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Diversity2022,14,532.https://doi.org/10.3390/d14070532www.mdpi.com/journal/diversity
Article
What’sInsidetheHole?AReviewofEuropeanDendrolimnetic
MothFlies(Diptera:Psychodidae:Psychodinae)
SantiagoJaume‐Schinkel
1,
*,AlessioMorelli
2
,GunnarMikalsenKvifte
3,
*andXimoMengual
1
1
ZoologischesForschungsmuseumAlexanderKoenig,Leibniz‐InstitutzurAnalysedes
Biodiversitätswandels,Adenauerallee127,D‐53113Bonn,Germany;x.mengual@leibniz‐lib.de
2
ViaMartiriUngheresi22,I‐65019Pianella,Italy;alessiomorelli89@alice.it
3
DepartmentofBiosciencesandAquaculture,NordUniversity,P.O.Box2501,N‐7729Steinkjer,Norway
*Correspondence:s.jaume@leibniz‐lib.de(S.J.‐S.);gunnar.mikalsen‐kvifte@nord.no(G.M.K.)
Abstract:Weconductedanextensiveliteraturereviewinsearchofrecordsofdendrolimnetic
Psychodinae,withadditionalfieldsamplingoftheEuropeanspeciesofPsychodinaeassociated
withwater‐filledtreeholes.Aftercheckingmorethan100publications,only11specificpublished
recordsinvolvingdendrolimneticPsychodinaewerefound.Ourresultsshowthatsixgenera,rep‐
resentedby13speciesofPsychodinae,areassociatedwith13speciesofplanttrees.Asaresultof
ourfieldsampling,wereportLepiseodinalatipennis(Sarà,1953)andTelmatoscopusbartai(Ježek,2004)
comb.nov.forthefirsttimeinGermany.Furthermore,weredescribeL.latipennisbasedonfreshly
collectedmaterialwithacloserexaminationoftheholotype.Derivedfromourfindings,wereview
thegeneraLepiseodinaEnderlein,1937andTelmatoscopusEaton,1904,providinganidentification
keyforthemalesofbothgenera.Inaddition,wesynonymyzeKrivosheinoscopusJežek,2001syn.
nov.underTelmatoscopus,changingcombinationofTelmatoscopusussuricus(Ježek,2001)comb.nov.
andTelmatoscopusbartai(Ježek,2004),additionally,wechangecombinationandasononymyof
Tematoscopuswagneri(Salmanna,1982)comb.etsyn.nov.underTelmatoscopusadvena(Eaton,1893).
Furthermore,wedescribeforthefirsttimethefemaleandeggsofTelmatoscopusadvena.Moreover,
weprovidethefirstpublishedDNAbarcodes(COI)forTelmatoscopusbartai,Lepiseodinalatipennis
(Sarà,1953),Lepiseodinarothschildi(Eaton,1912),andLepiseodinatristis(Meigen,1830).Finally,we
alsodiscussthetaxonomyandecologyoftheEuropeandendrolimneticspeciesofthesubfamily
Psychodinae.
Keywords:water‐filledtreeholes;DNAbarcoding;mothflies;newtaxa;integrativetaxonomy
1.Introduction
Phytotelmata(singularphytotelma)derivesfromtheGreekwords“phyto”,which
meansplant,and“telma”,meaningpond,andwasfirstproposedbyVarga[1],basedon
hisobservations,todefinethemicrohabitatsproductofwater‐filledbodiesinplantsur‐
faces[2].Withinthephytotelmata,wecanfindthedendrotelmata(singulardendrotel‐
ma),derivingfromtheGreekwords“dendron”,meaningtree,and“telma”,whichcir‐
cumscribethebodywatersonlytowater‐filledtreeholes.Thesedendrotelmataarecon‐
sideredanintegralmicrohabitatinsideforestecosystems,astheyprovideasuitable
spacefordevelopment,prey,andwatersourcesformanyorganismsrangingfromin‐
vertebratestovertebrates[3,4].Speciesthatrelatetoorinhabitthesewater‐filledtree
holesareknownasdendrolimnetic,fromtheGreekwords“dendron”,and“limnḗtēs”,
whichmeansrelatingtoorinhabitingtheopenwaterofabodyoffreshwater.
Dendrotelmataarecommonlyfoundinoldtrees,deadoralive,andareaconsidered
crucialcomponentofecosystems,especiallywiththemodernforestrymanagement
wherethenumberofmatureandover‐maturetrees(commonlyreferredasoldtrees)has
decreased[5].Themainsourcesofnutrientsflowinginthetreeholesconsistofleaflitter
Citation:Jaume‐Schinkel,S.;Morelli,
A.;Kvifte,G.M.;Mengual,X.What’s
InsidetheHole?AReviewof
EuropeanDendrolimneticMoth
Flies(Diptera:Psychodidae:
Psychodinae).Diversity2022,14,532.
https://doi.org/10.3390/d14070532
AcademicEditor:SimoneFattorini
Received:30May2022
Accepted:24June2022
Published:30June2022
Publisher’sNote:MDPIstays
neutralwithregardtojurisdictional
claimsinpublishedmapsand
institutionalaffiliations.
Copyright:©2022bytheauthors.
LicenseeMDPI,Basel,Switzerland.
Thisarticleisanopenaccessarticle
distributedunderthetermsand
conditionsoftheCreativeCommons
Attribution(CCBY)license
(https://creativecommons.org/license
s/by/4.0/).
Diversity2022,14,5322of36
andarthropodcadavers,whilethequalityandcompositionofthesenutrientsvaryacross
speciesandhabitats[6,7].Thefactthatthesemicrohabitatshaveasmallsize,discrete
boundaries,andarenaturallyreplicatedinnature,makesthemattractiveforstudiesof
communitystructureandfunctionalities[7].Researchershavereportedtheusageof
dendrotelmatabyamphibiansduringdevelopment,asawatersourceforreptilesand
smallmammals,and,asbathingsitesforbirdsandbats[4].Nonetheless,mainlyinver‐
tebrateswithaquaticdevelopmenthavebeenreporteddevelopinginsidethesemicro‐
habitats,withsomereportsofotherinvertebratesthatusethemasawatersource[4,8].
Amongthem,themostcommonorganismsfoundinsideareimmaturestagesofDiptera
andColeoptera[7–10].
Mothflies(Insecta:Diptera:Psychodidae)arecommonlyfoundinwaterbodies,as
themajorityofpsychodidspeciesdevelopinwaterduringlarvalstages,withafewex‐
ceptionsthatdevelopinsoil,dung,orfungi[11–13].Therearesixsubfamiliesrecognized
worldwide,namelyBruchomyiinae,Horaiellinae,Phlebotominae,Psychodinae,Sy‐
coracinae,andTrichomyiinae;allofthemexceptHoraiellinaearepresentinEurope
[14,15].SpeciesinthesubfamilyTrichomyiinaedevelopinsidetreeholesorrottingwood
andtheirlarvaeareconsideredxylophagous(fromGreek“xylon”,meaningwood,and
“faguein”,meaningeating)[5].SpeciesofBruchomyiinaeandPhlebotominaehavebeen
reporteddevelopingonthegroundandleaflitterfeedingondecayingorganicmatter
[15].ThelarvalstagesofSycoracinaespecieshavebeenfounddevelopinginaquatic
mossesandleaflitter[15].Onthecontrary,speciesofthesubfamilyPsychodinaeare
commonlyreporteddevelopinginwaterbodiessuchaspondsandstreams,butthereare
afewgenerapresentinEuropewhoselarvaedevelopinsideoftreeholes,namelyClog‐
mia,Clytocerus,Lepiseodina,Pneumia,Psychoda,andTelmatoscopus[13,16–25].Althoughthe
larvaldevelopmentandhabitatforsomeEuropeanspeciesareknown,themothflies
associatedwithwater‐filledtreeholeshavebeenpoorlystudiedinEurope[25].
Inthepresentstudy,werevisetheavailableliteratureabouttheEuropeanspeciesof
thesubfamilyPsychodinaeknowntodevelopindendrotelmata,withaspecialemphasis
onthegeneraLepiseodinaandTelmatoscopus.Additionally,westudythedendrolimnetic
mothfliescollectedinGermanyresultingfromtheGermanBarcodeofLife(GBOL)pro‐
ject[26](www.bolgermany.de,accessedon02April2022).Asanoutcome,weredescribe
Lepiseodinalatipennis(Sarà,1953)fromGermany(Nordrhein‐Westfalen)andItaly(Sicily)
basedonnewmorphologicalandmoleculardata,andwesynonymizeKrivosheinoscopus
Ježek,2001syn.nov.underTelmatoscopusEaton,1904.Furthermore,wechangecombi‐
nationandsynonymizeTelmatoscopuswagneri(Salamanna,1982)comb.nov.etsyn.nov.
underTelmatoscopusadvena(Eaton,1893),andwedescribeforthefirsttimethefemale
andeggsofTelmatoscopusadvena(Eaton,1893)basedonmorphologicalandmolecular
data.Moreover,weprovidethefirstrecordofLepiseodinalatipennis(Sarà,1953)andTel‐
matoscopusbartaiJežek2004comb.nov.fromGermany.Furthermore,weprovidethefirst
COI(5′‐endofthecytochromecoxidasesubunitI)sequences,alsoknownasDNAbar‐
code,forTelmatoscopusbartai(Ježek,2001)comb.nov.,Lepiseodinalatipennis(Sarà,1953),
Lepiseodinarothschildi(Eaton,1912),andLepiseodinatristis(Meigen,1830).
2.Materialandmethods
2.1.GeographicScope
WefollowtheproposedgeographicboundariesofEuropebydeJongetal.[27]with
thefollowinglimits:East:Ural(E60°),West:AtlanticOcean(W30°),South:Mediterra‐
nean(N35°),andNorth:AtlanticIslands(N82°).
2.2.LiteratureRecords
Literaturesearchwasconductedbytrackingreferencesfromknownliteraturewith
thehelpofsearchengines(e.g.,www.scholar.google.com,accessedon5Jaunary2022)
andscientificdatabases(e.g.,www.jstor.org,www.scopus.com,www.webofscience.com,
Diversity2022,14,5323of36
5January2022)usingthesearchkeywords“Psychodidae,Psychodinae,dendrolimnetic,
Diptera,water‐filledtreeholes”.Literatureusedforthestudyencompassesrecordssince
thedescriptionofthespeciestothemostrecentpublishedworksuntilthebeginningof
2022,focusingonPsychodinaeanddendrolimneticstudies.Morethan100itemswere
analyzed;however,only11includedrecordsofdendrolimneticPsychodinae(aslistedin
Table1).
2.3.Sampling
SpecimenswerecollectedusingMalaisetrapsduringtheyears2013–2021aspartof
theGermanBarcodeofLife(GBOL)project[26](www.bolgermany.de,accessedon2
April2022).Specimenswerepreservedin96%ethanolandstoredat−20°Cuntilthey
weredissectedforDNAextractionandpreparingpermanentslides.Allsampledspeci‐
mensarestoredattheZoologischesForschungsmuseumAlexanderKoenig,Bonn,Ger‐
many(ZFMK).Furthermaterialwasbredfromorganicmattersampledfromden‐
drotelmata,and,occasionallycollectedwithaMalaisetrapanddirectlycollectedinItaly,
during2010–2022(AM).
Additionalexaminedmaterialisdepositedinthefollowingnaturalhistorycollec‐
tionsmentionedinthetextusingtheiracronyms:
AM:AlessioMorelliprivatecollection,Pianella(PE),Italy;latertobedepositedatthe
NaturalHistoryMuseumofGenova.
BNHM:BritishNationalHistoryMuseum,London,UnitedKingdom.
MNGD:MuseodiStoriaNaturaleGiacomoDoria,ComunediGenova,Genova,Italy.
NMP:NárodníMuzeum,Prague,CzechRepublic.
ZFMK:ZoologischesForschungsmuseumAlexanderKoenig,Bonn,Germany.
2.4.StudyoftheCollectedMaterialandTerminology
Afterlysis(seeGeneticsbelow),thespecimenswereclearedusingNaOH10%,dis‐
sected,andpermanentlymountedusingEuparal(WaldeckGmbH&Co.KG,Division
Chroma,HavixbeckerStraße,Münster,Germany)asmountingmedium,followingthe
proceduredetailedinIbáñez‐Bernal[28],withthemodificationthatpriortothediaph‐
anizationprocess,weperformedthedissectionofthehead,wings,andterminalab‐
dominalsegmentstomaceratetheminNaOH10%andcontinuewiththeprocedure,
whilepreservingtheremainingtissue(thorax,legs,firstabdominalsegments)inethanol
forposteriorDNAextraction.AdditionalmaterialwasmaceratedinKOH10%,trans‐
ferredinaceticacid10%,anddehydratedinacetone99%.Specimensweredissectedin
cloveoilandpartsmountedonmicroscopeslideswithCanadabalm.Fortheadditional
material,somespecimenswerephotographedtokeepaqualityimageofthehabitus.
WefollowthegeneralterminologyproposedbyCummingetal.andKvifteand
Wagner[15,29],exceptforthemaleterminaliathatweusetheterm“hypopods”forthe
posteriorgenitalicappendages,whichhavebeentreatedintheliteratureascercopodiaor
surstylioriginatinginthe9thabdominalsegment,or10thsegment,oracombinationof
both,asproposedbyKvifteandWagner[30].ForthefemaleTerminaliawefollowKo‐
trba[31].EggterminologyfollowsDeAlmeidaetal.[32].
Intheexaminedmaterialofeachspecies,acodeisprovidedforeachexamined
specimen(e.g.,ZFMK‐DIP‐000852020),andallthelabelinformationisfoundasSup‐
plementaryMaterial(TableS1).
2.5.Genetics
SpecimenswereprocessedattheZFMK,wherelysisandPCRwereperformedfol‐
lowingtheprotocolandprimersby[33,34].AfterthePCR,samplesweresenttoBeijing
GenomicsInstitute(BGI)forbidirectionalsequencing.Rawdatawerecuratedmanually
usingGeneious(v.7.1.9).FinalCOIsequenceswere658bplong.Allsequenceswillbe
Diversity2022,14,5324of36
publiclyavailableatwww.bolgermany.de.BOLDandgenebankaccessionIDscanbe
foundintheSupplementaryMaterial.
3.Results
Basedonourliteraturereview,wereport13speciesofPsychodinaeassociatedwith
dendrotelmatabelongingtosixgenera(Table1),namelyClogmia[C.albipunctata(Willis‐
ton,1893)],Clytocerus[C.xylophilusVaillant,1983],Lepiseodina[L.latipennis(Sarà,1953),L.
rothschildi(Eaton,1912)andL.tristis(Meigen,1830)],Pneumia[P.canescens(Meigen,1804)
andP.trivialis(Eaton,1893)],Psychoda[P.alternataSay,1824,P.cinereaBanks,1894andP.
minutaBanks,1894]andTelmatoscopus[T.advena(Eaton,1893),T.bartai(Ježek,2004)
comb.nov.,andT.thuringicusBeran,Doczkal,Pfister&Wagner,2010].Additionally,we
listTelmatoscopusbartai(Jezek,2004)comb.nov.asapotentialdendrolimneticspecies,
givingargumentsforthisdecision.
Table1.DendrolimneticPsychodinaetaxareportedforEurope,treespecieswhereitwasreported
andreferences.Doubleasterisk(**)denotatesanewrecordoftreespeciesforthePsychodinae
species.
TaxonTreeSpeciesReference
Clogmiaalbipunctata(Williston,1893)Oak(Quercussp.),notspecified[35,36]
ClytocerusxylophilusVaillant,1983Limetree(Tiliasp.)[37]
Lepiseodinatristis(Meigen,1830)
Ash(Fraxinussp.),beech(Fagussp.),birch(Betulasp.),
cherry(Prunussp.),elm(Ulmussp.),maple(Acersp.),oak
(Quercussp.),mulberry(Morussp.),lime(Tiliasp.),not
specified,**PopulusnigraL.
[13,22,25,36,38]
Lepiseodinarothschildi(Eaton,1912)Maple(Acersp.),oak(Quercussp.),notspecified[9,13,38]
Lepiseodinalatipennis(Sarà,1953)**Mapletree(Acersp.),**Oak(Quercussp.)
Pneumiacanescens(Meigen,1804)Notspecified[18,25]
Pneumiatrivialis(Eaton,1893)Notspecified[22,25]
PsychodaalternataSay,1824Notspecified[23]
PsychodacinereaBanks,1894Apple(Malussp.),oak(Quercussp.),**Hornbeam
(CarpinusorOstryasp.)[25]
PsychodaminutaBanks,1894Maple(Acersp.),oak(Quercussp.)[25]
Telmatoscopusadvena(Eaton,1893)Ash(Fraxinussp.),birch(Betulasp.),elm(Ulmussp.),oak
(Quercussp.),sycamore(Platanussp.)[13,22,25,38]
TelmatoscopusthuringicusBeran,
Doczkal,Pfister&Wagner,2010Notspecified(assumptionofdevelopment)[5]
TelmatoscopuslaurenceiFreeman,1953Limetree(Tiliasp.)[39]
3.1.KeytotheMalesofEuropeanPsychodinaeGeneraFoundinDendrotelmata
Differentialdiagnosis.AdultsofthesubfamilyPsychodinaecanbeeasilydifferen‐
tiatedfromtheadultsoftheexclusivelyxylophagoussubfamilyTrichomyiinae,which
canalsobefoundintreeholes,bythepresenceofaneyebridgeinPsychodinae(absentin
Trichomyiinae)andwingveinRwithfivebranchesinPsychodinae,withtwolongitu‐
dinalveinsbetweenradialandmedialforks(veinRwithfourbranchesinTrichomyiinae,
withonelongitudinalveinbetweenradialandmedialforks).
1. Antennawithatleastflagellomeres2–10nodiform,dividedintoabasalnodanda
distalneck(Figure1B,D)…3
‐ Antennawithflagellomerescylindricalorfusiform(Figure1A,C)…2
2. Cornicula(evertedsac‐shapedstructuresonthebacksurfaceofthehead(pre‐
sumedscentorgans);alsoknownaspatagia)usuallypresent;antennalscapemore
Diversity2022,14,5325of36
thanthreetimeslongerthanpedicel;flagellomere1withadistalbrushofwavy
setae(Figure1C)…ClytocerusEaton,1904
‐ Corniculaalwaysabsent;antennalscapelessthanthreetimesthelengthofpedicel;
flagellomere1withoutadistalbrushofwavysetae(Figure1A)…PneumiaEnder‐
lein,1935
3. Ascoidswithanteriorandposteriorbranches(ascoidsY‐shaped)(Figure1D);hy‐
popodsofmaleswithonlyoneapicaltenaculum…PsychodaLatreille,1797
‐ Ascoidswithasinglecurvedbranchorcarryingonlyanteriorbranches(ascoids
digitiformandnotY‐shaped)(Figure1B);hypopodsofmaleswithfourormore
apicaltenacula…4
4. Ascoidsbifurcate(asinKvifte&Wagner[15],Figure16;alsoinIbáñez‐Bernal[40],
Figures55and56);tenaculadistallyknife‐shapedandshorterthanbasalwidthof
hypopods(asinIbáñez‐Bernal[40],Figure58)…ClogmiaEnderlein,1937
‐ Ascoidswithasingledigitiformorleaf‐shapedbranch,notbifurcate(Figure1B);
tenaculadistallyfeatheredandlongerthanthebasalwidthofhypopods(Figures
2B,3B,4D,5B,6B,7B,8A,C,9A–C,10Band11B)…5
5. Aedeagusasymmetrical,withejaculatoryapodemebroaderthandistalelements
(Figures3C,4D,5C,6Band7C);ascoidsdigitiformorS‐shaped…Lepiseodina
Enderlein,1936
‐ Aedeagussymmetrical,withejaculatoryapodemenarrowerthandistalelements
(Figures12Cand13C);ascoidsleaf‐shapedordigitiform…TelmatoscopusEaton,
1904
Diversity2022,14,5326of36
Figure1.Firstantennalsegments(leftantenna)of:(A).Pneumiatrivialis(Eaton,1893),(B)Telmato‐
scopusadvena(Eaton,1893),(C).Clytocerusocellaris(Meigen,1818),(D).Psychodasp.Abbreviations:
asc=ascoids,flm=flagellomere,ped=pedicel,scp=scape.Scale(A–C)inmillimeters(mm).
Diversity2022,14,5327of36
Figure2.Lepiseodinalatipennis(Sarà1953),malegenitaliaadaptedfromtheoriginaldescriptionby
Sarà(1953).(A).Gonocoxitesandgonostyli.(B).Hypopodsandhypoproct.Abbreviations:gnx=
gonocoxite,gns=gonostyli,hpd=hypopod,epi=epiproct,ten=tenacula.Noscaleavailablebased
onoriginaldrawing.
Diversity2022,14,5328of36
Figure3.Lepiseodinarothschildi(Eaton,1912),malegenitalia.(A).Hypandrium,gonocoxitesand
gonostylus.(B)Epandrium,hypopods,tenacula,hypoproct.(C)Aedeagus.Abbreviations:aed=
aedeagus,conplt=condyleplate‐like,eja=ejaculatoryapodeme,epi=epiproct,gnx=gonocoxite
gns=gonostylus,ten=tenacula,hyp=hypandrium.Scale(A–C)inmillimeters(mm).
Diversity2022,14,5329of36
Figure4.Lepiseodinalatipennis(Sarà,1953),male.(A)Head.(B)Wing.(C)Flagellomeresandas‐
coids.(D)Genitalia.Scale(A–D)inmillimeters(mm).
Diversity2022,14,53210of36
Figure5.Lepiseodinalatipennis(Sarà,1953),malegenitalia.(A)Hypandrium,gonocoxitesandgon‐
ostylus.(B)Epandrium,hypopods,tenacula,hypoproct.(C)Aedeagus.Abbreviations:aed=ae‐
deagus,epa=epandrium,eja=ejaculatoryapodeme,epi=epiproct,gnx=gonocoxite,gns=gono‐
stylus,ten=tenacula,hpd=hypopods,hyp=hypandrium.Scale(A–C)inmillimeters(mm).
Diversity2022,14,53211of36
Figure6.Lepiseodinalatipennis(Sarà,1953),male.SEMpictures.(A)Firstflagellomere(3rdantennal
segment),redcirclehighermagnificationofsensilla.(B)Genitalia,lightblue=hypandrium,orange
=aedeagalsheath,green=gonocoxites,purple=gonostyli,red=aedeagus,darkblue=hypopods,
yellow=tenacula.Scale(A,B)inmicrometers(μm).
Diversity2022,14,53212of36
Figure7.Lepiseodinatristis(Meigen,1830),malegenitalia.(A)Hypandrium,gonocoxitesandgon‐
ostylus.(B)Epandrium,hypopods,tenacula,hypoproct.(C)Aedeagus.Abbreviations:aed=ae‐
deagus,conplat=condylesplate‐like,epa=epandrium,eja=ejaculatoryapodeme,epi=epiproct,
gnx=gonocoxite,gns=gonostylus,ten=tenacula,hpd=hypopods,hyp=hypandrium.Scale(A–
C)inmillimeters(mm).
Diversity2022,14,53213of36
Figure8.Wingsvariabilityof:(A–D)Lepiseodinalatipennis(Sarà1953).(E,F)L.tristis(Meigen,1830),
(G,H)L.rothschildi(Eaton,1912).Scale(A–H)inmillimeters(mm).
Diversity2022,14,53214of36
Figure9.Genitaliaof:(A)Lepiseodinatristis(Meigen,1830),(B)Lepiseodinarothschildi(Eaton,1912),
(C)Lepiseodinalatipennis(Sarà,1953).Scale(A–C)inmillimeters(mm),scalelines=0.10mm.
Diversity2022,14,53215of36
Figure10.Genitaliaof:(A)Lepiseodinalatipennis(Sarà,1953),(B)Lepiseodinatristis(Meigen,1830).
Bluearrowsshowdifferencesinhypandrium,gonocoxite,andgonostyli.Scale(A,B)inmillimeters
(mm),scalelines=0.10mm.
Diversity2022,14,53216of36
Figure11.GenitaliavariabilityofLepiseodinalatipennis(Sarà,1953):(A–C)Aedeagalcomplexin
dorsalview.(D)sameinposterodorsalview.(E)HabitusofL.latipennis(Sarà,1953),(F)Habitusof
L.rothschildi(Eaton,1912).Scale(A–D)inmillimeters(mm);withoutscale(E,F).
Diversity2022,14,53217of36
Figure12.Telmatoscopusadvena(Eaton,1893),malegenitalia.(A)Hypandrium,gonocoxitesand
gonostylus.(B)Epandrium,hypopods,tenacula,hypoproct.(C)Aedeagus.Abbreviations:epa=
epandrium,epi=epiproct,eja=ejaculatoryapodeme,gnx=gonocoxitegns=gonostylus,hpd=
hypopod,ten=tenacula,hyp=hypandrium.Scale(A–C)inmillimeters(mm).
Diversity2022,14,53218of36
Figure13.Telmatoscopusbartai(Ježek,2004),malegenitalia.(A)Hypandrium,gonocoxitesand
gonostylus.(B)Epandrium,hypopods,tenacula,hypoproct.(C)Aedeagus.Abbreviations:aed=
aedeagus,epa=epandrium,epi=epiproct,eja=ejaculatoryapodeme,gnx=gonocoxites,hpd=
hypopod,hyp=hypandrium,ten=tenacula.Scale(A–C)inmillimeters(mm).
3.2.SystematicAssessment
GenusClogmiaEnderlein,1937
ClogmiaEnderlein,1937:87.Typespecies:PsychodaalbipennisWilliston(=albipunctata
Williston).
Diversity2022,14,53219of36
Diagnosis.Scapelessthan2.5timesitswidth;flagellomeressymmetricallynodiform,
withascoidsvariableinshape;flagellomere14withelongatedapiculustaperingto‐
wardsapex;wingwithradialforkbasaltomedianfork;ejaculatoryapodemeY‐shaped,
narrowindorsalview;aedeagussymmetricalindorsalview.
SpeciespresentinEuropeassociatedwithdendrotelmata:C.albipunctata(Williston,
1893)[35,36](Table1).
Clogmiaalbipunctata(Williston,1893)
PsychodaalbipunctataWilliston,1893:113.Typelocality:Cuba,LaHavana.
PericomameridionalisEaton,1894:194.Typelocality:EastAfrica.
PsychodasnowiiHaseman,1907:311.Typelocality:USA,Texas,Galveston.
PsychodalegnothisaSpieser,1909:44.Typelocality:Tanzania.
PsychodaerectaCurran,1926:102.Typelocality:WestIndies.
PschodanocturnaAbreu,1930:115.Typelocality:Notgiven,probablyCanaryIslands
(seeIbáñez‐Bernal[40]).
Psychodanocturnavar.nigrithoraxAbreu,1930:115.Typelocality:notgiven,probably
CanaryIslands.
TelmatoscopusharantiMirouse,1958:93.Typelocality:MididelaFrance.
Telmatoscopusalbipunctatus(Williston):[41](p.185).
Clogmiaalbipunctata(Williston):[42](p.42);351[11](p.351)(seeIbáñez‐Bernal[40]).
Diagnosis.Antennawithsymmetricallynodiformflagellomeres,eachflagellomere
withabifurcateascoids;eyesseparatedby1facetdiameter,eyebridgewith4facet
rows;aedeagussymmetrical,ejaculatoryapodemestraightandnarrowindorsalview;
hypopodswithfive‐sixtenacula,tenaculadecreasinginsizetowardstheapexofhy‐
popod.
Examinedmaterial:ZFMK‐DIP‐00081299,ZFMK‐DIP‐00081508,
ZFMK‐DIP‐00081514,ZFMK‐DIP‐00081543,ZFMK‐DIP‐00081624,ZFMK‐DIP‐00081591,
ZFMK‐DIP‐00082118[ZFMK].
Distribution.ThisisoneofthemostwidespreadspeciesofPsychodidaeinthe
world,highlyinvasiveandsynanthropic.InEurope,itisrecordedinAzoresArchipela‐
go,Belgium,CanaryIslands,Corsica,Croatia,CzechRepublic,Denmark,Finland,
France,Germany,Greece,Hungary,Italy,Luxemburg,MadeiraIslands,Sardinia,Slo‐
vakia,Slovenia,Spain,SwedenandUnitedKingdom[9,43–47].
GenusClytocerusEaton,1904
ClytocerusEaton,1904:59.Typespecies:Clytocerusocellaris(Meigen,1804:44)bysub‐
sequentmonotypyofMalloch(1907)(see[48]).
Diagnosis:Cornicula(evertedsac‐shapedstructuresonthebacksurfaceofthe
head)areusuallypresent;eyebridgewith3–6facetrows,separatedby1–3facetdiam‐
eters;antennalscapeelongate,three‐sixtimeslongerthanwide;flagellomereIandII
fusedtoformacompoundflagellomere,withadistalbrushofwavysetae[14].
SpeciespresentinEuropeassociatedwithdendrotelmata:C.xylophilusVaillant,
1983[37](Table1).
ClytocerusxylophilusVaillant,1983
ClytocerusxylophilusVaillant,1983:355.Typelocality:France.
Diagnosis.Eyesseparatedby2facetdiameters,interocularsutureabsent.Antennal
scape4.75timeslongerthanwide.Hypopodswith6–7tenacula.
Examinedmaterial.None.
Distribution.France[43].
Remarks.Vaillant[37]mentionsunderthedescriptionofC.xylophilusthatthelar‐
vaeofClytoceruspulvereusVaillant,1983canbefoundinthesamesubstrateasC.xy‐
lophilus;however,itisreferringtothe“blackhumus”foundnearariver(samesubstrate
Diversity2022,14,53220of36
mentionedbyKrek[49]),andlaterVaillant[37]clarifiesthatonlyC.xylophilusisknown
asadendrolimneticspecies,.
GenusLepiseodinaEnderlein,1937
LepiseodinaEnderlein,1937:91.Typespecies:PsychodatristisMeigen,1830:272.
Diagnosis.Fronsandclypeusseparatedandnotprotrudingovereyemargin;scape
short,lessthan2timeslengthofpedicel;flagellomeressymmetricallynodiformindorsal
view,withapairofdigitiformsinuousascoids;flagellomere14withelongatedapiculus;
wingveinR2+3notconnectedtoR4;apexofR5endingon,orbelowwingapex;gonocoxites
short,withdorsalprojectionundeveloped;ejaculatoryapodemenarrowindorsalview;
aedeagalcomplexasymmetric,withaparameralstructuremoreorlesssclerotized,con‐
nectingaedeagustothegonocoxalapodemes;hypopodswithindistinctlyfringedtenac‐
ula;epandriumwithasingleforamen.
SpeciespresentinEuropeassociatedwithdendrotelmata:L.rothschildi(Eaton,
1912),L.latipennis(Sarà,1953),andL.tristis(Meigen,1830)[9,13,23,25,36,38](Table1).
Lepiseodinalatipennis(Sarà,1953)
Figures2,4–6,7A,8,C,9C,10and11E.
TelmatoscopuslatipennisSarà,1953:13.Typelocality:Italy,Messina.
TelmatoscopuslatipennisSarà:[50](p.53).
Clogmialatipennis(Sarà):[51](p.60).
Lepiseodinalatipennis(Sarà):[52](p.146).
Diagnosis.Malehypandriumbroadalongitsentirelength;gonocoxalalveolidis‐
tributedintheentiregonocoxalsurface;gonostyliwithoutalveolionthebase,twicethe
lengthofgonocoxites;epandriumnotdivided,withasinglekidney‐shapedforamen;
aedeagusincurved;paramerenotstronglysclerotized;hypoprocttongue‐shaped.
Differentialdiagnosis.ThisspeciesiscloselyrelatedtoLepiseodinatristis,butitcan
bedistinguishedbythecombinationofthefollowingcharacters:hypandriumbroad
alongitsentirelengthwithconvexityinthebasalmargininL.latipennis(hypandrium
narrowwithamedialprojectioninthebasalmargininL.tristis);gonocoxalalveolidis‐
tributedintheentiregonocoxalsurfaceinL.latipennis(gonocoxalalveolirestrictedtothe
apicalmarginintwoorthreeirregularrowsinL.tristis);gonostyliwithoutalveolionthe
baseinL.latipennis(gonostyliwithalveoliinthebaseinL.tristis);epandriumnotdivided
intheapicalmargin,withakidney‐shapedforameninL.latipennis(epandriumdivided
intheapicalmarginwitharoundedforameninL.tristis);aedeaguscurvedinL.latipennis
(aedeagusstraightinL.tristis);parameresnotstronglysclerotizedandalmostnotvisible
inL.latipennis(parameresstronglysclerotizedinL.tristis).
LepiseodinalatipenniscanbedifferentiatedfromLepiseodinarothschildibythegono‐
stylitwicethelengthofgonocoxitesinL.latipennis(gonostylilessthanhalfofthelength
ofgonocoxiteinL.rothschildi),apicalmarginofhypandriumconvexinL.latipennis(apical
marginofhypandriumconcaveinL.rothschildi),andparameresnotstronglysclerotized
andalmostnotvisibleinL.latipennis(paramerestronglysclerotizedinL.rothschildi).
Redescription.Measurementsinmm(mean,SD=0.005,n=2).Winglength2.53
(2.20–3.30),wingwidth0.98(1.02–1.32).Headlength0.50(0.48–0.59),headwidth0.63
(0.53–0.65).Antennalsegments,scapelength:0.11(0.09–0.11),pedicel:0.08(0.70–0.90),
postpedicel:0.10(0.10–0.13),flagellomeresaveragelength0.11;Palpomeres1:0.10(0.08–
0.10),2:0.25(0.20–0.27),3:0.23(0.20–0.24),4:0.29(0.24–0.29).
Male.Head:eyebridgewithrowsoffourfacets(rarely3facets);frontalpatchun‐
divided,withanirregularrowofalveoliextendingtowardstheinterocularsuture,the
wholefrontalpatchtogetherwiththeirregularrowresembletheshapeofahandbell;
interocularsutureasaninverted“v”.Noalveoliofsupraocularsetaearepresent.Labella
bulbous,longerthanwide,with10–12smallsetae.Antennawithscapecylindrical,1.34
timeslongerthanitswidth,1.43(1.2–1.4)timesthelengthofpedicel;pedicelspherical;14
nodiformflagellomereswithbasalbulbanddistalneck,apicalflagellomerewithlong
Diversity2022,14,53221of36
apiculus,apiculusalmosthalfthelengthoftheflagellomere(Figure4A);antennalascoids
slightlyflattened,digitiform,sinous(S‐shaped),totallengthabout1.88thelengthof
flagellomeres;flagellomereswithasensilla(asshowninFigure6A).Palpalsegment
proportions1.0:2.63:2.37:3.03.
Thoraxwithnoparticularcharacters.
Wing:Lengthabout2.5(2–2.5)timesitswidth,variableinshape,withanterior
marginmoreorlesscurved;membranebareexceptonveins;hyalinewithaslightin‐
fuscationoncostalcell;Scstraight,endingatleveloftheoriginofR2+3;OriginofR2+3not
joiningR4,alittledistaltotheoriginofM1+2;originofM1+2broadandrounded;forksof
R2+3andM1+2almostatthesamelevel,theforkofR2+3beingslightlydistaltoM1+2;R5
endingatwingapex;CuAendinginwingmarginatthelevelofR2+3fork.
Abdomen.Withoutanyparticularcharacters.
Genitalia.Hypandriumplate‐like,broad,basalmarginwithmedialprojection
widelyrounded,distalmarginalmoststraight;gonocoxiteslongerthanwide,coveredin
alveolionalmostallthesurface;gonostyliabout1.5timesthelengthofgonocoxites,ta‐
peringtowardsapex,almoststraight;gonocoxalapodemespoorlydistinguishablein
dorsalview,inventralviewplate‐like,stronglysinuous,fusedandnarrowatlivelofthe
midline;parameresformingasingleslightsclerotizedstructure,pyriform,resembling
theupperhalfofabowlingpinconnectinglikeabridgeaedeagustogonocoxal
apodemes(asinFigures4D,5C,6B,9Cand10A);epandriumsubrectangular,abouttwice
widerthanlongwithasinglekidney‐shapedforamen;hypopodsslightlyoutcurved,
apexrounded,with9–13tenacula(asinFigures4B,5Cand9C);epiproctshort,triangular
andcoveredinpilosity;hypoproctbroadandtongue‐shaped,coveredinpilosity,ex‐
tendingtowardsmidofhypopods.Aedeaguswithejaculatoryapodemedigitiform,
narrow7.5timeslongerthanitswidth,about1.5timesthelengthofgonocoxites;dis‐
tiphallusaboutthesamelengthofejaculatoryapodeme,incurved,extendingtowardsthe
apexofgonostyli,bothejaculatoryapodemeanddistiphallusformasinglecomplex,
jointedwithparameralstructureandencirclesbyamembranousparameralsheath.
Female.Unknown.
Remarks.Theholotype(slidemounted)isquitedark,makingtheobservationof
structuresdifficult.Theheadisdissected,andquitedifficulttoseeclearly,oneantennais
dissectedandtheotherantennaismissing,onecompletepalpusisdissected,theother
palpusismissing.Thehypopodsaredissectedandplacedapartfromeachother,one
gonocoxitesandgonostylusaredissectedandplacedapartfromthegenitalia,there‐
mainingpartsofthegenitalia(aedeagalcomplex,onegonocoxitesandgonostylus)are
foundtogether.Onewingisdissectedandseparatedfromthethorax.Ontheoriginal
descriptionSarà(1953)mentionstheshapeofthehypoproctbeingtrilobed(Figure2),
afterexaminationoftheholotypeinthepreparationthehypoproctlooksindeedtrilobed;
however,weconsiderthisamalformationontheslideitself,andnotthenaturalshapeof
thehypoproct,allotherexaminedmaterialpresentabroadtongue‐shapedhypoproct.
OntheoriginaldescriptionSaràabstainedtoillustrateordescribetheaedeagalcomplex;
however,afterexaminationwearesureallourspecimensbelongtothesamespecies.
Biology.Somespecimens(No.0190‐0193)emergedfromorganicmattersampled
fromanAcersp.dendrotelma(Figure14),inamixedsubmontanforest,additionally,the
specimenusedfortheSEMpictureswascollectedinaMalaisetrapnexttodendrotel‐
mataofanoak(Quercussp.)tree,suggestingthatalsoL.latipennisisadendrolimnetic
species.InGermanyaltituderangesfrom250–280ma.s.l.(metersabovesealevel),while
inItalyithasarangefrom40–800ma.s.l.
Examinedmaterial.Holotypeexamined[MNGD],ZFMK‐DIP‐00081595,
ZFMK‐DIP‐00081552[ZFMK],SpecimenNo.0039,0488,0487,0439,0190,0191,0192,0193
[AM].
Distribution:PreviouslyknownonlyfromthetypelocalityinMessina(Sicily,Italy).
TherecordsherereportedarethefirstfromGermany(Rheinland‐Pfalz)andfornorthern
andcentralItaly.
Diversity2022,14,53222of36
Figure14.(A,B)HabitatofPsychodacinereaBanks,1894,dendrotelmaofhornbeam(Carpinusor
Ostryasp.).(C,D)HabitatofLepiseodinatristis(Meigen,1830),rottingtreeholewithbasalden‐
drotelmaofPopulusnigra.Allinamixedsubmontanforest(ForoValley,centralItaly).
Lepiseodinarothschildi(Eaton,1912)
Figures3,8G,Hand9B
TelmatoscopusrothschildiiEaton,1912:7.Typelocality:England,London,HydePark.
Diversity2022,14,53223of36
Telmatoscopusrotschildi(Eaton):Lapsuscalamiof[50],followedbySalamannainDahlet
al.[53](seebelow).
Clogmiarothschildi(Eaton)[ICZN(1999):art.33.3.1:incorrectsubsequentspellingin
prevailingusageisdeemedtobeacorrectoriginalspellingandmaintained;seeJežek
(2004)]:Vaillant(1982a):298;(1982b):206;Wagner(1990):60;Bernotienè(2002):7.
Clogmiarotschildi(Eaton):Dahletal.[53](p.33).
Lepiseodinarothschildi(Eaton):Ježek[52](p.146).
Diagnosis.Malegonostyliarehalfthelengthofgonocoxites;paramerestrongly
sclerotized,overlappingtothelateralbranchofaedeagus;hypandriumW‐shaped,broad
ontheentiresurfacewiththeapicalmarginconcave.
Examinedmaterial:ZFMK‐DIP‐00081311,ZFMK‐DIP‐00081322,
ZFMK‐DIP‐00081323,ZFMK‐DIP‐00081324,ZFMK‐DIP‐00081327,ZFMK‐DIP‐00081328,
ZFMK‐DIP‐00081329,ZFMK‐DIP‐00081330,ZFMK‐DIP‐00081510,ZFMK‐DIP‐00081537,
ZFMK‐DIP‐00081551,ZFMK‐DIP‐00081558,ZFMK‐DIP‐00081559,ZFMK‐DIP‐00081567,
ZFMK‐DIP‐00081574,ZFMK‐DIP‐00081623,ZFMK‐DIP‐00082122,ZFMK‐DIP‐00082126,
ZFMK‐DIP‐00082147,ZFMK‐DIP‐00082149[ZFMK].Specimensnumber:13666,13689,
13694,13625,13628,13822,18655,18490,18225,19261,19110,16352,16489,18408,19469,
17902,21769,12319,12320,21533,21114[NMP].Specimensnumber:0489[AM].
Distribution.Austria,Belgium,Bulgaria,CzechRepublic,Finland,France,Germa‐
ny,Ireland,Italy,Lithuania,Netherlands,Slovakia,Spain,UnitedKingdom[9,13,54,55].
Lepiseodinatristis(Meigen,1830)
Figures7,8E–F,9Aand10B
PsychodatristisMeigen,1830:272.Typelocality:Notspecified,probablyBelgiumor
Germany.
Psychoda(Pericoma)tristis(Meigen):[56](p.16).
Pericomatristis(Meigen):[57](p.17).
Telmatoscopustristis(Meigen):[58](p.170).
Lepiseodinatristis(Meigen):[42])(p.91);[52](p.146).
Clogmiatristis(Meigen):[53](p.33);[59](p.7).
Diagnosis.Malehypandriumnarrow,withasmallabruptmedianprojection;gon‐
ocoxalalveolirestrictedtotwoirregularlinesontheapicalsurface;gonocoxiteswitha
patchofalveoliatbase;twoparameresstronglysclerotized.
Examinedmaterial:ZFMK‐DIP‐00081512,ZFMK‐DIP‐00081513,
ZFMK‐DIP‐00081563,ZFMK‐DIP‐00081583,DIP‐ZFMK‐00081622,ZFMK‐DIP‐00082119,
ZFMK‐DIP‐00082124,ZFMK‐DIP‐00082125,ZFMK‐DIP‐00082128,ZFMK‐DIP‐00082130,
ZFMK‐DIP‐00082133,ZFMK‐TIS‐2628581,ZFMK‐TIS‐2628542,ZFMK‐TIS‐2628588,
ZFMK‐TIS‐2628587,ZFMK‐TIS‐2628608[ZFMK].Specimennumber:24339,13701,13668,
13633,13823,17331,17270,20777,20243,20077,20076,20080,20079,20088,20067,3226,
33282,34041,34042,34043,34044,20987,21365,23772,24339[NMP].Specimennumber:
0490,0491[AM].
Distribution.Algeria,Austria,Belgium,Croatia,CzechRepublic,France(incl.Cor‐
sica),Germany,Ireland,Italy,Lithuania,Slovakia,andUnitedKingdom
[25,36,43,50,53,59,60].InItaly,thespeciesisknownonlyfromtwooldanduncertain
recordsforthenorthernandpeninsularregion[53,61].Thespecimensherereported
confirmtheoccurrenceofthisspeciesintheItalianpeninsula.
KeytotheEuropeanadultmalesofLepiseodina
1. Gonostylilongerthangonocoxites…2
‐ Gonostyliabouthalfthelengthofgonocoxites(Figure3A)…L.rothschildi
2. Hypandriumnarrow,withasmallmedialprojectiononbasalmargin(Figure7A);
alveoliingonocoxitesrestrictedintwoorthreeirregularrowsintheapicalmargin
Diversity2022,14,53224of36
(Figures9Aand10B);gonostyliwithalveolionthebase(Figures9Aand10B);two
parameresstronglysclerotized(Figure7C)…L.tristis
‐ Hypandriumbroadinitsentirelength,withoutmedialprojection(Figure5A);al‐
veoliingonocoxitesscatteredinthewholesurface(Figure5A);gonostyliwithout
alveolionthebase(Figure5A);oneparameralstructurenotstronglysclerotized
(Figure5C)…L.latipennis
GenusPneumiaEnderlin,1935
PneumiaEnderlein,1935:247.Typespecies:PericomapalustrisMeigen,1804(see[62]).
Diagnosis.Eyebridgewith6facetrows.Antennalflagellomeresbarrel‐shaped,
basalflagellomereslackingspinesorclustersofstiffsetae;apicalflagellomerewitha
digitiformapicalprotuberanceaboutaslongasorlongerthanthebasalpartofthe
flagellomerecarryingit.Aedeagalcomplexsymmetrical;gonostylibulboseandelongate
atthebase,taperingtowardstheapex.
SpeciespresentinEuropeassociatedwithdendrotelmata:Pneumiacanescens
(Meigen,1818)andPneumiatrivialis(Eaton,1893)[18,23,25](Table1).
Pneumiacanescens(Meigen,1818)
TrichopteracanescensMeigen,1818:45.Typelocality:notgiven,probablyGermany.
Pericomacanescens(Meigen):20(p.156).
Pneumiacanescens(Meigen):[63](p.124).
Diagnosis.Eyebridgewith5–6facetrows:wingveinforkR2+3withabackward
projection(asin[11](Figure12));hypopodswith10tenacula;ejaculatoryapodeme
wideratthebasethanapex;aedeagalcomplexlongerthangonocoxites,taperingto‐
wardsapex.
Examinedmaterial.NMP‐2687,NMP‐2913,NMP‐10787,NMP‐17243[NMP].
Distribution:Afghanistan,Armenia,Austria,Azerbaijan,Belgium,Bulgaria,China,
CzechRepublic,Denmark,France,Georgia,Germany,Greece,Hungary,Kyrgyzstan,
Lithuania,Netherlands,Poland,Romania,Russia(NovosibirskRegion),Slovakia,Swe‐
den,TurkeyandUnitedKingdom[40,56,57].
Pneumiatrivialis(Eaton,1893)
PericomatrivialisEaton,1893:121.Typelocality:GreatBritain.
Pneumiatrivialis(Eaton):[23](p.202);[63](p.116).
Diagnosis.Eyebridgewith7facetrows;fronsnotextendingbeyondfirstpalpal
segment;allpalsegmentsofsimilarwidth;foretibiastraightandnotengrossed;wing
veinforkR2+3withoutabackwardprojection;hypopodswith7orfewertenacula;ejacu‐
latoryapodemestraight,notwideratbase;aedeagalcomplexshorterthangonocoxites,
nottaperingtowardsapex,distalpartofaedeagussmoothwithoutwrinkles.
Examinedmaterial.ZFMK‐DIP‐00081529,ZFMK‐DIP‐00081536,
ZFMK‐DIP‐00081541,ZFMK‐DIP‐00081576,ZFMK‐DIP‐00081601,ZFMK‐DIP‐00081602,
ZFMK‐DIP‐00081603,ZFMK‐DIP‐00081604,ZFMK‐DIP‐00081605,ZFMK‐DIP‐00081606,
ZFMK‐DIP‐00081607,ZFMK‐DIP‐00081610,ZFMK‐DIP‐00081999,ZFMK‐DIP‐00082001,
ZFMK‐DIP‐00082002,ZFMK‐DIP‐00082003,ZFMK‐DIP‐00082006,ZFMK‐DIP‐00082007,
ZFMK‐DIP‐00082008,ZFMK‐DIP‐00082009,ZFMK‐DIP‐00082010,ZFMK‐DIP‐00082011,
ZFMK‐DIP‐00082012,ZFMK‐DIP‐00082013,ZFMK‐DIP‐00082014,ZFMK‐DIP‐00082015,
ZFMK‐DIP‐00082016,ZFMK‐DIP‐00082017,ZFMK‐DIP‐00082029,ZFMK‐DIP‐00082031,
ZFMK‐DIP‐00082032,ZFMK‐DIP‐00082033,ZFMK‐DIP‐00082034,ZFMK‐DIP‐00082035,
ZFMK‐DIP‐00082036,ZFMK‐DIP‐00082037,ZFMK‐DIP‐00082039,ZFMK‐DIP‐00082040,
ZFMK‐DIP‐00082041,ZFMK‐DIP‐00082042,ZFMK‐DIP‐00082043,ZFMK‐DIP‐00082044,
ZFMK‐DIP‐00082045,ZFMK‐DIP‐00082046,ZFMK‐DIP‐00082047,ZFMK‐DIP‐00082048,
ZFMK‐DIP‐00082060,ZFMK‐DIP‐00082061,ZFMK‐DIP‐00082062,ZFMK‐DIP‐00082063,
ZFMK‐DIP‐00082064,ZFMK‐DIP‐00082065,ZFMK‐DIP‐00082066,ZFMK‐DIP‐00082068,
Diversity2022,14,53225of36
ZFMK‐DIP‐00082069,ZFMK‐DIP‐00082070,ZFMK‐DIP‐00082071,ZFMK‐DIP‐00082072,
ZFMK‐DIP‐00082073,ZFMK‐DIP‐00082074,ZFMK‐DIP‐00082076,ZFMK‐DIP‐00082077,
ZFMK‐DIP‐00082078,ZFMK‐DIP‐00082079,ZFMK‐DIP‐00082080,ZFMK‐DIP‐00082081
[ZFMK].
Distribution:Austria,Azerbaijan,Belgium,Bosnia–Herzegovina,Bulgaria,Croatia,
CzechRepublic,Denmark,Finland,France,Georgia,Germany,Hungary,Ireland,Neth‐
erlands,Norway,Poland,Portugal,Serbia,Slovakia,Slovenia,Spain,Sweden,Switzer‐
land,Turkey,UkraineandUnitedKingdom[43,55,64,65].
GenusPsychodaLatreille,1797
PsychodaLatreille,1796:152.Typespecies:TipulaphalaenoidesLinnaeus,1758by
subsequentdesignationofQuate[41](p.191).
Diagnosis.Specieswithshortvertex;labellumflattenedandcarryingdigitiform
setae;eyebridgewithoutinterocularsuture;antennawith12–14flagellomeres,those
beyond11thalwaysreducedinsizeandshowingdifferenttypesoffusion;flagellomeres
nodiformwithaneck(exceptapicalflagellomeres);ascoidsusuallywiththreebranches,
twoanteriorandoneposteriorbranch,Y‐shapedorwithfourbranches,shapedlikea
plussign(+);aedeagusoftenasymmetrical;hypopodswithasingleapicaltenaculum.
SpeciespresentinEuropeassociatedwithdendrotelmata:P.alternataSay,1824,P.
cinereaBanks,1894andP.minutaBanks,1894.[23,25](Table1).
PsychodaalternataSay,1824
PsychodaalternataSay,1824:358.Typelocality:USA,Pennsylvania,Philadelphia.
PsychodatripunctataMacquart,1838:85.Typelocality:notgiven,probablyFrance.
PsychodamarginepunctatavonRoser,1840:50.Typelocality:notgiven.
PsychodasexpunctataPhillipi,1865:631.Typelocality:notgiven.
PsychodaschizuraKincaid,1899:21.Typelocality:USA,Seattle,Washington.
PsychodanocturnalaHaseman,1907:319.Typelocality:USA,Missouri,Columbia.
PsychodafloridicaHaseman,1907:324.Typelocality:USA,Florida.
PsychodabengalensisBrunetti,1908:371.Typelocality:India,CalcuttaandSimla.
PsychodaalbimaculataWelch,1912:411.Typelocality:USA,Illinois.
PsychodadakotensisDyar,1926:109.Typelocality:USA,SouthDakota.
Psychodaalternatavar.marmosaAbreu,1930:123.Typelocality:notgiven.
Psychodaalternatavar.floridicaHaseman:Johannsen,1934:25.
Tineariaalternata(Say):[66](p.142);[63](p.114)[67](onlinecatalogue);[51](p.46);;
[68](p.96);[69](p.89);[70](p.107);
Psychoda(Tinearia)alternataSay:Bravoetal.[71](p5,11).
PsychodaalternataSay:[40](p.97).[41](p.218)[72](p.216);[73](p.195);[74](p.16);[75]
(p.12);[76](p238);[77](p.67);[78](p.21);[79](p.48).
Diagnosis.Eyesseparatedby1–3facetdiameters,withoutinterocularsuture;an‐
tennawith13flagellomeres,lastthreeflagellomeressmall,flagellomere11–12fused,
flagellomere13smallerandpartiallyfusedto12;labellumwithoneshortandfourlong
teethandfoursetae;aedeagusasymmetrical,withonparamerethickerthanaedeagus,
ejaculatoryapodemebroadposteriorly,longerthanaedeagalcomplex;hypopods2.3
timeslongerthangonostyli.[40].
Examinedmaterial.None.
Distribution.Cosmopolitan[40,55].InEurope,itispresentinAustria,Balearctic
Islands,Belgium,Bulgaria,Croatia,Cyprus,CzechRepublic,Dennmark,Finland,France,
Germany,Greece,Hungary,Ireland,Italy,Netherlands,Norway,Poland,Portugal,Ro‐
mania,Sardinia,Slovakia,Slovenia,Spain,Sweden,Switzerland,andUnitedKingom
[55].
PsychodacinereaBanks,1894
PsychodacinereaBanks,1894:331.Typelocality:USA,NewYork,SeaCliff,L.I.
Diversity2022,14,53226of36
ThreticuscomparEaton,1904:57.Typelocality:Algeria,England,Ireland,andMaderia.
PsychodaprudensCurran,1924:219.Typelocality:Canada,Alberta.
Psychodochacinerea(Banks):[66](p.135);[80](p.100).
Diagnosis.Eyesseparatedby1–2facetdiameters,withoutinterocularsuture;an‐
tennawith14flagellomeres,last3separatednotfused;labellumwithfourterminaldigi‐
tiformsetaeandupto5setiformsetae;medialforkwithbasalswelling;aedeagalcom‐
plexslightlyS‐shaped,asymmetrical,withunpairedbentparamere;aedeagussubtrian‐
gular,coveringasquarishstructure;ejaculatoryapodemetriangularatbaseindorsal
view;hypandriumtrapezoidal;hypopodsaslongasgonostyli,graduallynarrowingto‐
wardtheapex,butwithoutbasalswelling.
Examinedmaterial.Specimensnumber:No.0480‐4081[AM]Rearedfromden‐
drotelmataofahornbeam(CarpinusorOstryasp.).
Distribution.Cosmopolitan[71].InEurope,itispresentinAustria,Azoresarchi‐
pelago,Belgium,Bosnia–Herzegovina,Bulgaria,CanaryIslands,Corsica,CzechRepub‐
lic,Denmark,Finland,France,Germany,Greece,Hungary,Iceland,Ireland,Italy,Lux‐
emburg,MadeiraIslands,Netherlands,Norway,Poland,Romania,Sardina,Serbia,Slo‐
vakia,Slovenia,Spain,Sweden,Switzerland,andUnitedKingdom[43,49].
PsychodaminutaBanks,1894
PsychodaminutaBanks,1894:331.Typelocality:USA,NewYork,nearSeaCliff.
PsychodamarylandanaDelRosario,1936:111.
PsychodaspretaTonnoir,1940:57.
Psychodulaminuta(Banks):[52](p.56).
Diagnosis.Eyesseparatedby1facetdiameters,withoutinterocularsuture;antenna
with16flagellomeres,last4flagellomeresreducedinsizeandfused;labellumwithfour
terminaldigitiformsetaeandtwotrichiformsetae;aedeagusasymmetrical,flankedby
twolargetriangularplatesofmorphologicallyunknownorigin;gonostylinarrowingto
taperedpointinapicalfifth;hypandriumnarrow,nottrapezoidal;hypopodsmuch
longerthangonostyli,withbasalswelling.
Examinedmaterial.None.
Distribution.Holarctic.InEurope,itispresentinAustria,BalearicIslands,Belgium,
Bulgaria,Corsica,Cyprus,CzechRepublic,Denmark,Finland,France,Germany,Greece,
Hungary,Ireland,Italy,Lithuania,MadeiraIslands,Netherlands,Norway,Romania,
Sardinia,Slovakia,Slovenia,Spain,Sweden,Switzerland,andUnitedKingdom
[25,43,59,81].
GenusTelmatoscopusEaton,1904
TelmatoscopusEaton,1904:58.Typespecies:PericomaadvenaEaton,1893,bydesignation
ofQuate[82].
SciriaEnderlein,1935:247.Typespecies:PericomaadvenaEaton,1893,byoriginaldes‐
ignation(see[83]).
KrivosheinoscopusJežek,2001:57Typespecies:Telmatoscopusussuricus(Ježek,2001)syn.
nov.
Diagnosis(modifiedfromKvifte,[83]).Fronsandclypeusseparatedandnotpro‐
trudingovereyemargin;flagellomeresasymmetricallynodiform,withpaired
leaf‐shapedtodigitiformascoids;flagellomere14withelongatedapiculus;wingveins
R2+3notconnectedtoR4;apexofR5endingatwingapex;ejaculatoryapodemenarrow
indorsalviewanddistallyendingintwoshortbrancheswithmembranousconnection
toaedeagalcomplex;aedeagalcomplexsymmetrical;aedeagalcomplexencapsulatedin
aparameralsheath;hypopodswithindistinctlyfringedtenacula;epandriumwitha
singleforamen.
Remarks:Ježek[84]describedKrivosheinoscopusbasedonfivemalespecimensofthe
typespeciesK.ussuricusJežek,2001(nowTelmatoscopusussuricuscomb.nov.)fromthe
RussianFarEast.Later,hedescribedK.bartaiJežek,2004(nowTelmatoscopusbartaicomb.
Diversity2022,14,53227of36
nov.)fromtheCzechRepublic(Ježek2004).Ježek[84]providedadiagnostictableof
closelyrelatedtelmatoscopoidgenera,namely,Lepiseodina,Sciria(=Telmatoscopus),
Iranotelmatoscopus,Krivosheinoscopus,Telmatoscopusauctt.(=Seoda)[84](p.60).
Inthetable,sevenmorphologicalcharacterswereprovidedtodifferentiate
KrivosheinoscopusfromSciria(=Telmatoscopus),theyarenumberedanddiscussedbelow.
1. Thescape/pedicelproportionis2:1inKvirosheinoscopus(1:1inSciria);however,even
whiletheproportionofscape/pedicellengthproposedbyJežek[84]is2:1inT.us‐
suricus,thisisnotthecaseforT.bartaiwhereitis1:1.
2. Theascoidsareverylong,thinandcoiledinKrivosheinoscopus(large,flat,leafor
hood‐shapedinSciria).ThischaracteriscontradictedbyTelmatoscopuslaurencei[38],
whichhasdigitiformascoidsandcoiled,andthesecharacterstatesarepolymorphic
inVaillantodesWagner,2001andPanimerusEaton,1913.Intermediaryformsalso
occurinsomegeneraofTelmatoscopoids.Duetowidespreadpolymorphismofthis
characterwedonotconsiderthisareliablegenus‐levelcharacterunlesssupported
byother,independentlinesofevidence.
3. Thefirstpalpalsegmentveryshortandkeg‐shapedinKrivosheinoscopus(longand
cylindricalinSciria).ThischaracterisvariableinsidethegenusTelmatoscopus(e.g.,
longinTelmatosocpusadvenaandshorterinTelmatoscopusthuringicusBeran,Doczkal,
Pfister&Wagner,2010),thereforeitishereconsideredasintragenericvariability,
andthusnotadiagnosticcharacter.
4. Thepositionoftheradialandmedialforksofwingvenation,atthesamedistancein
Krivosheinoscopus(medialforkdistadtoradialforkinSciria).Thisisanothercharac‐
terpresentingasvariablewithinthegenusTelmatoscopus(e.g.,medialforkdistalin
T.advena,medialforkbasalinT.thuringicus,forksatthesamelevelinTelmatoscopus
bartai).
5. TwopairsofprotuberancesintheaedeagalcomplexinKrivosheinoscopus(onepairin
Sciria).Accordingtothehomologizationof[83],theabsenceofprotrudingpara‐
meresinTelmatoscopusarenotduetoabsenceofparameres,theyarepresentas
transversescleriteswithintheaedeagal–parameralcomplex.Thedifferencebetween
T.advena,T.bartaiandT.ussuricusintheparameresisaquestionofdegreeofde‐
velopmentratherthanaclear‐cutpresence/absencequestion,andintermediatecases
occurinNearcticspecies(e.g.,T.patibulusQuate).
6. Thehypoprocthasaterminalprojection,longandnarrowinKrivosheinoscopus
(withoutaterminalprojection,triangularinSciria).Thischaracteriscontradictedby
Telmatoscopusbartaicomb.nov.wherethehypoproctistriangular,andnotthinand
elongated.
WiththreesharedcharactersbetweenKrivosheinoscopusandTelmatoscopus(=Sciria
sensuJežek)includingthewingveinR5endingatwingapex,veinCuendingdistaltoM1+2
and,theejaculatoryapodemelaterallycompressed(=basalapodemeoftheaedeagal
complexinJežek)[84].
Furthermore,Ježeklistedseven(fivearecommentedhere)morphologicalcharacters
separatingKrivosheinoscopusfromSeodaEnderlein(consideredasTelmatoscopusauctt.In
Ježek[84]),including:
1.Thescape/pedicelproportion,being2:1inKrivosheinoscopus(3‐4:1inSeoda);this
remainsadiagnosticdifference.
2.Firstpalpalsegmentshortandkeg‐shapedinKrivosheinoscopus(longandcylin‐
dricalinSeoda).ThischaracterisvariableinsideTelmatoscopus,andisthusnotadiagnos‐
ticcharacter.
3.VeinCuendingdistaltoM1+2inKrivosheinoscopus(samellevelordistadtoM1+2in
Seoda)yetagain,avariablecharacter;
4.WingveinR5endingatwingapexinKrivosheinoscopus(endingbelowwingapexin
Seoda),thischaracterisdiagnosticbetweenSeodaandTelmatoscopusaspointedoutin[83].
Diversity2022,14,53228of36
5.TheejaculatoryapodemelaterallycompressedinKrivosheinoscopus
(dorso‐ventrallycompressedinSeoda)alsoadiagnosticcharacterpointedoutin[83].
DiagnosticmorphologicalcharacterstoseparateTelmatoscopusfromSeodapresented
in[83]includesinglepairofdigitiformtofiliformascoids(anadditionalringofsmallse‐
tiformascoids+themainpairofdigitiformascoidsinSeoda);fronsclearlyseparatedfrom
clypeusnotprotrudingoverthemesalmarginofeyesinTelmatoscopus(fronsfusedwith
clypeausandprotrudingoverthemesalmarginofeyesinSeoda);parameralscleritesnot
fusedinTelmatoscopus(parameralscleritesfusedinSeoda).
OurcurrentlyreformulatedconscriptionofTelmatoscopusincludes10speciesglob‐
ally,namelyT.advena(Eaton,1893),Palaearctic;T.bartai(Ježek,2004)comb.nov.,Eu‐
rope;T.dendrophilusVaillant,1983,USA;T.frondeusTokunaga&Etsuko,1955,Japan;T.
laurenceiFreeman,1953,Europe;T.pappi(Wagner,1979),Afghanistan;T.patibulusQuate,
1955,USA;T.ussuricus(Ježek,2001)comb.nov.,Russia;T.tanegashimensis(Ježek&Mogi,
1995),Japan;T.thuringicusBeran,Doczkal,Pfister&Wagner,2010,Europe.
Notes.Previously,Kvifte,[83]includedPanimeruswagneriSalamanna,1982asSeoda,
howevertheinclusionofT.wagneri(=T.advena)comb.nov.etsyn.nov.insideofTel‐
matoscopusisbasedonthemalegenitalmorphologypresentedintheoriginaldescription,
thereforethenewcombinationfromSeodatoTelmatoscopus.Additionally,thecharacters
separatingT.wagneri(=T.advena)syn.nov.and,T.advenaarequiteinconspicuous,
however,wedidnotexaminetheholotypeofT.wagneri(=T.advena)syn.nov.,nonethe‐
less,basedontheoriginaldescriptionandfigures,wedidnotfindanymorphological
charactersthatsupportthemasseparatesspecies,andwetreatT.wagneriasanewsyn‐
onymofT.advena.
AdditionalcharactersandcloserexaminationofthespeciesT.pappi,T.frondeus,and,
T.tanegashimensisisdesirableastheysharesomecharacterswiththegenusLepiseodina,
however,thesespeciesareoutsidethegeographicalscopeofthiswork,andthereforeare
includedasTelmatoscopusfollowing[83]untilabroaderrevisionofthegenusisavailable.
SpeciespresentinEuropeassociatedwithdendrotelmata:T.advena(Eaton,1904),T.
bartai(Ježek,2004)comb.nov.,T.laurenceiFreeman,1953,T.thuringicusBeran,Doczkal,
Pfister&Wagner,2010[5,13,25,38,39](Table1).
Telmatoscopusadvena(Eaton,1893)
Figures12–14
PericomaadvenaEaton,1893:127.Typelocality:GreatBritain.
Telmatoscopusadvena(Eaton):[81](pp.205–209)
Sciriaadvena(Eaton):[85](p.87).
Telmatoscopusadvenus(Eaton):[86](p.86).
Panimerusadvenus(Eaton):[50](p.80).
PanimerushavelkaiWagner,1975:1;(see[83]).
PanimeruswagneriSalamanna,1982syn.nov.
TelmatoscopusseguyiVaillant,1990:378;(see[83]).
Diagnosis.Male.Ascoidsbroad,leaf‐shapedandcoiled;phallomeresincurved,
symmetrical,lessthanhalfthelengthofejaculatoryapodeme;hypopodswithtenacula
restrictedtoapex.
Femaledescription(Figures15and16)basedonthedescriptionofthemalein[83]
thefemaleissimilartothemaleexcept:Headslightlylongerthanwide(0.5mmwide,0.6
mmlength).Eyebridgewithfourfacetrows,separatedby1.5facetdiameter;onlyfirst
palpalsegmentpresentinexaminedmaterial;apicalantennalflagellomeresabsentin
examinedmaterial,ascoidss‐shaped,coiled,thin(notbroadleaf‐shapedasinmale).
Winglength,wingwidth,lengthxtimesitswidth.
Diversity2022,14,53229of36
Figure15.Telmatoscopusadvena(Eaton,1983),female.(A)Head.(B)Genitalchamber.(C)Egg.(D)
Cerci.Scale(A–D)inmillimeters(mm).
Diversity2022,14,53230of36
Figure16.Telmatoscopusadvena(Eaton,1893),female.(A)Firstantennalsegments.(B)Sternite8and
genitalchamber.(C)Egg.(D)Cerci.Abbreviations:AP=anteriorpole,asc=ascoids,cerc=cerci,
flm=flagellomere,genchamb=genitalchamber,ped=pedicel,PP=posteriorpole,scp=scape,stn
=sternite.Scale(A–D)inmillimeters(mm).
Diversity2022,14,53231of36
Wing2.7timeslongerthanwide(1mmwide,2.7mmlong),Sclong,endingat
junctionofR4+R2+3.InfuscatedinareabetweenCandR1.
Sternite8(Subgenitalplate)aboutthesamelengthandwidth(0.25mmwide,0.24
mmlong),waistedinthemiddle;basalmarginstraight,apicalmargincoveredinsmall
setaeandstronglyconcaveinthemiddle.Cercilong(0.4mmlength),1.6timesthelength
ofthesternite8,with10setaeonthedorsalsurface,thesurfaceinthebaseofthecerci
presentsmultipleplate‐shapedprotuberances,resemblingthetextureofcrocodileskin.
Subgenitallobesrectangular.
Eggdescription.(Meanlength0.32mm,width0.1mm,n=15)Ovoid,longerthan
wide(0.1mmwide,0.32long).NoMicropylestructureisvisibleintheanteriorpole;
however,itdoesseemtobeacircularaperturewherethemicropylecouldbelocated.
Exochorionwithbothlongitudinalandtransversalsingle,flattenedridgesthatwhen
joined,formcells(mainlyrectangularorhexagonal)acrosstheexochorion.
Materialexamined.Holotypeexamined:no.235444[BMNH],ZFMK‐DIP‐00081296,
ZFMK‐DIP‐00081297,ZFMK‐DIP‐00081298,ZFMK‐DIP‐00081306,ZFMK‐DIP‐00081307,
ZFMK‐DIP‐00081308,ZFMK‐DIP‐00081309,ZFMK‐DIP‐00081310,ZFMK‐DIP00081313,
ZFMK‐DIP‐00081314,ZFMK‐DIP‐00081315,ZFMK‐DIP‐00081316,ZFMK‐DIP‐00081317,
ZFMK‐DIP‐00081318,ZFMK‐DIP‐00081319,ZFMK‐DIP‐00081325,ZFMK‐DIP‐00081320,
ZFMK‐DIP‐00081331,ZFMK‐DIP‐00081571,ZFMK‐DIP‐00081572,ZFMK‐DIP‐00081573,
ZFMK‐DIP‐00081581,ZFMK‐DIP‐00081582,ZFMK‐DIP‐00081620,ZFMK‐DIP‐00081621,
ZFMK‐DIP‐00081312,ZFMK‐DIP‐00081500,ZFMK‐DIP‐00081501,ZFMK‐DIP‐00081502,
ZFMK‐DIP‐00081509,ZFMK‐DIP‐00081511,ZFMK‐DIP‐00081517,ZFMK‐DIP‐00081520,
ZFMK‐DIP‐00081522,ZFMK‐DIP‐00081538,ZFMK‐DIP‐00081545,ZFMK‐DIP‐00081546,
ZFMK‐DIP‐00081547,ZFMK‐DIP‐00081549,ZFMK‐DIP‐00081550,ZFMK‐DIP‐00081553,
ZFMK‐DIP‐00082120,ZFMK‐DIP‐00082121,ZFMK‐DIP‐00082123,ZFMK‐DIP‐00082127,
ZFMK‐DIP‐00082129,ZFMK‐DIP‐00082131,ZFMK‐DIP‐00082132,ZFMK‐DIP‐00082134,
ZFMK‐DIP‐00082135,ZFMK‐DIP‐00081570,ZFMK‐DIP‐00081579,ZFMK‐DIP‐00081596
[ZFMK].Specimennumber:10047,13667,13602,13527,12687,18493,13193,18215,18237,
20784,20750,20823,0,11356,11264,11357/34235,21212,21315,22648,22649,22650,22651,
22652,22653[NMP].
Distribution.AeganIslands,Belgium,Finland,France,Germany,Ireland,Norway,
Slovakia,UnitedKingdom[13,43,87].
Telmatoscopusbartai(Ježek,2004)comb.nov.
Figure13
KrivosheinoscopusbartaiJežek,2004:117,Typelocality:CzechRepublic:Bohemiaor.,
ŽeleznéhoryMtsProtectedlandscapearea.
Diagnosis.Male.Ascoidsdigitiform,thin,long,andcoiled;phallomeressymmet‐
rical,outcurved,morethanhalfthelengthofejaculatoryapodeme;hypopodswithte‐
nacularestrictedtoapex.
Examinedmaterial.Holotypeexamined:Cat.No.34243[NMP].
ZFMK‐DIP‐00081598,ZFMK‐DIP‐00081597,ZFMK‐DIP‐00081599[ZFMK].
Distribution.CzechRepublic[88],Germany.Ourthreereportedspecimensarethe
firstrecordofthisspeciesinGermany.
TelmatoscopuslaurenceiFreeman,1953
TelmatoscopuslaurenceiFreeman,1953:71.Typelocality:GreatBritain.Herts,
Harpenden,RothamstedExperimentalStation.
Diagnosis.Male.Ascoidsdigitiform,thin,coiled;phallomerescoiled,lessthanhalf
thelengthofejaculatoryapodeme;hypopodswithtenacularestrictedtoapex.
Examinedmaterial.None.
Distribution.OnlyknownfromGreatBritain[13,39,43].
TelmatoscopusthuringicusBeran,Doczkal,Pfister&Wagner,2010
Diversity2022,14,53232of36
TelmatoscopusthuringicusBeran,Doczkal,Pfister&Wagner,2010:63.Typelocality:
Germany,Thuringia,NationalParkHainich,Weberstedt,Birkensee.
Diagnosis.Male.Ascoidsarebroad,leaf‐shaped,taperingtowardstheapexand
coiled;phallomeresoutcurved,lessthanhalfthelengthofejaculatoryapodeme;hy‐
popodswithaclusterofapproximately30tenaculaatapex,andmoretenaculascattered
alongtheentiresurfaceofhypopods.
Examinedmaterial.None.
Distribution.OnlyknownfromGermany[5].
KeytotheEuropeanadultmalesofTelmatoscopus
1. Morethan30tenacula;tenaculanotrestrictedtotheapicalportionofhypopods,
scatteredinallthehypopodslength(asin[5](Figure10))…T.thuringicus
‐ Lessthan30tenacula;tenacularestrictedtotheapicalportionofhypopodsandnot
scatteredinthehypopodslength…2
2. Ascoidsbroad,almostasbroadasthebasalnodeofflagellomerecarryingthem
(Figure1B);aedeaguswithparameresnotcoiledandstronglyincurved(Figure
12C)…T.advena
‐ Ascoidsnarrow,neverbroaderthan1/3ofthewidthofthebasalnodeofflagello‐
merecarryingthem;paramerescoiledoroutcurved(Figure13C)…3
3. Paramerescoiledwithstrongincurvationafterthecoilandapicaltipshook‐out
curved(asin[38](Figure2D));hypopodswithlessthan20tenacula…T.laurencei
‐ Parameresnotcoiledandoutcurved(Figure13C);hypopodswithmorethan20te‐
nacula…T.bartai
4.Discussion
Alloftheabove‐mentionedspecies(alsoinTable1)arerecordedtobeassociated
withdendrotelmatatoacertaindegree,whethertheyarespecializedtocompletetheir
lifecycleinthisecosystemortheyareonlyopportunisticisadifferentmatter.Asmen‐
tionedbyOboňaandJežek[25]theextremevariationofenvironmentalconditionssuch
aspH,temperature,frequentwaterlosswithrapidfloodingand,oxygendeficitcan
causethedeathofnon‐specializedspecies,especiallyinlarvalstagesthatareusingden‐
drotelmataasanirregularbreeding/developingsite,whilespecializedspeciescanendure
theseharsh822environmentalvariationsthrivingthroughalltheirlifecycle.
TheoccurrenceofClogmiaalbipunctata,C.xylophilus,Pneumiacanescens,P.trivialis,
Psychodaalternata,P.cinerea,andP.minutaindendrotelmataisratherincidentalorhighly
understudied[23,25].Inotherwords,thepresenceofthesespeciescouldbeanextension
oftheirregulardevelopmentsites(e.g.,smallwaterbodies,streams,etc.)andadults
happentofindwater‐filledtreeholesthatareapotentialdevelopmentsitefortheiroff‐
spring,thus,theycoulddevelopinDendrotelmataandsurvive,buttheirlong‐termus‐
ageofthissiteisnotwelldocumented.Furtherstudiesindifferenthabitatcouldprovide
keyinformationtobetterunderstandtheecologicalrelationshipbetweenwater‐filled
treeholesandthespeciesofPsychodinaethatdevelopinthem.Tothedate,onlya
handfulofstudiesspecificallytargetedmothfliesindendrotelmata.
ClogmiaalbipunctataisbroadlydistributedbothinEuropeandworldwideandisthe
mostsynanthropicspeciesinsidethePsychodidaefauna,itcanbecommonlyfoundin‐
sidebuildings,andsewagetreatmentplantsinalmosteverycity.Thehighsynanthropy
issharedwithsomespeciesofthegenusPsychoda,whicharealsooftenfoundincities,
includingP.alternataandP.cinerea,thisspeciescancertainlyadaptanddevelopinawide
rangeofhabitats,includingdendrotelmata,therefore,thesespeciescouldbeclassifiedas
generalistswhenitcomestositesforlarvaldevelopmentandnotboundtodevelopin
water‐filledtreeholes.SomefurtherPsychodaspeciesmaybefoundindendrotelmata,
butthedevelopmentofmostspeciesisstillunknown.
Diversity2022,14,53233of36
Lepiseodinatristis,L.rothschildi,TelmatoscopusadvenaandT.laurenceiare
well‐documentedspeciesthatdevelopinsidedendrotelmata(Table1)withmultiple
recordsofbothadultsandlarvae.Onthecontrary,TelmatoscopusthuringicusandT.bartai
comb.nov.,areonlyassumedtobedendrolimneticbasedonobservationsoftheclosely
relatedspeciesT.advena[5].InthecaseofthehereindescribedspeciesLepiseodina
latipennis—somespecimenscollectedinItalywererearedfromdecayingorganicmatter
collectedinadendrotelmatafromamapletree(Acersp.),furtherspecimenscollectedin
GermanycamefromaMalaiseTrapplacednexttoawater‐filledtreewholeinanOak
tree,thusprovingthatthisspeciesdevelopinsidedendrotelmataascongenericspecies.
5.Conclusions
InEurope,only13speciesofPsychodinaeareknowntodevelopinsidewater‐filled
treeholes,afterourextensiverecordsearchwereportthatthePsychodinaespeciesde‐
velopin13differenttreespecies,includingtwonewtreespeciesinwhichnoprevious
recordoflarvaldevelopmentwasreported.WeredescribedLepiseodinalatipennisthrough
holotypeandnewmaterialexamination,andwereportitforthefirsttimeinGermany.
WealsoreportTelmatoscopusbartaicomb.nov.forthefirsttimeinGermany,andwe
provideagenericdiscussionofTelmatoscopus.Water‐filledtreeholes(dendrotelmata)are
usuallyassociatedwitholdtreeswhicharecommonlyendangeredthroughtheforest
managementstrategiesappliedinEuropeancountries,andtheyarebecomingrareto
findinsideforests.Tosummarize,thereisagapintheknowledgeoftheecologicalin‐
teractionsinsidethePsychodinaeandtheirenvironment,furtherstudiescanpotentially
providenewinformation,newrecordsandnewinteractionsthatwouldbebeneficialto
betterunderstandtheecosystems.Furthermore,oldtreeindividualsremainakeycom‐
ponentintheforests,astheyharborhighbiodiversitythatisstillunderstudiedandthey
shouldremainuntoucheduntilthenaturaldecompositiontakesplace.
SupplementaryMaterials:Thefollowingsupportinginformationcanbedownloadedat:
https://www.mdpi.com/article/10.3390/d14070532/s1,TableS1:ExaminedmaterialcollectionData.
AuthorContributions:Conceptualization,S.J.‐S.,G.M.K.andX.M.;investigation,S.J.‐S.,A.M.,
G.M.K.andX.M.;Methodology,S.J.‐S.,A.M.,G.M.K.andX.M.;Resources,S.J.‐S.,A.M.,G.M.K.and
X.M.;Writing—originaldraft,S.J.‐S.;Writing—review&editing,A.M.,G.M.K.andX.M.Allau‐
thorshavereadandagreedtothepublishedversionofthemanuscript.
Funding:TheresearchwasfundedbytheBundesministeriumfürBildungundForschung,Berlin,
Germany,project“GermanBarcodeofLifeIII:DarkTaxa”(FKZ16LI1901B).
InstitutionalReviewBoardStatement:Notapplicable.
InformedConsentStatement:Notapplicable.
DataAvailabilityStatement:Notapplicable.
Acknowledgments:TheauthorsthankBjörnMüllerforhisincrediblehelpduringtheDNAex‐
tractionprocessofspecimens.WeextendourgratitudetoBjörnRulikforcollectingthesomeofthe
specimensexaminedforthiswork.WearethankfultoMichalTkočforallowingthefirstauthorto
visitthePsychodidaecollectioninPrague.WearethankfultoalltheGBOLIII:DarkTaxateamfor
theirconstantsupport.LastbutnotleastwearethankfultoGregCurlerandWeiaReinboudfor
valuablediscussionsontaxonomyofdendrolimneticPsychodidae.
ConflictsofInterest:Theauthorsdeclarenoconflictofinterest.
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