cid:2), 43(cid:1)). Fin- Basedonthedarklypigmentedparameresand land: Sb: Savonranta, 10.VIII.–4.IX.1996, 3 (cid:2), some other details in the structure of the male 4.IX.–2.X.1996, 15 (cid:2), 38 (cid:1); Obb: Rovaniemi terminalia, L. badia appears to be distantly re- mlk, 25.VII.–27.VIII.1996, 4 (cid:2), 5 (cid:1) (all P. latedtothePalaearcticL.macrostigma(Schum- Martikainenleg.);inFinnishMuseumofNatural mel,1829)andL.alpicola(Lackschewitz,1928), History,ZoologicalMuseum,Helsinki,incoll.J. inwhich,however,thewingpatternisextremely Salmela, Jyväskylä and in coll. J. Starý, Olo- reduced,thetarsalclawsbearatleastthreeteeth, mouc.Allspecimensoriginallypreservedineth- andtheparameresaremuchmoreslender,spine- anol,3(cid:2)dissectedonslides,3(cid:2)and3(cid:1)dried like. ENTOMOL.FENNICAVol.15 (cid:127) RedescriptionandbiologyofL.badia 45 Closer affinities of L. badia probably occur wahlgreni, including Gnophomyia acheron, a with the Nearctic L. indigena (Osten Sacken, speciesnewtoEurope,sofarknownfromWest 1860),includingthesubspeciesL.indigenajack- Siberia, Russian Far East and Japan (cf. Sav- soni(Alexander,1917)andL.indigenaloloensis chenkoetal.1992;forcompletelist,seeAppen- Alexander,1958.Thelatterspeciesissomewhat dix). similar to L. badia in general appearance, also sharingthereducednumberofteethonthetarsal Acknowledgements. We would like to thank Dr Petri claw,butthewingpatternismoreextensive.The Martikainen(UniversityofJoensuu,Finland)forthesam- plingofthematerial.DrPeterW.Kovarik(OhioStateUni- pattern consists of a broad crossband along the versity,Columbus,USA),MrNigelWyatt(TheNatural so-calledcord, andanother one(moreproximal History Museum, London, England) and Dr Pekka andbroken)formedbycloudsinthecellsR,M, Vilkamaa (University of Helsinki, Finland) were instru- CuandA.Theparameresaredarkened,butslen- mentalinloaningtheimportanttypematerial.Mr.Teemu der, rathe1r ribbon-shaped, strongly curved and Nieminenkindlytookthephotographofthewing.Thanks to Dr. Hanna Kokko for correcting the English of the truncateatapex. manuscript.Theworkwassupportedbyagrantforthese- niorauthorfromtheGrantAgencyoftheCzechRepublic (No.206/01/1437).WealsothankDrPjotrOosterbroek 4.2.BiologyofL.badia andDrPekkaVilkamaafortheirhelpfulcomments. TheFinnishspecimensofL.badiawerecollected References withtrunk-windowtrapsplacedondeadaspens (Populus tremula L.) in two localities, namely Sb: Savonranta (62°15’N, 29°00’E) and Obb: Alexander,C.P.1920:ScientificresultsoftheKatmaiEx- pedition of the National Geographic Society: The Rovaniemi (66°10’N, 25°55’E) (for details, see crane-flies(Tipulidae,Diptera).—OhioJour.Sci.20: Martikainen 2001). The studied trees were in 193–203. clear-cuts(openhabitat)orinold-growthforests Alexander,C.P.1943:FamilyTipulidae.—In:Crampton, (shadedhabitat),eitherrecentlydiedormoreor G.C.,Curran,C.H.&Alexander,C.P.(eds.),The lessdecayed(barkpartly loose).Only treesand Diptera or true flies of Connecticut. First fascicle: 196–486.Conn.StateGeol.AndNat.Hist.Surv.,Bull. snagswithadiameteratbreastheightof>25cm 64.509pp.[reprintedin1966]. wereselectedforsampling.Thesamplingperiod Alexander,C.P.1962:ThecranefliesofMaine.—Univ. was from21 May to 2 October 1996 in Savon- Maine,Agr.Exp.Stat.Bul.T4,Technicalseries.24 ranta,andfrom28Mayto12September1996in pp. Rovaniemi. Both localities, particularly Savon- Alexander,C.P.1965:FamilyTipulidae.—In:Stone,A., Sabrosky, C. W., Wirth, W. W., Foote, R. H. & ranta, areaspen-richstandsandharbour several Coulson,J.R.(eds.),AcatalogoftheDipteraofAmer- rareandthreatenedbeetlesandpolyporefungias- icanorthofMexico:16–90.U.S.DepartmentofAgri- sociated with aspen (Martikainen et al. 2000, culture, Washington D.C., Agriculture Handbook Martikainen2001). 276.iv+1696pp. OurdataindicatethatL.badiaisanautumnal Alexander,C.P.1966:FamilyTipulidae.—In:Crampton, G.C.,Curran,C.H.&Alexander,C.P.(eds.),The species,sincethemajorityofthespecimenswere Diptera or true flies of Connecticut. First fascicle: caughtinSeptember.Virtuallyallcollectedspeci- 196–486.Conn.StateGeol.andNat.Hist.Surv.,Bull. menswereteneral,suggestingthatlarvalhabitat 64.509pp. waslocatedclosetothetrap:thespeciesmaywell Beling, T. 1886: Dritter Beitrag zur Naturgeschichte beasaproxyliclimoniidlivingindeadanddecay- (Metamorphose)verschiedenerArtenausderFamilie derTipuliden.—Verh.zool.bot.Ges.Wien36:171– ingaspenwood.Larvaeofsomeothermembers 214. ofthegenuswerereportedtoinhabitrottenwood, Brinkmann, R. 1991: Zur Habitatpräferentz und Phäno- suchasLimoniamacrostigma(cf.Beling1886). logiederLimoniidae,TipulidaeundCylindrotomidae InadditiontoL.badia,somerareandpoorly- (Diptera) im Bereich eines norddeutschen Tief- knownTipuloideawerecollectedonthesameoc- landbaches.—Faun.-Ökol.Mitt.Suppl.11:1–155. Hennig,W.1954:FlügelgeäderundSystemderDipteren casion, such as Elephantomyia krivosheinae, unter Berücksichtigung der aus dem Mesozoikum Lipsothrix ecucullata,Ula bolitophila, Tipula beschriebenen Fossilien. — Beitr.Entomol. 4: 245– (Lunatipula) circumdata and T. (Pterelachisus) 388. 46 Starý&Salmela (cid:127) ENTOMOL.FENNICAVol.15 Lindner, E. 1958: Pilzbewohnende Limoniidenlarven withnotesontheirhabitatrequirementsandconserva- unter besonderer Berücksichtigung von Limonia tionimplications.—Karstenia40:79–92. quadrinotataMeigen(Diptera).—Tijdschr.Entomol. Savchenko,E.N.,Oosterbroek,P.&Starý,J.1992:Family 101:263–281. Limoniidae.—In:Soós,(cid:1).,Papp,L.&Oosterbroek, McAlpine, J. F. 1981: Morphology and terminology – P.(eds.),CatalogueofthePalaearcticDiptera:183– adults.—In:McAlpine,J.F.,Peterson,B.V.,Shewell, 374.HungarianNaturalHistoryMuseum,Budapest1. G.E.,Teskey,H.J.,Vockeroth,J.R.&Wood,D.M. 463pp. (eds.),ManualofNearcticDiptera1:9–63.Research Starý, J. 1992: Phylogeny and classification of Tipulo- Branch,AgricultureCanada,Ottawa,MonographNo. morpha,withspecialemphasisonthefamilyLimo- 27.vi+674pp. niidae.—ActaZool.Cracov.35:11–36. Martikainen,P.2001:Conservationofthreatenedsapro- Tjeder,B.1958:AsynopsisoftheSwedishTipulidae1. xylicbeetles:significanceofretainedaspenPopulus Subfam. Limoniinae: tribe Limoniini. — Opusc. tremulaonclearcutareas.—Ecol.Bull.49:205–218. Entomol.23:133–169. Martikainen,P.,Penttilä,R.,Kotiranta,H.&Miettinen,O. Walker,F.1848:Listofthespecimensofdipterousinsects 2000: New records of Funalia trogii, Perenniporia inthecollectionoftheBritishMuseum.—London1: tenuisandPolyporuspseudobetulinusfromFinland, 1–229. ENTOMOL.FENNICAVol.15 (cid:127) RedescriptionandbiologyofL.badia 47 Appendix.AlistofspeciesofTipuloideacollectedwithtrunk-windowtrapsatSavonrantaandRovaniemimlk 1996,P.Martikainenleg.*=<10ex.;**=11–20ex;***=>20ex. Species Savonranta Rovaniemimlk Limoniidae Austrolimnophila(Archilimnophila)unica(Osten-Sacken,1869) * – Euphylidoreaphaeostigma(Schummel,1829) * – Phylidorea(Paraphylidorea)fulvonervosa(Schummel,1829) ** – Erioptera(E.)sordidaZetterstedt,1838 * – GnophomyiaacheronAlexander,1950 * – Ormosia(O.)ruficauda(Zetterstedt,1838) * – Dicranomyia(D.)autumnalis(Staeger,1840) * – Dicranomyia(D.)didyma(Meigen,1804) – * Dicranomyia(D.)handlirschiLackschewitz,1928 * – Dicranomyia(D.)modesta(Meigen,1818) – * Dicranomyia(Idiopyga)stigmatica(Meigen,1830) – * Discobolaannulata(Linnaeus,1758) *** – Discobolacaesarea(Osten-Sacken,1854) * – ElephantomyiakrivosheinaeSavchenko,1976 * – LipsothrixecucullataEdwards,1938 * – Neolimoniadumetorum(Meigen,1804) ** – Metalimnobiaquadrimaculata(Linnaeus,1761) ** – Metalimnobiazetterstedti(Tjeder,1968) – * Limoniabadia(Walker,1848) *** ** Limoniasylvicola(Schummel,1829) * – Rhipidia(R.)maculataMeigen,1818 ** * Rhipidia(R.)uniseriataSchiner,1864 * * Pediciidae – – UlabolitophilaLoew,1869 * – UlamixtaStar(cid:2),1983 * – Ulasylvatica(Meigen,1818) ** – Tipulidae – – Tipula(Lunatipula)circumdataSiebke,1863 – *** Tipula(Lunatipula)limitataSchummel,1833 – * Tipula(Pterelachisus)irrorataMacquart,1826 * – Tipula(Pterelachisus)wahlgreniLackschewitz,1925 * – Tipula(Savtshenkia)limbataZetterstedt,1838 * * Tipula(Savtshenkia)signataStaeger,1840 * – Tipula(Schummelia)variicornisSchummel,1833 * – Tipula(Tipula)paludosaMeigen,1830 * – Tipula(Vestiplex)nubeculosaMeigen,1804 * – Tipula(Vestiplex)scriptaMeigen,1830 * * Cylindrotomidae – – Triogmatrisulcata(Schummel,1829) * –