Table Of ContentElyt,・a,「otkyo,26(2):223-248,Novemberl5,l998
ReorganizationoftheOreocarabusComplex(Coleoptera,Carabidae)
BasedonEndopha11icMorphologyandMolecularPhylogeny
Yuki IMURA
DepartmentofGynecology,TokyuGeneralHospital,Kita-senzoku,
1-45-6,0ta-ku,Tokyo,145-0062Japan,
Zhi-HuiSU,Cheong-GenKIMandSyozoOsAwA
JTBiohistoryResearchHall,1-1Murasaki-Cho,
Takatsuki,Osaka,569-1125Japan
Abstract EighteentaxaofthegenusCa1abus(s.lat)belongingto thesubgenus
0,-eoca1・abus(sensuIMuRA&MlzUsAwA,1996)havebeenexaminedforendophallicmor-
phologyandmitochondrial ND5genesequencestoanalysetheirphylogeneticrelation-
ships.Thisgroup,whichhasbeenintegratedtoasinglesubgenus0,-eocarabus,ispoly-
phyleticinaND5genealogical tree,andshouldbeseparatedintoat least8distinct lin-
eages. 0neof themincludesCa,・abus(“0t・coca,abus”)fat・lothatsharescommonancestry
withthreespeciesbelongingtothesubgenusR/7igoca1・abtls.Theother7lineages, 0,-tno-
, ca1-a加s,Cyr1/oca,-a加s, llpo1,oca,,a加s, Pill・1coca1,a加s,Ca,pat ﾍop/1izfls 「 αff nocaMMs
andCa、,azzut1ocatabusarewellisolatedonthetree.Astheirbranchingpointsareancient
ontheND5genealogicaltree,itwouldbeappropriatetoregardeachofthemasaninde-
pendentsubgenus.
Introduction
01-eocarabusGEHIN,1876(sensuIMURA&MlzUsAwA,1996)isafairly largesub-
genus in thegenusCarabus (s lat),containingover 20speciesdistributed in the
EurasianContinent. It is characterizedmainly byheptap1oidelytra1sculpture,and is
otherwise rather poor in its own morphological peculiarity. All the Oreocarabus
speciesbearsuchcharactersasbisetosepenultimatesegmentofthelabialpalpus,hairy
submentum, thepresenceofhairlessventraldepressions(=thiridium)onthemalean-
tenna,well-developedhindanglesofthepronotumandratherprominentelytra1shoul-
ders,etc.,butnoneofthemcanberegardedasanautapomorphyfor thesubgenus.0n
theotherhand,endopha11icstructureofthemalegenitalorganrevealsaconsiderable
diversification,suggestingthat Oreocat・abus isnotnecessarily uniform,andmaybe
classifiedintoatleastseveralspecies-groups(IMURA&MIzUsAwA,1996).
Inthispaper,arevisionalstudyhasbeenmadeusingtotally180reoca1-abustaxa
fromEurope,Turkey,CaucasusandChina,byexaminingdetailedendopha11icmor-
224 YILlkiIMUR^etal
pho1ogyofthemalegenitalorganandbyconstructingagenealogicaltreebasedonthe
mitochondrialND5genesequences.
Materialsand Methods
The followingspecimensbelongingtothesubgenusOreocalabus(sensuIMURA
&M1zusAwA,1996) fromEurope,Turkey,CaucasusandChinawereused.Localities
andDDBJ/EMBL/GenBankaccessionnumbersoftheND5genesequencesaregiven
inparentheses.
1. Cal・abusp11tzeysiamlsputzeyslanusGEHIN(NW.Italy,AB017451)
2. C.p.laynaudiamlsDEUvEetSIMARD(NW.Italy,AB017448)
3. C.p.pedemontanusGANGLBAUER(NWItaly,AB017452)
4. C.p germanaeCAsALEetCAvAzzUTI(NWItaly,AB017453)
5. C.p.omensisBORN(NW Italy,AB017454&ABC17456)
6. C.fairmalrelpelvicusCAvAzzuTI(NWItaly,AB0l7449&AB0l7450)
7. Cheteromorp17usDANIEL(NW.Italy,AB017455&AB017457)
8. Cconco1ol DANIEL(C.Switzerlan AB017458)
9. C.sylvestrisPANZER(E.Austria,AB017459)
10. C.latrei11eamlsCsIKl(NW.Italy,AB017478)
11. C.linneiPANZER(NE.Slovakia,AB017472;N.Slovakia,AB0l7473)
12. C hortensisLINNE(W. Austria,AB017463;W.Slovakia,AB017464;NE.
Slovakia,AB017465)
13. Ccribr・,atusQUENsEL(NE.Turkey,AB017460)
14. C.oribi・atusQUENsEL(C.Caucasus,AB017461)
15. CporrectangulusGEHIN(NE.Turkey,AB017462)
16. C glabratusPAYKuLL(WAustria, AB017466;N.&E.France,D86207;
NE.Slovakia,AB017467)
17. CgtabratusPAYKuLL(Mt.Caste,AB017468;Guardabosone,AB017469;
Andrate,AB017470;Cane11i,AB017471;all fromNW.Italy)
18. C.lat1-o1mnshanenslsDEuvE(Sichuan,China,AB017476)
19. C titanusBREUNING(Sichuan,China,AB017474)
20. C.su1IMURAetZHoU(Beijing,China,AB017475)
Inaddition to theabove, theND5gene fromonemaleexampleofC. (
1go-
calabus) pusio SEMENov from thePassGongga Ling of northern Sichuan, China
(AB017477),wassequenced forcomparison.
ForeachtaxonexceptC.cribratus(C.Caucasus),endophallusofthemalegeni-
taliawas fullyeverted,air-driedandfigured.For theanalyticalmethodsoftheND5
genesequencesandconstructionofagenealogical treebytheneighbor-joining(NJ)
method(SAIToU&NEI,1987),seeSUetal. (1996a,b, '98),IMURAeta1. (1997)and
IMURA,KIMeta1. (1998).ToknowthephylogeneticpositionofOreoca1-abusspecies,
theND5genesequencesofsomeothercarabinespecieswereused forconstructingthe
tree.
Reorganizationofthe01・coca,abusComplex 225
ResultsandDiscussion
OreocarabuswasfirstproposedbyGEHINin1876(p. 12).Heestablished this
“vagabond” higher taxon for the following 20 species: guadarr,amus, heyden1
(=steuartii),erransGORY,1839[necFISCHER,1823](=amplipennisLApoUGE,1924),
deyro11e1,ghi11ianli(=ghaiam),hortensis,linnei,carinthiacus,latrei11eiDEJEAN,1826
[necFISCHER,1822](=1atrei11eanusCsIKI,1927),alpinus(=conco1or),putzeyslanus,
) fochm owf (=fosc ﾝnf ｽovf , syfvesfrz oppef (=afpesfrfs oppz1), fafrmazrez, fzneofa_
? ttes (=massagettls), oysz,mane前,sfo'zowf (=fosc(cid:0) (cid:18553)ovz' ) a fruncafzco fs. In
viewofthepresentcriteria, theyarenothingbutanassemblageofvarioustaxaby
meresuperficialresemblance,andsomeofthemshouldbetransferredtoseveralother
subgenera.GEHINdidnotdesignatethetypespeciesatthattime.
In1896,REITTERplacedOreocarabusasoneofthe ft subgeneraoftheCarabi
multistriati,andrestrictedit toseveralIberianspeciessuchasghilian1,errans(=am_
plipennls),guadarramusandsteuarti.
LAPOuGE(1931)regardedOreocarabusasadistinctgenusbelongingtothedivi-
sionMultistries,andrecognizedinit fourcomponentsubgenera,namely, 0,eocarabus
(s.str), 0rinocar,abus,CaucasocarabusandNelocarabus. 0,-eoca,abus(s.str) inhis
sensecomprisesonlyasinglespecies,crt・ans(=amplipennis).
Inhismonograph,BREUNING(1932,'33) treatedOreocarabusasoneofthesub-
generainthedivisionMultistriati,anddivideditintothe following6sections:Oreo-
ca'abus(s.str),Titanocarabus, 0rinocarabus,Pac/nvcal,abus,U1ocarabusandMega-
nebrius.Thesection01-eocarabusinhissensecomprisesthefollowing10species:am-
fpemfs, seudostetfarfl, guadarl'amtls, gﾝlfzam orfensis, wfadsfmfrs f (=vfadf-
mi''s '!'),orle''afMs, geme afus, faeda加sandgfab,,a加s. At thesame time,hedesig-
nated Ca''a t'sampflpe,mzsasthe pcspeciesof0,-eoca,,abMs.
AccordingtoDEuvE(1994), 0reocarabusisoneofthe65subgeneraoftheLobi-
fora,thelargestdivisionofthegenusCa・labus(s lat).Hesynonymizedthefollowing
8highertaxawiththesubgenusOreoca1-abus:Carpathophilus,Cyti1ocarabus,Euporo-
carabus,Hypsoca1-abus,0rinocarabus,Phricocarabus,PiocarabusandTitanocarabus.
Recently, anewsystemwasproposedby IMURA(1996), andwas followedby
IMuRAandMIzusAwA(1996).Accordingtohisclassification, 0reocarabusisoneof
the17subgenerainthesubdivisionLatitarsiofthedivisionMultistriati.Itiscomposed
ofover20speciesdistributedintheEurasianContinentandmaybeclassifiedinto5
species-groups.
Thus, thereisnoconsensus ofpreviousauthorsastothetaxonomy of Oreo-
ca1-abus(s lat).Suchaconfusionseemstohavebeencausedmainlybydifficulty in
determiningautapomorphy for thespeciesbelonging to thishigher taxon,exceptthat
theyfundamentallyhaveheptap1oidelytra1sculpture.
Figure1showsagenealogicaltreeconstructedfromthemitochondrialND5gene
sequencesincludingall thetaxaanalysedin thisstudy togetherwithsomeother cara-
binespecies.Eightwell-definedlineagesarerecognizedincluding“01-eocarabus”taxa,
226 YukilMURAeta1
mostofwhichemergedca.30MYA(fordating,seeKIMetal.,1998),andwereinde-
pendent fromallothercarabinetaxa,exceptthatCala'bus(“0reocarabus”) latroWas
clusteredwith theRhigocalabusspeciessupportedbyahighbootstrap value. This
wouldmeanthattheemergenceoftheselineagesmaybetracedbacktoaboutthesame
timeofthecarabineradiation(SU,0KADAeta1.,1996;Su,KIMeta1., l998).
Thefirstcluster iscomposedof5species,putzeysianus, fairmaire1,heteromor-
phus,conco1orandsylvestris,andmaybedividedintotwosubclusters.Theseparation
ofthetwosubclusters israther oldandmaybeestimatedtohavetakenplace t2-16
MYA.Evolutionarydistancesbetweenthetaxa(speciesorsubspecies) ineachsub-
clusteraresmall,especiallyinthefirstsubcluster.Thefirstsubclustercontains4of5
subspeciesofC. putzey.slanus, i.e.,nominotypica1putzeyslanus,pedemontanus,go'-
manaeand1・aynaudianus,allhavingthe putzeysianus”一typemalegenitalia, together
withC fafrmairez efvfcus,themalegenitaliaofwhichareofthe“fazmaz''of一' 一
ThesecondoneiscomposedofC.putzeysianusomensls十C heteromorphusandC.
conco1or十C.sylvestns. 0menslsismorphologicallyregardedbeyonddoubtasasub-
speciesofC.putzeyslanus,asseenbythe putzeyslanus”一typegenitalicstructure(see
Fig.15).FromtheND5genesequences,however, it ismostcloselyrelatedtoC he-
teromorphushaving the'f(alrmalret”一typegenitalia.Sincewehavebeenunable to
analysetheotherracesdistributedintheinterveningareabetweenthoseofp.omensls
and eferomorp us,i.e.,anothersubspeciesoffazmazrezorcemszus(seeFig 46),itis
tooprematuretodiscusstherelationbetweenphylogenyandmorphologicaldifferenti-
ationamongthesecomplicatedalpinecarabids.
Astothesubgenericname for thisgroup,weadoptOrlnocalabusKRAATz(type
species:Csylvestris).Presumablythefollowingspeciesareincludedinthissubgenus:
adame icofa, afpestris, ertofzm,car加fhiacus, casfanopterusand cemszus.Judging
fromtheendopha11icmorphology, 0reocarabus(s.str)(typespecies:C amplipennis
fromtheIberianPeninsula,seeFigs 40& 45)couldbeused,but thedistributional
areasofthesetwogroupsarecompletelyseparated fromeachother(Fig 47). Inour
view,useofOreocarabus(s.str)shouldberestrictedtoseveralspeciesdistributedin
theIberianPeninsula(e.g.,ghilian1,guadarramusandpseudosteuarti,etc.).
Thesecond cluster is formedby C. oribi,atusandC. porrectangulus. Carabus
cribratusfromCaucasusandC.porrectangulusfromnorthernTurkeyareclusteredto-
gether,andtheevolutionarydistancebetweenthemisveryclose. Ca,abuscribratus
fromnortheasternTurkeyistheoutgroupofthem.Inotherwords,thetwospeciesare
notdividedfromeachotheronthephylogenetictree.Ascribr,atusandporrectangulus
areclearlydistinguishablefromoneanothermainlybytheshapeofaedeaga1apex(see
Fig. 1. Genealogical treeofthemitochondrialND5geneoftheOreoca,・abuscomplexwithsomeother
carabinespecies. ANJ-treewasconstructedbySINCAsequenceanalysispackageusing KIMURA's
two-parameterevolutionarydistance.Thebootstrapconfidencelevel(%)(basedon500resamplings)
isshownateachbranchingpoint.TheND5genesequencesoftwoCych・tusspecieswereusedasan
outgroup.Thesubgenericnamesproposedinthispaperforthe0,,cecal・abuscomplexareindicatedon
therightcolumn.
Reorganizationofthe0,・eoca1abtlsComplex 227
0.08 0.06 0.04 0.02
Distance
100 Cychrusthibetanus (Sichuan.China) Subgenericname
Cymoraw1lzi(Japan) adoptedinthispaper
Caabus(0reocarabus)putzeys1a'nusra)/naudlanus
(LaghidiCollalunga.ltaly)
C(0)fat,ma1re1pe/vlcus (ColtediBellino.Italy)
C(0)fpe/vlcus (ColtediBeliino,Italy)
50 C.(0)p.puteJ/slanus (ColtedeltaLombarda,Italy)
100 4972 CC.((00.))ppgpeemdea/nr1aoneta(nMuts.S(Cacoclalaredleiol.TIetarmly)ini.Italy) Orlnocarabus
C(0)pomensls(ColdelMule,Italy)
C. (0)heteromophus (Champorcher, Italy)
6 C.(0)pomens1s(ColdelMule,Italy)
100 C(0.)1/eteromolphus(Champorcher,Italy)
C.(0)conco/or(Furka-Pass,Swiss)
C.10)sy/vestris (Baumgartner,Austria)
C.(Mesocarabu)prob/ematcus (France)
Pentacarabus)hafnanc/l (Japan)
C (0reocarabus)crlbratus (Artvin,Turkey)]
- C(0.)crlbratus(Mt.Khumaratkhok.Russia) Cyt7iocarabus
100
9 C(0.)porrectangu/us(Gijmushane.Turkey)
C. (0.)hortens1s(Innsbruck Austria) ]
100 C(0)hortens1s(Trencln.Slovakia) Euporocarabus
71 C. (0)hortensls(HighTatras,Slovakia)
96 C.(0)g/abatus(Innsbruck,Austria) l
3 10
C(0)g/abratus(Bourgogne,France)
C(0)g/abratus(HighTatras,Slovakia)
98 C.(0)g.1ator(Mt.Caste,Italy) Phnco 0us
C(0)g./ator (Guardabosone.Italy)
100
C(0)g./ator (Andrate,Italy)
50
81 c.(0.)g./ator (cane1ll.、taly)
1 C(0.)/lnne1(HighTatras,Slovakia) ]
Ca,pathophiius
C (0)/1nne1(Zilina,Slovakia)
100 l- c (0.)manus(Jiuzhaigou.Sichuan.China) ] r,'tanoca 0us
l_ C(0)sui(Beijing.China)
32 c (Leptocarabus)proceu/us (Japan)
C.(Au/onocarabus)kurilensls(Japan)
ll
c(Rhigocarabus)cateniger (Qinghai,China)
(cid:0)(cid:0)(cid:0) 100 ー」 (0reocaaCbu(Fsl)hjglaotceara(bJしiuIsz)hpauigsolou,(SSicichhuuaann..CChhininaa)) コ RhlgOCafabuS
2 _ C (R)confucius(Sichuan.China)
carabus)decolor(Russia)
(T)firatercu/us(China)
(Asttlenocarabus)epact」1us (Japan)
30
39 、.(Scambocarabus)kruberi (China)
)carabus)/atrel//eanus (Champorcher,Italy) コCavazzutiocarabus
7
(Lirnnocarabus)maackaquati1/s(Japan)
100 1Eu/eotocarabus)porectlco//Is (Japan)
C. (Hemicarabus)tubeof」/osus (Japan)
C(Homoeocarabus)maeander (Japan)
23 100 C.(01lomopterus)1nsu/;cola(Japan)
C. (0)de1laanii (Japan)
4 C(S.St「.) vanνo/xeml(Japan)
c(s.slr)arvens1sconc/ator (Japan)
,。。
, c(s.str)aarvens1s(France)
C.(Archicarabus)nemoralsi (France)
Chrysocarabus)auronltens (France)
C.(Copto/abus)firuhstorelri (Japan)
100
c(Damasetl)blaptolclesrugipennls (Japan)
100 (D)bb/aptoides(Japan)
70 P,ocrustes)coraceus (France)
81 C.(Megodontus)ko/bel (Japan)
99 C(M)v1olaceus(France)
228 Yuki IMURAelal
Figs.29-31),morphological transformationshouldhavetakenplaceduringevolution
withinrelativelyshort time.Sincethey formadiscreteclusteron theND5tree, the
subgenericnameCyt1iocarabus(REITTER,1896)maybeadoptedforacceptanceofthis
group.A unique Iranianspecies,geme11atusMENETRIEs, belongspresumablytothe
samesubgenus,thoughitsrangeisomittedfromthedistributionmapofCyt1iocarabus
showninFig 47.
Calabus hortensis hasbeen regardedas atypical member of Oreocalabusby
mostauthors.However,itbelongstotheunique(3rd)clusterintheND5tree,andwe
adoptEupo1,ocarabus(REITTER,1896) for lt. Thiscarabidseemstohaveanaffinity
withthenext(4th)clustercomposedsolelyofC glabratus,thoughthebranchingpoint
ofthesetwoclustersisconsiderablydeep.Thesetwogroupsarealsosimilarwitheach
otherinmorphologyoftheendopha11us,especiallyofpeculiarlyshapedparapraeputia1
lobes(seeFigs 32-34).
The4thclustercontainsC glabratus.Noteworthyistherelationbetweenthepop-
ulationofnorthwesternItalyandthoseoftheotherregions(France,AustriaandSlova-
kia).Itiswell-knownthatthegeographicalvariationofC glabratusisnotremarkable
despiteitswidedistributionalrange. IndeecしtheItalianraceishardlydistinguishable
fromthoseofFrance,AustriaandSlovakiabothintheexternalandgenitalicstruc-
tures,except thatthe formerhasalittlewiderpronotum(seeFigs 24-25,33-34).In
theND5tree,however,theyareclearlydividedintotwoclusterswithadeepbranching
point,indicatingthatthemorphologicaldifferentiationbetweenthetworaceshasbeen
veryslow incontrasttoalonghistoryaftertheirseparation.Thesimilarsituationsmay
berecognizedincertaingroupsoftheChineseApotomopterus(Suetal.,1998b)orof
theJapaneseDamaste, (s.str)carabids(SU,ToMINAGAeta1., l998),whichalsoshow
very littlemorphologicaldifferenceslongafter theirdiversification.Crossingexperi-
mentsremaintobedonetoknowwhether thesetwoglab1-atusracesareregardedas
siblingspeciesorsimplyassubspeciesretainingmatingabilitybetweenthem.Weten-
tatively recognizethenorthwesternItalypopulationasagoodsubspeciesunder the
namelatior,whichwasdescribed fromPiemontebyBORN(1895) asavarietyofC
glab1-atus.ThetwosubspeciesareapparentlyborderedbytheEuropeanAlps(Fig48).
Carabus11nneihasbeenplacedinOrinocarabusbymostauthors,but itbelongs
to a discrete lineage far remote from that of Orinoca1-abus, and we adopt Car-
pathop11ilus(REITTER,1896) forthisspecies.Thistreatmentissupportedbycharacter-
isticallyshapedendopha11usasshowninFig35.
CarabustitanusandC.su1, thelatterofwhichwasdescribedveryrecently from
Beijing(IMURA&ZHoU,1998),belongtoalineageindependent fromother “Oreo-
cal'abus”species.Thetreesuggeststhatthetwospecieswereseparatedratherrecently.
TheapplicablesubgenericnametothisclusterwouldbeTitanoca1-abus,whichwases-
tablishedbyBREUNING(1932) forC titanus.
Carabus lat1'o is endemic to northern Sichuan and has been one of the least
knownspeciesoftheChineseCarabinauntil recentyears. Itssystematicpositionhas
beenvariableaccordingtoauthors.AlthoughSEMENov(1898)establishedamonotypi-
Reorganizationofthe0,・coca,・abusComplex 229
cal higher taxon, Hypsocal・abus, for thisspecies, it was synonymizedeither with
Orinoca1・abus(BREUNING, 1932, '33)orwithOreoca'abus(DEUVE,1994; IMURA&
M1zUsAwA,1996).FromtheND5genealogical tree,however,C. fatl'oobviouslybe-
longstotheclustercontainingsuchRhigocalabusspeciesascatenlge1,confuciusand
Inviewoftheexternalmorphology,C. latroalmostagreesinthe fundamental
pus1o.
structurewithmostspeciesofRhigoca,・abus.Alsofromtheendophallicmorphology,it
seemstobeconsistentwiththecriteriaestablished foralltheothermembersofRhigo-
ca1-abus, thoughthemembraneousprojectionsshowconsiderablevariationamongthe
Rhigocarabusspecies.InFigs41-42areshownthepeculiarlyshapedendopha11usof
C. latro,thedetailsofwhichin fulleversionareillustrated forthefirsttime.Thus,C.
fatl,oshouldberegardedasamemberofthesubgenusRhigocalabus(REITTER,1896).
Thisspeciescomprisesthreesubspecies(withtheexceptionofqinlingensisIMuRAof
theQinlingMountains,whichshouldberegardedasanindependentspecies).Allot
these are distributed in theeasternmost periphery of the rangeof thesubgenus as
showninFig 50.
Cal・abuslatrei11eanusisasmall-sizedspecieswithanarrowdistributionalrange,
whichislimitedtothePennineAlpsandtheGranParadiseMassifinthewesternpart
oftheAlps(Fig47).Almostallthepreviousauthorshaveregardeditasnothingbuta
regularmemberofOrinoca1-abusfromitsexternalmorphology.However,theND5tree
showsthatthisspeciesbelongstoanisolatedlineagewhichiscompletelyindependent
not only from that of Orinoccu,abus but also from the other 71inea9es mentioned
above.Theendopha11icstructureofCfatle,i11eamlsisalsoveryuniquelyspecializedin
shape(seeFigs 43&44).Onthebasisofthesefindings,IMuRA(1998)proposeda
newsubgenusunderthenameCavazzutiocalabus forthisuniquealpinecarabid.AC-
cordjngtopersonalcommunicationfromProf.P.CAvAzzuTI,whokindlycollectedfo「
usaserjesofalcohol-treatedcarabidsamplesfromnorthwesternItaly,C. fatl'e' i11eanuS
jscharacterizedalso fromthephenologicalviewpoint. Inthespring,itstartsintoaC-
tjonmuchearlierthantheothersympatrica11yoccurringOrinoca1a'busspeciesSuchas
chote・lo1norphusandC.conco1o1.Astothehabitat,itprefersenvironmentsimilarto
thatofPlatyca1-abusratherthantotheaveragebiotopeofOrinoca1'abuS.
ReorganizationoftheTaxaTreatedinthisStudy
1. SubgenusOrinocalabusKRAATz,1878
(Typespecies:C.sylvestrisPANZER)
1. C.(0rinocarabus)putzeysianusputzeysianusGEHIN,1876
2. C.(0.)p raynaudianusDEUvEetSIMARD,1977
3. C.(0.)p.pedemontanusGANGLBAUER,1892
4. C.(0.)p germanaeCAsALEetCA?AZZUTI,1977
5. C.(0.、)p.omenslsBORN, l901
6. C.(0.)fcairma1i-eipelvlcusCAVAZZUTI,1973
230 Yuki IMURAeta1
7 C.(0.)heteromorphusDANIEL,1896
8 C.(0.)conco1orDANIEL,1896
9 C.(0.)sylvestrisPANZER,1793
11. SubgenusCyt11ocarabusREITTER,1896
(Typespecies: C.cribratusQUENsEL)
1. C.(〔:yt11ocarabus)cribratusQUENsEL,1806(NE.Turkey)
2. C.(Cy)cribratusQUENsEL,1806(C.Caucasus)
3. C.(Cy)porrectangulusGEHIN,1885
111. SubgenusEuporocarabu‘sREITTER, l896
(Typespecies: C hortenslsLINNE)
C.(Euporocarabus)hortenszsLINNE,1758
IV SubgenusPhricocarabusREITTER,1896
(Typespecies:CglabratusPAYKULL)
l. C.(Phricocarabus)glabratusglabratusPAYKULL,1790
2. C.(P)g.latterBORN,1895,bonasubspecies
SubgenusCarpathophilusREITTER,1896
(Typespecies:C.linneiPANZER)
C.(Ca atopazfus)f加 ez'PANZER,1812
VI. SubgenusTitanocarabusBREUNING,1932
(TypeSpecies:CtitanusBREUNING)
1. C.(Titanocarabus)tltanusBREUNNIG,1932
2. C.(T)sui lMURAetZHoU,1998
VIi. SubgenusRhigocarabusREITTER,1896
(TypeSpecies:CmorawttzianusSEMENov)
C.(Rhzgocarabus)latrominshanenslsDEUvE,1987
VIii. SubgenusCavazzut1ocarabusIMURA,1998
(Typespecies:C.latre111eanusCslKl)
C.(Cavazzutiocarabus)latre111eanusCslKl,1927
Reorganizationo「the0,・coca,ab1lsComplex 231
Appendix
Morphologyof fullyevertedendopha11usofeachtaxonusedinthisstudyisgiven
below.TheirdistributionalareasareshowninFigs 46-50.
1) Carabus(0rinocarabus)putzeysianusputzeyst'anusGEHIN,1876
(Figs 2, ll)
Malegenitalia. Aedeagus peculiarlyshaped, very longandslender, with the
apicallobeshortanddolabriforminlateralview;ostiumlobemoderatelysized,with
thetipslightlybilobed;neitherbasallateral lobesnorparaligulaisvisible,medianlobe
alsonotclearlyrecognizedthoughthemembraneouswall for itspositionslightlyin-
flated;prepraeputia11obesrecognizedasasmallhairyareawith faintinflation;para-
praeputial lobesrathersmallandsymmetrical;praeputia1padmoderatelydeveloped
anddome-likeinshape;apicalportionofendopha11usrathershortandsimple,apical
lobessmallandpodian1obesvestigial.
Specimensexamined. 1 (for drawingofendophalIus), Coltedelta Lombarda,
Vinadio,Cuneo,Piemonte,NW Italy,7-VII-1968,P.CAvAzzuTl leg;1 (for DNA
analysis),samelocality(2,300m),15-VI-1997,P.CAvAzzUTlleg.
2) Carabus(0rmocarabus)putzeysianusraynaudianusDEUvEetSIMARD,1977
(Figs3,12)
Malegenitalia. Aedeagusasinthenominotypicalsubspecies,butthedorsaltip
ofapical lobealittlelessstronglyprotrudent;position formedianlobealittlemore
stronglyinflatedthaninthenominotypicalsubspecies;praeputia1padlargerandmore
stronglyprojecteddorsad.
Specimensex;amlned. 1 (for drawing of endopha11us), Laghi di Co11alunga,
2,600m,Vinadio,Cuneo,Piemonte,NW Italy,11-VII-i993,P.CAvAzzUT11eg;1(S
(forDNAanalysis),samelocality(2,700m),20-VII-l997,PCAvAzzuTIleg.
3) Carabus(0rmocarabus)putzeysianuspedemontanusGANGLBAUER,1892
(Figs 4,13)
Malegenitalia. Aedeaguswiththebasal portionnarrower andslenderer,and
moreacutelybentventradatbasalthirdthaninthenominotypical form;apicallobeof
aedeagusmoregentlyroundedasawhole;positionsforbasal laterallobesandmedian
lobealittlemorestronglyinflatedthaninthenominotypica1form;podian1obesobvi-
ouslyprotrudentventra andapicallobesalittlelarger.
Specimens examined. 1 (for drawing of endophallus), Colladel Termini,
2,000m, Mt. Cinaiera, 、f. Corsaglia, Cuneo, Piemonte, NW Italy, 23-VI-1997,P.
CAvAzzUTl leg; 1 (forDNAanalysis),ValCasotto,2,000m, inthesamemountain,
232 YukiIMURAeta/
15-VI-1997,P.CAVAZZUTIleg
4) Carabus(0rmocarabus)putzeysianusgerlnanaeCAsALEetCA?AZZUTI,1977
(Figs5,14)
Malegenitalia. Aedeagussimilartothatofsubsp.pedemontanus,but thebasal
portionnotsostronglynarrowedandalittlelessacutelybentventrad,withtheapical
marginof apical lobe less roundlyshaped; ostium lobe the largest and the most
strongly projected dorsa though itsapexisnotremarkablybilobed; prepraeputia1
lobes also the most remarkably recognized in all the subspecies of putzeyslanus,
thoughmuchsmallerascomparedwiththeotherspeciesofthesamegroup;praeputia1
padalittlelargerthanthatof1,aynaudianus;podian1obealittlesmallerthaninpede-
montanus.
Specimense)camined. 13(fordrawingofendopha1lus),Mt.Saccare1lo,2,200m,
Viozene,、JTanaro,Cuneo,Piemonte,NW. Italy,24-VI-1994, L.&P.CAvAZZUTI leg;
1(S(forDNAanalysis),samelocality,2-VI-1997,P.CA?AzzUTIleg.
5) Carabus(0rmocarabus)putzeysianusomensisBORN,1901
(Figs6,15)
Malegenitalia. Aedeagusaml ostthesameasinthenominotypica1subspecies
ofputzeysianus,butmorestronglybentventradatapicalthird;apicallobeofaedeagus
lesswidelydilatedinlateralview,withtheapicalventralcornerhardlyprotrude api-
caldorsalonemoresharplypointed;basalapicalportionofendopha11usmoderately
inflatedandmedianlobeunclear;prepraeputia11obesvestigialbutpraeputia1padlarge
andstronglyprojecteddorsadasingermanae;apical lobeswell-recognizedbutpodian
lobesvestigial.
Specimensex::amined. 1,3(fordrawingofendophallus),ColdelMule,2,300m,
Castelmagno,Cuneo,Piemonte,NW.Italy,19-VI-1996,P.CAvAzzUTI leg; 2 (for
DNAanalysis),samelocality,14-VI-1997,P.CAvAzzUTIleg.
6) Calabus(0rmocarabus)f‘aimtaireipe、l,icusCAvAzzuTI,1973
(Figs7,l6)
Malegenitalia. Shapeofaedeagusless specialized than in putzeysianus, with
theapical lobeshort andgradually narrowed towardsapex; ostium lobe largeand
bilobedattip; basalportionofendopha11ussimple,neitherparaligulanorbasal lateral
lobesvisible;prepraeputia11obesrecognizedasapairofhairyweakmembraneousin-
flation; parapraeputia11obesmoderately developedandsymmetrical;praeputia1pad
withapairofhump-likeinflationin frontalpart;podian1obeswideandrobust.
Specimens eMmlned. 1 (for drawing of endopha11us), Colte di Be11ino,
2,800m,Acceglio, Cuneo,Piemonte,NW. Italy,14-IX-1997,PCAvAzzUTl leg;2
