February 1992 J.Mamm.Soc.Japan16(2):71−95
IdentificationandMorphologicalCharacteristicsof
〔ソ(仙・/り/∼りノノ叩J7(バれ/JJ/J∫、苦り/ん川り川.l・べ・ヾん川∫(ゾ//∫.
E.inez andE.evafromtheUSSR,Mongolia,
and Northern and CentralChina
YukibumiKANEKO β/(−/(融、〟/上‘血)ノ再(−り・.凡〔、J〟小耳&7机・‘〟ね′∼.んぜ〟=・〟〔J′∼扉,さ巾・・ 7妻ゑα郁αね〟76玖ノ(砂α乃 (Accepted12July1991)Abstract.Museumspecimensof236red−backedvolesfromthesouthernpart
oftheUSSR,Mongolia,andtheProvincesofInnerMongolia,Hopei,Shansi,
Shensi,Kansu,Chinghai,Hupeh,andSzechwaninChinawerecomparedwith
69specimensofCleth7iononv)S明わcanusfromFinland・Thedistancefromthe
mostp?Sterioredgeoftheupper3rdmolar(M旦)tothemostanteriorpointontheincISOr(I−M3)correlatedpositivelywithagesdeterminedbythedevelop−
mentalstagesofmolarrootsintheyear−rOundsamplesfromFinland・Inthe
samplesfromFinlandandfromeightlocalitiesintheUSSRandMongolia46D
Normore(GroupA),mOStOftheskullsmeasuringI−M3=14・6mmormoredid
nothavealveolarcapsulestoindicaterootdevelopment.Ontheotherhand,in
the samples fromlllocalitiesinnorthern China situated from370to410N,
alveolarcapsulesindicatingrootlessmolarswereobservedeveninskullswhere
I−M3=14.6mmormore.Inthesamplesfrom18localitiesincentralChina
from3lOto390N,allskulls had anI−M30f14.6mm orless,and alveolar
capsuleswerefoundeveninadultfemaleshavingprominentmammaeonthe
skin.AdultanimalslargerthanthesmallestI−M3class(0・3mminterval)of
the adult femalesin the respectivelocalities are dividedinto two groups
(GroupsBandC)bytherelationbetweenI−M3andinterorbitalwidth(IOW)・
GroupBanimalshadlargerI−M3andsmallerIOW,includingtheholotypeof
C7俗eOnWSShanseius,WhereasthoseofGroupChadsmallerI−M3andlarger
IOW.GroupCspecimenscouldbefurtherdividedintotwogroups(C−1andC−
2)basedontherelationbetweenトM3andtai11ength(TL)・Givenanincrease
inI−M3,SPeCimensinGroupCLlhadashorterTL,includingtheholotypesof
Microtus(Eothenonv)S)inezandM.(E.)nux,WhereasthoseofGroupC−2hada
longer TL,including the holotypes of M.(CaYyOnyS)eva,M・(Ca73)OnWS)al−
Cそnous,andCmseoプク研aquilus・GroupAspecimensareidentifiedasCleth−
nonoプ叩S ndbcanusbythepresenceofroots andtheagevariationofmolar
patternsonM3,Whereas,basedontheabsenceofroots,GroupBspecimensare
classifiedasEshanseius,GroupC−1asE.inez,andGroupC−2asE.eva.The
holotype of Minez jeholicuswithunwornM3is
.
demarcationlinebetweenC・n4bcanusandEothenonwsis
. r reviewed. 八一し_、・什・り・心‥=′/け/・刷りり、、叩′∴砧′〃・\‥古砧′′り′ノ”・→ん′′=/′′バ:/∵・血・・′′…り・、 inez;Eothenonv)S eua;EasternAsia.72 〟(7J汀血)
Except for ClethrionoククりS rutilus,Classification of other red−backed voles (Clethrionoプクり惜andEothenonws)fromtheProvincesofInnerMongolia,Hopei, Shansi,Shensi,Kansu,Szechwan,and HupehinChinahasbeenverycomplir Cated.Followlng the description of seven new forms,three classifications havebeenproposed.Hinton(1926)1umpedsixformsintoEvotonv)S(=Cleth− riononws)ndbcanus,eXCludingoneformwhichwasdescribedafterpublication Ofhiswork.Allen(1940)designated thevoles fromInner Mongolia,Hopei, andShansias C.n4bcanus,andrecognizedtwo orientalvolesasEothenonり′S inez from Shansiand Shensi,and E.eva from Shansi,Shensi,Kansu,and Hupeh.Corbet(1978)recognizedthreeorientalvoles,E.shanseiusfromHopei andShansi,E.inez andEeva.
MorphologicalvariationanddistributionofredLbackedvoleshavenever
been clarified for northern and centralChina for three reasons.First,the
precedingclassificationshavenotbeenuniformlyappliedbyothertaxonomists
(Ellerman,1941;Tokuda,1941;Ognev,1950;Ellerman&Morrison−Scott, 1951;Shou,1962;Gromov&Polyakov,1977;Honackiet al.,㌧1982;Hu& Wang,1984).Secondly,identification keys are very simple and only crude distributionmaps are available(Corbet,1978).Finally,manyfaunalstudies
recently publishedin China haveneither mentioned the classification system
adoptednortheidentificationmethodused(Li,1965;Tan et al.,1965;Zhao, 1978;Chenetal.,1980;Zheng,1982;Liueial.,1984;Zhang,1984;Laing& Zhang,1985;Zhang,1987;Wang,1990;Zheng&Zha丘g,1990). Inthepresentpaper,Icompareexternal,Cranialandmolarcharacteristics Ofred−backedvolesfromtheUSSR,Mongolia,andnorthernandcentralChina WiththoseofC.ndbcanusfromFinland.Ipresentmorepracticalcriteriato identifythevolesintheseregionsandshowthedistributionsofthespeciesin detail.
Materials and M:ethods
A totalof305specimens were examined:69specimens of C.n4bcanus COllectedfromKilpisjarvi,Finland(69003′N,20049′E)fromFebruarytoNovem− ber,1983,by A.Kaikusalo;and 236 specimens of Clethriononws and Eo−
thenonys from the southern part ofthe USSR,Mongolia,and northern and SOuthern China(the Provinces of Hopei,Shansi,Shensi,Kansu,Chinghai, Hupeh,and Szechwan)housedin the followinginstitutions:the British Museum(NaturalHistory)(BM),U.K.;theMuseumofComparativeZoology, HarvardUniversity(MCZ),theAmericanMuseumofNaturalHisory(AMNH), theUnitedStatesNationalMuseumofNaturalHistory(USNM),andtheField Museum of NaturalHistory(FMNH),U.S..A.;theInstitute of Zoology, Academia Sinica(ASZI),Beijing,and the Northwest PlateauInstitute of Biology,AcademiaSinica(ASNPIB),Sining,China;andtheYamashinaInsti− tuteforOrnithology(YIO),Chiba,Japan.
73
見通−∂α戊edlわJβざカⅥ∽蝕ねγ柁A5ぬ
COllected,muSeurh,and registration number of allspecimens examined are
listedinAppendixA.Latitudeandlongitudeofthelocalitiesweredetermined
fromgazetteersinZhuang(1983)andSu(1984).AppendixBincludesremarks
On the typelocalities of Cmseon砂S Shanseius Thomas,1908and Ca7yOnWS
aquilusAllen,1912.Some ofthesespecimenswerepreviouslydescribed and
identified(Thomas,1908a,b,1909,1910a,b,1911a,b,C,d,1912a,b;Clark&
Sowerby,1912;Hollister,1913;Allen,1912,1924,1940;Hinton,1926;Howe11, 1929;Kuroda,1939;Ellerman,1941;Ellerman&Morrison−Scott,1951;Tan
et al.,1965;Corbet,1978).
Taillength(TL)was recorded from the skinlabel.The presence of mammae and the number of nipples were checked for female specimens.
Incisor−3rdmolarlength(トM3)andinterorbitalwidth(IOW)weremeasuredon undamagedskullstothenearestO.1mmwithadialcaliperbytheauthor.The
ILM3isthedistancefromthemostanteriorpointontheincisorto themost posterioredgeoftheM3.TheIOWistheshortestdistanceofthefrontalbones
between orbits.The condylobasallength(CBL)was not applied,because
manyskullsweredamaged.
ThedisappearenceoftheM2alveolarcapsulewasrecordedfortheskulls
Of C.ndbcanus from Finland and allmuseum specimens(Prychodko,1951; Koshkina,1955).Three stages were recognized:a StrOng arChed alveolar CapSule;anintermediate stage,in which the capsuleis slightly swollen;a CapSuleflattenedandreducedtoastraightline. TheageofskullsofC.7頑canusfromFinlandwasdeterminedaccording tothedevelopmentalstagesoftheirmolarrootsfo1lowingthecriteriaofAbe (1976).Sixageclassesweredetermined:Ⅰ+ⅠⅠ,III+IV,Ⅴ,lessthan33%of rootratio(ⅤⅠ),from33to60%ofrootratio(VII),and61%ormoreofrootratio (ⅤⅠⅠⅠ).
Adultmuseumspecimensweredeterminedashavingasizegreaterthanthe
SmallestI−M3class(0.3mminterval)offemaleswiththepresenceofmammae Ontheskinintherespectivelocalities(Fig.3). EnamelpatternsofocclusalsurfaceonMl,M2,andM3weredrawnfor museumspecimensfromclose−uppicturesofthemolarrows(1.75×magnifica− tion)usingastereomicroscope(SMZ−10)producedbyNikonat6.6×magnifica−tionafterthepicturesweretakeninthemuseumwithanaccessoryclose−up
lensattachedtoanOlympuscamera.EnamelpatternsonM3wereclassifiedfirstintofourtypesaccordingto
Abe(1982).Type4+5iscomplexwiththreereentrantanglesonthebuccal Side;type6hasthreesalientanglesonthebuccalside,aShortposteriorloop,and a confluent dentalisthmus between triangles;Type9has three salient
anglesinwhichtheinternalposteriorenamellamellaisstraightto convex;
Type7+8hasthree・Salientangles onthebuccalside with slightly concave internalposteriorlamella,WhichType9doesnot.Type9andType7+8were
furtherdividedintoType9′andType(7十8)′,reSpeCtively,inwhichaconfluent dentalisthmusbetweenalltrianglesispresent.
74 助乃(戒0
TheoccurrenceofasmallextraposteriorLinternalangleonMlandM2
wasalsochecked.Theposterior−internalenamellamellawasclassifiedbasedOnthepresenceorabsenceofaprominentangle.
Results l.Vdriation〆Clethrionomysrufocanus InthelargesampleofC.ndbcanusfromFinland,I−M3correlatespositive− 1ywiththesixageclassesdefinedbyrootdevelopment(r=0.796,U=67,P< 0.001)and may be used as an approximateindicator of age(Fig.1).ThealveolarcapsuledisappearsonceI−M3reaches14.8mm.Amongthe16skulls inthe13.0−14.7mmI−M3range,OneSkullhasanintermediatecapsulestage (6.2%);amOngthelOskullsinthe14.8−15.OmmI−M3range,Sixskullshavean intermediatestageorlosethecapsule(60.0%);andwhereI−M3=15.1mmor more,allskullslose the capsule.Root development(ⅤⅠ,VIIand VIIIage classes)isobservedinthreeamongfourskullswithoutcapsulesinthe14.8−
−●鴎ゝ
・・−1W・
A ●● ●ム ●現品・
○。。。§13 14 15 16 17mm
I−M3Fig.1.Relationship betweenincisor−3rd molarlength(I−M3),age Classes(Ⅰ−ⅤⅠⅠⅠ),and
developmentofalveolarcapsuleinC.n4bcanusfromFin1and.SymboIs:CapSulepresent= OpenCircle;CapSuleabsent=Closedcircle;andintermediatestage=Opentriangle.
凡、lナーみ汀ん−√Jlt)ん・∫ノねJJJ長打Jりフナ.▲h山 75 15.OmmトM3range(75%),andin4lof43skullswithoutcapsuleswhereI−M3= 15.1mmormore(95.3%).Consequently,theabsenceofthecapsuleisagood indicatorforrootdevelopmentinskullswhereI−M3=15.1mmormore. ThedisappearenceofthealveolarcapsuleinrelationtoトM3iscompared amongthesamplescollectedindifferentmonths(Fig.2).In C.glanolus,the
developmentofmolarrootsisknowntobemoreretardedintheautumn−born
cohortthaninthespringorsummercohort(Lowe,1971;Zejda,1971).The capsule disappearsin specimens collectedin May.whereI−M3=14.6mm or more,WhereasthecapsuleisabsentinspecimenscollectedinJulyandSeptem− berwhereトM3=15.5mmormore,Whichsuggestsaretardationofmolarroot formation.Inthefollowingcomparisonofcapsuledevelopment,theabsenceofcapsulesindicatingrootdevelopmentisspecifiedintheskullclasseswhere
I−M3=14.6−15.2mm or more,and the museum specimens aregivenin two collectionperiods:thefirsthalfoftheyear(JanuarytoJune),andthesecond halfoftheyear(JulytoDecember). 二一.=/J・高//■り/イ.、小一九仇汁り小叫=≠.ヾ・り//ん一川Jh・/・!/−///、J二\ヾ化l/‖肘′//′′、・/′/′/ ∧「(〃軸げノブのJd艮びJビr〃C/J/JJ(7 PlotsofトM3,theabsenceofalveolarcapsuleandthepresenceofmammae
ontheskinhavebeenarrangedfromnorthtosouthforthemuseumspecimens
examined(Fig.3).Fiftyrtwo adult females with mammae on the skin are
foundamong230specimensfromLocaIitieslto37.SkullsfromLocalitiesl
to19exhibitI−M3=14.6mmormore,WhereasthosefromLocalities20to37 haveI−M30fless than14.6mm.The absence of capsulesindicating root developmentis foundin skullsin whichIrM3=14.6mm or more;allsuch
Fig.2.Monthlyvariationoffrequencydistributionsofincisor−3rdmolarlength(Ⅰ−M3)with development of alveolar capsulein C.ndbcanus from Finland.One squareindicates one SpeCimen.SymboIs:CapSulepresest=OpenreCtangle;CapSuleabsent=Smallclosedrhom− buswithinasquare;andintermediatestage=Ⅹ−1etterwithinasquare.
76 ん7仇−ん) 3 l. ..l.. .l.▲..l...l Z ⑧ 柑¢ ⑫ 皇 日誌指頭 訪日釘⑳逃避 118 128 1鳩 隠1mm トM3 11.O 128 1J..6 16,lmm トM3 Fig・3・Geographicalvariationoffrequencydistributionsofincisor−3rdmolarlength(トM3)
With development of alveolar capsulein samples of red−backed voles from the USSR,
Mongolia,andnorthernandcentralChina.Somelocalitiessituatedatthemorewestern。r easternregionsarearrangedparalleltoshowtheapproximatelysimilarlongitude.Number− edlocalities(1L37)refer to Appendix A.A numberedlocalitywith a circleindicates
Samples collected fromJanuary toJune,While an underlinedlocality numberindicates SamplescollectedfromJulytoDecember.Anunmarkedlocalitynumberindicatessamples
COllectedfromJanuarytoDecember.Theholotypesareabbreviatedasfo1lows:Mimtus
inezjeholicus=Mij;Micrvtus(Eothenonys)inez=MEi;C7WeO7町SShanseius=Cs;Mimtus
(Ca7yOnWS)eua=MCe;MicYVtuS(Eoihenonv7S)nux=MEn;Micntus(Cb7yOnVLS)alcinous=
MCa;C7WeOnySaquilus=Ca・ThebrokenlineindicatesanI−M30f14.6mm,fromwhich
the alveolar capsulehasdisappearedin C.ndbcanus fromFinland(Fig.1).Onesquare
indicatesonespecimen・SymboIs‥CapSulepresent=Verticallineswithinasquare;CapSule
absent=OpenSquare;intermediatestageofthecapsule=Ⅹ一1etterwithinasquare;andan
77 風姻−∂α虎βdl句Jぉ力℃肌艮び′eγ乃Asわ
specimenswerecollectedfromLocalitieslto8・Ontheotherhand,CapSules
indicatingalackofrootdevelopmentareobservedinotherskulls,including34
skullswhereI−M3=15.2mmormore(Localities9to19),andin36adultfemales
withfnammae(Localities9to37),Thus,thepopulationsfromLocalitieslto
8differ from those from Localities9to37with regard to the presence of
alveolarcapsulesandwillbereferredtohereafterasGroupA・
AsidefromGroupA,tWOOtherpatternsofI−M3androotdevelopmentare
foundamongspecimensfromLocalities9to37(Fig:3)・OnegroupispresentinLocalities9to19whereanimalshavealongトM30f14,6mmormore,and
theothertypeisfoundinLocalities20to37whereanimalshaveashortI−M3
0flessthan14.6mm.Fig.4showsthatadultanimalsfromLocalities9to37
canbedividedintotwoenclosuresbasedontherelationshipbetweenI−M3and
IOW,thoughallspecimensexaminedarenotclassifiedbythisrelationship・
1る mm12
13
14
15
ト仙3 Fig.4.Relationshipbetweenincisor−3rdmolarlength(I−M3)andinterorbitalwidth(IOW) insamplesofred−backedvolesfromLocalities9to37・NospecimenisgiveninLocality24 becauseofIOWbeingdamaged・Adultanimalsenclosedbybrokenlinesareclassifiedas GroupsBandC.Fornumberedlocalities(10−34),SeeAppendixA,andforabbreviationsof theholotypes,SeeFig・3・SymboIs:adultindividulal=Closedcircle;individualotherthan adults=OpenCircle.78 ん7〃し−ん)
Hereafterskn11swithlongI−M3andnarrowIOWarereferredtoasGroupB, and those withshortトM3■as Group C.Group B consists of adult animals
COllectedfromLocalities9to19,eXCludingLocalitieslOand14.GroupCis COmpOSedofadultanimalscollectedfromLocalities17,23,26,29,31and34.
AdultanimalsfromLocality17(Kolanchowpopulation)belongtobothGroup
BandGroupC.Inspiteofnotbeingclassifiedasadultfemalesduetoalack
Of mammae on the skin,SpeCimens from LocalitieslO and14,and from Localities21,25,30,32,33and35−37wereclassifiedasbeinginGroupsBand C,reSpeCtively,becausetheyareenclosedineitherarea andseparatedfrom each other.AIso,SpeCimensfromLocalities20,22,27and28areplacedin
GroupCbecausetheyarelocatedneartheenclosureofGroupC.
Group C can be further dividedinto two groups(C−1and C−2)by the relationshipbetweenI−M3andTL(Fig.5).WiththeincreaseinlengthofI− M3,GroupC−2specimensfromLocalities24,26,27,29,and31−36havealonger TLthanthoseofGroupC−1fromLocalities17,20−23,25,28and30.Although OneSkullwithshortトM3(Locality24;Fig.3)isnotplottedinFig.4because
the frontalis broken and theIOWis not available,it has alonger tailin
COmparison toI−M3andis classified as beingin Group C−2(Fig.5).One SpeCimenwith the missing tailand shortI−M3(Locality37;Fig.3)is not
illustratedinFig.5,butenteredinGroupC−2becauseoftheresemblanceofthe molarpatternontheM呈tothatoftheholotypeofMicntus(CalyOククりLS)alcinous
(minFig.8).
The fo1lowing four holotypes are plotted as adult animals:Cmseonws
Shanseius(BM8.8.7.85)inGroup B(Fig.4),M(E.)inez(BM9.1.1.188)in GroupC(Fig.4)andinGroupC−1(Fig.5),andM(C.)eva(BMll.2.1.223)and C7WeOnqySaquilus(MCZ7190)inGroupC(Fig.4)andinGroupC−2(Fig.5).On theotherhand,thefollowingthreeholotypesarenotshownasadultanimals. M(E・)n拡(BMlO・5.2.79;Locality30)andtheseriesfromthesamelocality areplacedintheadultsectionofGroupC(Fig.4)andtheninGroupC−1(Fig. 5)・Becausetheholotypeof〟.(C.)alcinous(BMll.9.8.136;Locality35)is locatedneartheenclosureofGroupCandtheseriesfromthesamelocalityare enclosedintheadultsectionofGroupC(Fig.4),theholotypeisclassifiedfirst asGroupCandthenGroupC−2(Fig.5).M.inezjeholicus(YIO857;Locality
13)iscategorizedasayoungspecimenbecausetheskullhasunwornM墨SOn
bothsides.Theholotypewascollectedtogetherwithadultspecimensclassi− fiedasGroupB(Fig.4),andtheholotypeissubsequentlyconsideredtobein GroupB. Somefurthercharacteristicsareexhibitedafterclassifyingspecimensas GroupsA,B,C−1,andC−2・AccordingtothefrequencyhistogramsofTL(Fig. 6),GroupsAandBdonotshowacline,thoughtheeasternspecimensofGroup B(Localities9,13,14,and16)haveaslightlylongerTLthanthewesternSPeCimens ofthe samegroup(LocalitieslO−12and17−19).The southern
formofGroupC−1(Locality30)hasaslightlylongertail.GroupC−2speci− menshavealongerTLthanthoseofothergroups,andGroupC−2specimens
79 凡1(′−わ爪、れ・(/11−ん−∫.わt川∼艮J5チトり∼.1ゴJ(7
havingaTLofmorethan41mmshowaclineincreasingfromnorthtosouth・
Regardingenamelpatternsofocclusalsurface on M旦(Fig・7),Group AskullswithshortI−M3areofType6;SkullswithlongI−M3areofTypes7+
8and9(a,b,andcinFig.8);andthecomplexformofType4+5appearsin smallnumbersindependent ofI−M3(6/66=9.1%;din Fig.8).In Group B, almosta11largersizedskullsareofTypes7+8and9(51/81=63.0%;eandf inFig.8),andsmall−SizedskullsareofType6,aSfoundinGroupA・Type4十 5,however,OCCurSmOrefrequentlyinGroupB(11/81=13.6%)thaninGroupAspecimens,and seven skulls have a confluent dentineisthmus between all
triangles:OneisofType(7+8)′andsixareofType9′(ginFig・8)・InGroup C−1,Type7十8predominates(19/33=57.6%;iandjin Fig.8),followedby Type4+5(12/33=36A%;hinFig.8),andanabsenceofType9specimens・In
GroupC−2,Onthecontrary,theType9isthemostcommon(35/50=70・0%;kミ
1,andminFig.8),followedbyType7+8(11/50=22.0%),andveryfewType4+ 5specimens(3/50=6.0%).TheenamelpatternofGroupC−1(h,i,andjinFig・ 8)showsaslightlylongerposteriorloopthanthatofGroupC−2(k,1,andmin Fig.8). 2934/
▼▼ G「0UPC−2
云 、て・・・ 0ノーー 26MEn
MEレ占こ .2J、、、≡9.一浩昏、刀2217
0 25 Gb O17 20 ノ・ ◎諏 訪0 00 刀/ ‘030Group
C・111
12
13
14
15mm
トM3
Fig.5.Relationshipbetweenincisor−3rdmolarlength(I−M3)andtaillength(TL)insamples ofred−backedvolesfromLocalities17and20to37.Fornumberedlocalities(17−37),See AppendixA,andforabbreviationsoftheholotypes,SeeFig・3・80 んJ〃t、如 I t l. 1 l
三____
. く2_∴_』___
l l l l l 3[R[
R#円
12 gd⊃0﹂り≡三
19 22 e 23 025 ∩自コ 28JR献ME∵
0 3 払 R ? U 芯231
(つ刀 。卜 漣 17 32 41幻 65汀.m TL 32 4150 伍mm TL Fig.6.Geographicalvariationoffrequencydistributionsoftaillength(TL)insamplesof red−backedvolesfromtheUSSR,Mongolia,andnortherhandcentralChina.Fornumbered localities(1−37),SeeAppendixA,andforabbreviationsoftheholotypes,SeeFig・3・Groups A,B,C−1,andC−2areclassifiedinthetext.81 凡J」柚木−‘/l′′′1)ん,り)p〃J且7∫/(ー7J▲心血 l. ‥.l..‥l l. −l l 口旬Pe6 回吋Peん5 四句Pe7†8 切旬Pe(7◆房 ■可Pe9 ロ句Pe9 , 2 34 5 くd⊃○占 8“ + ﹁ − ﹁ 9 3 ‘56 1 ▲l・l l 10 11 皿 n12 =I 占17 18 19
阿lm
□ 7123 5 12 22 2 TUdコ0占 払 巫27引 ?Udコ2り −■ 29 32 34 36 …… ノ・て三 1軍 146 16j mm トM3 11.0 12β 1昭 161mm l1ロ トM3 Fig.7.Geographicalvariationoffrequencydistributionsinthepatternsof3rdmolarin relationtoIrM3insamplesofred−backedvolesfromtheUSSR,Mongolia,andnorthernand centralChina.82 んJ〃し丸) c]
b c
d ef g
厨暫浸嵩灘
屡苫警笥
尿管悼藁簡
虚像望領
h l
J k
l m宏蔓算茶筒
展望蟹磨常■
宴いV・居嬰鰻苓職
庭草泣責慢
居留斧苫笥
Fig・8・EnamelpatternsofuppermolarsinGroupsA(a−C),B(dLg),C−1(h−j),andC−2 (kLm)・Thealveolarcapsuleisabsentina−C,butpresentind−m.a,MCZ21535(Locality 7,CBL=25・1mm,andトM3=14・8mm)withsimplexformonM旦(Type9);b,MCZ21365 (Locality6,CBL=25・9mm,andI−M3=15・3mm)withsimplex form on M旦(Type9);C,BM(NH)12・4・1・81(Locality2,CBL=26・6mm,andI−M3=15・7mm)withsimplexformonM墨 (Type9);d,MCZ21372(LocalitylO,CBL=24.9mm,andIrM3=15.Omm)withcomplexform OnM旦(Type4+5);e,MCZ21373(LocalitylO,CBL=25.5mm,andトM3=15.4mm)with SimplexformonM呈(Type7+8);f,BM(NH)8.8.7.85(theholotypeofC7那eOnり塔Shanseius, Locality19,CBL=25・7mm,andトM3=15・3mm)withsimplexform onM旦(Type9);g, USNM155055(Locality19,CBL=26・7mm,andI−M3=15.4mm)withconfluentdentalspaces Ofsimplex form on M量(Type9’);h,BM(NH)9.1.1.188(theholotype of Mimtus(Eo−
theno刑ツS)inez,Locality17,CBL=23・Omm,andトM3=‡3・7mm)withcomplexformonM墨 (Type4+5);i,BM(NH)10・5・2・79(theholotypeofMicrotus(且)thenonv7S)mLX・,Locality30, CBL=23・4mm,andトM3=13・7mm)withsimplexformonM呈(Type7+8);j,BM(NH)9.1. 1・189(Locality17,CBL=23・7mm,andトM3=14・Omm)withsimplexformonM旦(Type7+ 8);k,BM(NH)11・2・1・223(theholotypeofMimtus(Ca7yOnV7S)eva,Locality26,CBL=22.8 mm,andI−M3=13・2mm)withsimplexformonM呈(Type9);1,MCZ7190(theholotypeof C7那eOnV7Saquilus,Locality34,CBL=23・4mm,andトM3=13・7mm)withsimplexformonM墨 (Type9);m,BM(NH)11・9・8・136(theholotypeofMicYVius(Cb7yOクク研)alcinous,Locality35, CBL=23・4mm,andトM3=14・2mm)withsimplexformonM旦(Type9).
点gd−∂α虎gdlわ/ぉ力℃椚肋ねァ%Asね 83
Theoccurrenceofasmallextraposterio●r−internalangleonMlandM2
(Fig.9)ishigherinGroupC−1(M主,18/33=54.5%;M呈,19/33=57.6%)thani云 GroupC−2(M主,8/50=16.0%:M呈,7/50=14.0%)specimens. Thenumberofnipplesonthemuseumskinswasfoundtobefourpairson GroupsAandBspecimens,andtwopairsonthoseofGroupsC−1andC−2. 3.7bo〃()川/r C()ナ汀/∼/∫わチノ Thereisasharpcontrastinthepresenceorabsenceofalveolarcapsules betweenspecimensinGroupAandthoseinGroupsB,C−1andC−2(Fig.3).In GroupA,theabsenceofalveolarcapsules,indicatingrootedmolars,beginsto developfromIrM3=14.6mm,Type6molarpatternappearsinshortI−M3,and thesimplexformofTypes7+8and9appearsinlongトM3(Fig.7).These Characteristicsarethesameasdescribedfor C.ク頑canus fromFinland and fromnortheasternChinaandnorthernKorea(Ⅹaneko,1990).IconsiderGroup AspecimenstobeexamplesofClethriononv)S n4bcanus. 1■...l....1l■.■ −l.t..l け封22刀 25遡 TUd⊃0占 凸Mユ ロM呈 ■Ml&M2 n E M T lコ 34 N−Ud召占 27 」 11.0 128 仏6 161mm I−M3 11{)128 仏6 161mm トM3Fig.9・Geographicalvariation of frequency distributions()f the occurrence of a small postero−internalangleonMlandM2insamplesinGroupsC−1andC−2.
84 ん7〃し−ん)
Specimens of Groups B,C−1and C−2,in contrast,are allocated to Eo− thenomys.Allhaveapalatalshelfconstructionlike Clethriononv)Sbutlack
rootdevelopmenteveninoldage,aSindicatedbythepresenceofcapsulesin
adultspecimens.AIso,themolarreentrantfoldsarenarrowerthanthoseof Alticola,Which haslittle cementin the folds.This combination of traits conforms to the generic diagnosis of Eothenonys(Hinton,1926;Ellerman,
1941;Corbet,1978). GroupBspecimensdifferinsomecharacteristicsofmolarpattern,TL,and IOW(Figs.4,6,and7)fromthoseofEothenoプク砂S r曙ulusinKorea(Kaneko, 1990).Forexample,thecomplexformofType4+5inM旦islesscommonin GroupB(11/81=13.6%)thanin Y<gulus(69/97=71.1%);thesimplexformof Types7+8and9ism?refrequentinGroupB(51/81=63・0%)thaninrewlus (23/97=23.7%);TLisslightlyshorterinadultspecimensofGroupB(N=26, 夏±SD=35.7±4.8mm,range=28−47mm)thanin those of regulus(N=29, 豆±SD=45.0±2.6mm,range=40−49mm);andIOWisslightlynarrowerin adultspecimens ofGroup B(N=29,X±SD=3.86±0.17mm,range=3.6r4.3 mm)thaninthoseofク聯Ius(N=29,X±SD=4.23±0.19mm,range=3.8,4,6 mm).Theholotypesof C7WeOブタひSShanseiusThomas,1908and Microtusinez
jeholicus Kuroda,1939bothexhibittraitsofGroup B.Ithereforerecognize
Group B specimens asEothenon7yS Shanseius and consider the name M,inez
jeholicusKuroda,1939tobeajuniorsynonymofE.shanseius(Thomas,1908).
TheholotypesofM.(E.)inezThomas,1908andM(E.)n祝方Thomas,1910 areclassifiedasmembersofGroupC−1andappeartobeconspecific.Group
CrlspecimensareheredesignatedasEothenon砂Sinez,withthenameM(E.)
nuxThomas,1910asajuniorsynonym.
Group C−2specimens can be allocated as Eothenonws eva because the
holotypes of M.(C,)eua Thomas,1911,M.(C.)alcinous Thomas,1912,and
CYaSeOnySaquilusAllen,1912areclassifiedasGroupC−2andconspecific.The namesM.(Ca73}0プクV)S)alcinousThomas,1911andC7WeOプ叩SaquilusA11en,1912 arejuniorsynonymsofEothenoクク砂S eVa(Thomas,1911). ThelocalitiesoftheexaminedspecimensofC.ndbcanus,E.shanseius,E. inez,andE.evaareillustratedinFig.10. Discussion TojustifycomparisoOfmuseumspecimenswithClethrionoプ叩n4bcanus from Finland,IwillrevleW the classification history of red−backed voles
(Clethriononv)S and Eothenoクク砂S)from northern and centralChina,With the exceptionofC.ndilus.Sixspeciesandonesubspeciesofthered−backedvoles
have been described:Cmseoクク叩S Shanseius Thomas,1908;Microtus(Eoth−
enonws)inez Thomas,1908;M.(E.)nux Thomas,1910;M.(Ca7yOプ叩S)eva Thomas,1911;M(C.)alcinousThomas,1912;CYaSeOnV)SaquilusAllen,1912; and M.inez jeholicus Kuroda,1939.Severalfaunalstudies(Thomas,1909, 1910b,1911b,C,1912a;Clark&Sowerby,1912)classifiedspecimensunderthe scientificnamesmentionedabove,eXCePtforAllen(1924),Whousedsomeofthe
85
見通−∂αCゑβ♂lわgβざカ℃∽払おγ柁A5ゐ
p莞じ遥OU巴じ巨N遷座可pud§登記ぶり.叫−OSUで鼠sヨOqゴ貪一日OJ苛.d∴鳶pu乱dd∴おSへ︵トC−1︶s遥コ再U01p巴むq∈コu
ん(〃ハム1
86
samenamesbutidentifiedcertainspecimensfromnorthernChinaasEvotonws
(=ClethYiononws)n4bcanuswithoutcomment.
In his famous monograph of Microtinae(=Arvicolinae),Hinton(1926) 1umpedthesesixforms(M.inezjeholicusKuroda,1939isexcludedbecausethe
description was published at alater date)into one subspecies of Evotonws
ndbcanus.Hinton(1926)stated thatsome ofthe morphologicaldifferences
describedforthesetypesaretheresultofagevariationandmanytaxonomists
adoptedhisclassification(Ellerman,1941;Tokuda,1941;OgIiev,1950;Eller− man&Morrison−Scott,1951;Shou,1962;Gromov&Polyakov,1977).
LikeHinton(1926),Howe11(1929)andAllen(1940)classifiedthered−backed
volefrom Hopeiand ShansiProvincesin China as Clethriononv)S n4bcanus
becausethesku1lofC.ndbcanusshanseiusisapparentlyidenticaltothatofC. ndbcanus(Allen,1940).However,their classification differed from Hinton (1926)forvolesfromtheProvinces ofShansi,Shensi,Kansu,Szechwan,and Hupehin China.Howe11(1929)retained one vole of M.(Ca7yOnyS)inez, whereasAllen(1940)separatedbothEinezandE.evafromC.n4bcanusbased onfurcoloration,taillengthandmolarocclusalpattern.
Corbet(1978)1aterrevisedtheclassificationofEothenonws,andrecognized E.shanseius,E.inez,and E.eva from C.ndbcanus because of the absence rooted molars evenin older specimens.Honackiet al.(1982)followed the
ClassificationofCorbet(1978).
Insummary,thesedifferentclassificationswerebasedonwhetherornot
the red−backed vole ofnorthern and centralChina possesses molar rootsin
adult specimens.In this paper,Sku11size where molar roots appearin C.
n4bcanus from Finland was used toidentify museum specimens of the C.
radbcanus group.The absence ofalveolarcapsulesindicatingrootdevelop− mentappearsinskullswithトM3greaterthan15.2mmin C.ndbcanusfrom Finland(Figs.1and2).InthemeasurementtableinHinton(1926),thedental length(=Ⅰ−M3)foroneskullwasgivenas14.4mmwithclosingcementspaces onM2amongfivespecimensfromLapland,Norway,andSweden,Whereasfour
skullswithadentallengthofmorethan15AmmhadrootsonM2.Theabove
observationsaccordwithmyestimationthattheabsenceofcapsulesindicating
rootdevelopmentinC.ndbcanusbeginsinskullswithanI−M3morethan15.2 111111. Theholotypesofshanseius,inez,aquihiS,andevaareadultspecimensthat havecapsulesindicating alack ofroots(Fig.3).Althoughtheholotypesof nux and alcinous do not confirm as adult animals due to a lack of femalespecimenswithmammaein Localities30and35,thesetwo holotypescorre− spondtotheadultsectionofGroupCinthesizeofI−M3andIOW(Fig・4),and thepatternsonM3aresimilarbetweentheholotypesofn拡andinez,andthe samebetweentheholotypesofalcinous,eVa,andaquilus(Fig.8).Furthermore,
all164specimensexaminedfromLocalities9to371ackroots,eVeninskulls
withI−M3more than15.2mm,including36adult females with prominent
87
凡再−ム(7(、h、〟11)/(Yノ)…JJ且び/りTJ.小壷
adult specimens rather thanimmature ones,and the generic allocation of
Shanseius,inez,and evaisEothenonws(Allen,1940;Corbet,1978;Honackiet alリ1982),nOt ClethYiononws(Hinton,1926;Allen,1940).
TheholotypeofMicrotusinezjeholicusisayoungE.shanseius,Although Allen(1940)provisionallyfollowedKuroda’s(1939)identification,hequestioned
thelonger hind foot(17−21mm)for jeholicus compared to E.inez(16mm). Corbet(1978)and Honackiet al.(1982)expressedthesame doubts.Alarge hindfootispossibleinE.shanseius;forexample,thehindfeetmeasure20−21 mmintheseriesofE.shanseiusfromKweihwaCheng(LocalitylO).
In describing Mic7btus(E.)inez and M(E)nzα,Thomas(1909;1910b)
mentionedthatthepatternonMlandM2hasatendencyto develop a small
extrapostero−internalangle,Whichisverydifferentfromthelargeextraangle inE.melan聯Ster.Onthe otherhand,Thomas(1911b)notedthatthe addi− tionalanglesarereducedtoaminutesizeandscarcelyperceptibleinM(C.)
eva,butnotin M(C.)alcinous.Allen(1940)referredto the presence ofthe anglein E.eua eva,but did not mentionitin E.eva alcinous.The postero− internalangleissometimespresentinE.inez andE.eva,butthefrequencyof
theoccurrenceisgreaterininezthaninevaanddoesnotsuggestanygeograph−
icaldifference(Fig.9).
ThisstudyrefinesthedistributionrangeineasternAsiaforC.nめcanus.
The southernlimit of the rangein China was reported differently by Allen
(1940),Corbet(1978),andHonackiet al.(1982),thoughmytaxonomicconclu− Sionsupportstheirclassification.Allen(1940)showedthedistributionasthe Provinces of Shanseiand Hopei,because he considered E.shanseius as a memberofC.ndbcanus.Corbet(1978)notedthesouthernlimitasthesouthern
Ural,Altai,ManchuriaandKorea,andexcludedMongolia,WhereasHonacki etal.(1978)includedMongolia.Fig.10indicatesthatthesouthernlimitlies450
Nin Mongolia,Which accords with the figure of Mallon(1985).Thus,the
Currently understoodlimit of C.n4bcanuslies at approximately 450Nin
Mongolia,and400Nin Korea and northeastern China(Manchuria;Kaneko, 1990),and41.50NinthesouthernpartofHokkaido,Japan(Ota,1984).
On the basis of the specimens examined andliterature references,the
habitatsofC.ndbcanusmaybesummarizedasfollows.Theedgeoftheforest
fo1lowingalongthenorthernedgeofMongoliaiscomposedofmixedforests
SuCh aslarches,firs and pines,aS Wellas oaks and birches(Allen,1940). Tapucha(Locality13)containsaforestofnutpines(」巧nus cembm;Ho11ister, 1913).Hence,the habitats of C.n4bcanus aremiⅩed or coniferous forests, WhicharethesameasthoseinManchuria(Kaneko,1990).
AnunsoIvedproblemconcernstheidentificationofonespecimen(BM8.8. 8.83=OriginalNo.1625)collectedfromtheChiaoChengShan(Locality19)in December,1907.IhavenotlistedinAppendixA;however,Thomas(1908b) identified the specimen as C7ⅥSeOプクり唱 Shanseius.Its capsules have almost
disappearedandthemolarsareabouttoformroots.Hinton(1926)mentioned
〃(J〃一丸I
88
1engthof15.4mmsuggestingaCBLofapproximately25.8mm.Heclassified
aIlthe volesincIuding this specimen from northern and centralChina as
Clethrionoクワ砂S ndbcanus.My examination of this specimen confirms the development of roots andI−M3=15Amm.This specimen and C.ndbcanus fromFinland,therefore,havethesameI−M3size,Whichsuggeststheabsence Ofalveolarcapsulesindicatingrootdevelopment(Fig.1).However,identifica−
tionofthisspecimenas C.ndbcanusshouldbesuspendeduntilwecanobtain
moresamplesofC,r8dbcanusfromthesamedistrict,eSpeCiallybecauseLocal− ity19isisolatedfromtherangeof C.ndbcanusgivenhere.
Regarding the red−backed vole from HopeiProvince,Thomas(1908b) identified one specimen(BM8.8.7.81=OriginalNo.1549)from theImperial Tombs(Locality15)as Cmseoプ叩Sク聯Ius.Allen(1924,1940)recognized all five specimens collected from thatlocality as Evotoクク砂S(or Clethriononws) γゆcanusshanseius,andHinton(1926)agreed.Kuroda(1939)identifiedlarge− Sized voles from Altolian(Locality9)and Mt.Mulei(Locality13)as C.
ndbcanusYqulus,andsmallervolescollectedfromthesamelocalitiesasanew
SubspeciesofMic7VtuSinezjeholicus,WhichareidentifiedhereasE.shanseius. Corbet(1978)andHonackietal.(1982)describedEク聯IusasexistinginHopei
Province and E.shanseius as possibly presentin Hopei.In the present tax− OnOmicconclusion,E.r曙ulusandE.shanseiusdifferintaillength,interorbital
width,andcrownwearpatternsonM3,andthemuseumspecimensfromHopei
Province areidentified as E.shanseius.The demarcationline between the
SpeCiesofY<gulusandshanseiusmaybetheLiao−hePingyuanRiverlocatedin
northeastern China.HabitatsofE.shanseius,basedonthespecimensexaminedandliterature references,includebushes atthefootofatalus−SlideintheImperialTombs (Locality15;Thomas,1908b),brush−COVered valley−bottoms at the Chiao ChengShan(Locality19;Thomas,1908b),1argerocksinorneardensespruce Orlarchwoods,andovergrowngulliesandcanyonsinKolanchow(Locality17; Thomas,1909).Thisvolefeedsuponleavesofyoungplantsgrowingamong rocksunderwhichitburrows(Locality19;Clark&Sowerby,1912).E.shan− Seiusapparentlyprefersrockyareas,aSdoesEク聯hiS(Kaneko,1990).
According to Allen(1940),Corbet(1978)and Honackiet al.(1982),the distribution of E.inezis the Provinces of Shanseiand Shensiand extends to
HopeiProvince(=Nekka orJehol).Apparently HopeiProvince was mentionedasKuroda(1939)referredtoitinhisdescriptionofM.inezjeholicus. However,becausejeholicusisidentifiedhereasEshanseius,thedistributionof
E.inezis now confined to the Provinces of Shansiand Shensi.E.shanseius
and E.inez occur sympatricallyin Kolanchow(Locality17;Fig.4),Which COnfirmsthedescriptionbyThomas(1909),Whonotedthattheformerismuch less common than thelatter.
Thefo1lowingarethehabitatsdescribedpreviouslyandnowidentifiedas
thoseofE.inez.Thespecieswascommonlyfoundinthebottomsofcertain narrow,WOOdedandovergrowngullies,inburrowsinthesoftsoilbeneathbush
89 見通−∂αC烏gdlわJβぶ函桝丘おねγ犯Aざ由 inKolanchow(Locality17;Thomas,1909),andinthelowerregionsofwell− vegetatedloessravinesinYen−an−fu(Locality22;Clark&Sowerby,1912)− ThegeographicrangeforE.evahasbeencitedastheProvincesofKansu, Hupeh,andSzechwan(Corbet,1978)ortheProvincesofKansu,Hupeh,Sze− chwan,and Shensi(Allen,1940;Honackiet alリ1982),because Allen(1940) examinedspecimensfromtheTai−pei−Shan,ShensiProvince(Locality29).I confirmedAllen’sidentificationinthisstudy,andthusthedistributionincludes ?hensiProvince・Withkeysforidentification,HuandWang(1984)reported thatE.evaappearsatZoigeXian(33D12′N,102054′E),PingwuXian(32024′N,104D 30′E)andHeishuXian(32006′N,103006′E),SzechwanProvince,Whicharelocat− edintherangeofE.eva(Fig.10).ThedemarcationbetweenE.inezandeua
isnotclear.Thetwospecieswereco11ectedintheQinglingandDabaShan
Mountains,ShensiProvince(Zheng,1982;Wang,1990),andthesouthernpart ofKansuProvince(Zheng&Zhang,1990).However,becausetheiridentifica−tions have not been verified,further studies should be carried outin these
districts.
ThedistributionofC.n4bcanusisseparatedfromthatofthethreespecies
of Eothenon砂S by the GobiDesert.The range of these three Eothenonys
indicatesthattheyoccurmostlyinthesoutheasternborderofthePalaearctic
Regionin China(EditorialCommittee of China NaturalGeographyin
Academia Sinica,1979)exceptforE.eva,Whichcanbefound alonginto the northernborder(HupehProvince)oftheOrientalRegion. .・l−★/ハー甘ん沃ル机〃小
IgratefullyacknowledgetheassistanceofAskoKaikusalo,Whoprovided
mewithexcellentspecimensfromFinland,andthemuseumstaffwhoallowed
metoexaminespecimensintheircare:IanR.Bishop,BritishMuseum(Natu− ralHistory);MariaE.Rutzmoser,MuseumofComparativeZoology,Harvard University;GuyG.MusserandWolfgangK.LH.Fuchs,AmericanMuseumof NaturalHistory;MichaelD.Carleton,United States NationalMuseum Of NaturalHistory;Bruce D.Patterson,Field Museum of NaturalHistory;WangSung,WangZongren,andQuanGuoqing,InstituteofZoology,Academia Sinica;Zhen Changlin,Northwest PlateauInstitute of Biology,Academia
Sinica;and K.Mommose,R.Kakizawa and N.Kuroda,YamashinaInstitute for Ornithology.Ifurther thank M.D.Carleton and Don Maybin,Kagawa
University,forcheckingandimprovinganearlierdraftofthispaper;M.E・ RutzmoserforlocatingthefieldsnotesofW.R.ZappeyhousedintheMuseum ofComparativeZoology,HarvardUniversity;JackFooden,FieldMuseumof NaturalHistory,forshowingsomereferencesforgazetteersinChina;andM・ Koyama,Tomakomai−nishiSeniorHighSchool,forinformingmeofcollecting locationsinvestigatedbyH.Orii.ThisworkwaspartlysupportedbyaShort TermVisitorGrantfromtheSmithsonianInstitution,theKarlP.SchmidtFund oftheFieldMuseumofNaturalHistory,andan“0verseasScientificGrantfor 1982(62041089)and1983(63043061)”givenbytheMinistryofEducation,Science
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AppendixA
.}−t−・ん拓・直上l・′J′,//仙(/
Locality numbers usedin this study(see Fig.10),locality,1atitude and longitude,mOnthandyearcollected,muSeum,andregistrationnumberofa11
specimensexaminedarelistedbelowfromnorthtosouth.Localitynumbers
l−8referto ClethriononりLS ndbcanus,numbers9−16,17(selectedspecimens), 18and19toEothenoククV}SShanseius,numbers17(selectedspecimens),20,23,25, 28and30toE.inez,andnumbers24,26,27,29and31−37toE.eua. approximately52010′N,104012′E;July, 83−86,15.3.9.35−40. 1.Leestvineechnoya,Irkutsk,USSR 1914;BM14.11.1.78−81,14.11. .,l 2.SyanskMts.,100milesWofLakeBaikal,USSR;5lO42′N,102009′E;June, 1910;BM12.4.1.79−83. 3.Tapucha,Altai,USSR;apprOXimately5lOO4′N,85037′E;August,1912; USNM175454−56,175458,MCZ14322. 4.Tchegan−BurgaziPass,Altai,USSR;49O38′N,89026′E;July,1912; USNM175453. 5.Katon−Karagai,Bukhtarmo River,Altai,USSR;49010’N,85038′E;No− vember,1927;BM28.6.19.41/August,1930;BM28,6.19.40,28.6.19,42・ 6.45milesNEofUrga(=Ulan Bator),Mongolia;48021′N,107034′E;July, 1919;AMNH46232,FMNH30525−26,MCZ21364,21370/August,1919; BM29.3.17.145,USNM259911−12,AMNH46047,46049,46237,FMNH 30524,30528−30,30532,30534−35,MCZ21365. 7.15milesNEofUrga(=UlanBator),Mongolia;48003′N,107008′E;Septem− ber,1919;AMNH46305,46314,MCZ21368−69,21535.
8.Sain Noin Khan(=Uyanga),Mongolia;46017′N,102012′E;June,1922; AMNH57483−84,57488,57491,57496,57500−01,57503−06,FMNH30537,
MCZ21366−67.
9.Altorian(=nearSandaoheize),HopeiProvince,China:40048′N,119000′E: October,1935;YIO862−66,890−95.
10.KweihwaCheng(=HohhotShi),InnerMongolia,China;40048′N,11lO42′E; November,1919;AMNH45429,45436(the skinis housedin FMNH as
93 点gd−ゐαdgdlわJぉ力℃∽肋ね用A5ブα
33719),FMNH33717−18,MCZ21372−74/December,1921;AMNH56344. 11.Mts.30miles W of Kweihwa Cheng(=Hohhot Shi),Inner Mongolia,
China;40O48′N,111007′E;May,1912;USNM175527−28,AMNH36896. 12.Mt.Wula Shan,Inner Mongolia,China;40O42′N,109024′E;June,1978;
ASZI21962−78. 13.Mt.Mulei(=Mt.WulingShan),HopeiProvince,China;40036′N,117030′E; September,1935;YIO857(theholotypeofMicYVtuSinezjeholicusKuroda, 1939),858−61,884−89. 14.Xinglong Xian,HopeiProvince,China;40024′N,117030′E;September, 博76;ASZI26643. 15.ImperialTombs(=Qing Dongling),Beijing,China;40006′N,117036′E; September,1907;BM8.8.7.81/Decembe;,1921;AMNH56349/January, 1922;AMNH56347. 16.Shijiayan,Beijing,China;39048′N,115036′E;September,1970;ASZI 24956/September,1987;ASZIfieldnumbers=K5703−04(theregistration numbersareunknown). 17.Mts.12milesNWofKolanchow(=KelanXian),ShansiProvince,China; 38050’N,111025′E;May,1908;BM9.1.1.188(the holotype of Micntus
(EothenonvLS)inez Thomas,1908),9.1.1.187,9.1.1.189−90/June,1908; BM9.1.1.181r86,9.1.1.191−201.TheregistrationnumbersBM9.1.1. 196−201refertoEshanseius,andtheothernumberstoE.inez. 18.Mt.50milesNWofTaiyuan−fu(=TaiyuanShi),ShansiProvince,China; 38023′N,111055′E:December,1909;USNM172591r92,172594−95/Janu− ary,1910;USNM172596. 19.ChaoChengShan(=Mt.GandiShanorMt.NanyanShan),ShansiProv− ince,China;37054′N,11lO30′E;November,1907;BM8.8.7.82,8.8.7.89− 91/December,1907;BM8.8.7.85(the holotype of Cmseonqys shanseius
Thomas,1908),8.8.7.84,8.8.7.86−88,8.8.7.92−93/October,1908; USNM155050−55.RestrictionofthetypelocalityofCmseoプク砂SShanseius isexplainedinAppendixB. 20.Zanhuang Xian,HopeiProvince,China;37036′N,114024′E;June,1963; ASZI23270. 21.Guanshang,ShansiProvince,China;37012′N,111012′E;July,1960;ASZI 16857. 22.12milesSofYen,an−fu(=Yan’anShi),ShensiProvince,China;36025′N, 109030’E;January,1909;USNM155049. 23.QinshuiXian,ShansiProvince,China;35O42′N,112012′E;May,1962; ASZI19488. 24.Near Guru,ChinghaiProvince,China;35036′N,102D36′E;July,1983; ASNPIB88021,88023. 25.Yuanqu Xian,ShansiProvince,China;350i8′N,11lO36′E;June,1962; ASZI19490,19492/July,1962;ASZI19489,19491.
26.Tauchow(=Lintan Xian),Kansu Province,China;34042′N,103024′E: April,1910;BMll.2.1.223(the holotype of Mim)tuS(Ca7yOnyS)eva
抽ノ汀ん1 94 Thomas,1911),11.2.1.224−26/April,1911;FMNH・19082−83/September, 1911;BM12.8.5.49−51,12.8.5.54r56,12.8.5.59/October,1911;BM12. 8.5.52−53. 27.Choni(=JoneXian),KansuProvinc占,China;34O36′N,103036′E;Septem− ber,1925;MCZ24464/October,1926;MCZ24200−01. 28.LonanHsien(=LuonanXian),ShensiProvince,China;34006′N,111006′E; December,1909;BMlO.5.2.87. 29.Taトpei−Shan(=Mt.TaibaiShan),ShensiProvince,China;33054′N,1070 42′E;January,1910;BMll.6.1.55/September,1921;AMNH56380(the Skinis housedin MCZ as21294),56383,FMNH39391/October,1921;
AMNH56369,56371,56373,56375,56377−78,56379(theskinishousedin MCZas21295),56381−82,FMNH39392.
30.Shang−ChouDistrict(=ShangXian),ShensiProvince,China;33054′N,1090 54′E;November,1909;BMlO.5.2.79(the holotype of Micntus(Eo−
thenonqys)nux Thomas,1910),10.5.2.75,10.5.2.81−82,10.5.2.84−86,
USNM171967−69.
31.WenXian,KansuProvince,China;32054′N,104′420E;August,1987;ASZI fieldnumber=K5684(theregistrationnumberisunknown).
32・Fang Hsien(=Fang Xian),Hupeh Province,China;320060N,110’420E; December,1907;MCZ7191,7194. 33.BarkamXian,SzechwanProvince,China;3lO54′N,102030,E;June,1961; ASZI20824−25. 34・Showlungtan,Hupeh Province,China;apprOXimately31036′N,110006′E; May,1907;MCZ7190(theholotypeofCmseonv)SaquilusAllen,1912),7196, BM13.9.13.10.RestrictionofthetypelocalityofCクWeOnyS aquilusis explainedinAppendixB. 35.Weichoe(=WenchuanXian),SzechwanProvince,China;3lO24′N,103030′ E;November,1910;BMll.9.8.136(theholotypeofMicYVtuS(Ca7yOnyS) alcinous Thomas,1911),11.9.8.128,11.9.8.130,11.9.8.133−34,11.9.8. 137−38. 36.Wansonshan(=Mt.WantiaoShan),HupehProvince,China;31012′N,110O 30’E;June,1907;MCZ7189. 37.GoanShidDwe(=nearGuanXian),SzechwanProvince,China;3lOOO’N, 103036′E;August,1934;FMNH45760. AppendixB f&ing the73少eLocali&〆Craseomysshanseius77u)maS,19(娼and
CaryomysaquilusAllen,1912
1.C7WeO〝り′SShanseiusThomas,1908(BM8.8.7.85;Locality19)The typelocality has beengiven as“100miles N.W.of Tai−Yuen−Fu, Shansi(8000’),China”(Thomas,1908b;Hinton,1926;Allen,1940;E11erman&
Morrison−Scott,1951;Corbet,1978;Honackietal.,1982).However,Malcom
見通−∂α戊edl句放り毎朝八弘扉紺柁A該 95
1908b),nOtedthat“On17thNovemberIbeganworkatChao−Cheng−Shan,a mountainoflO,OOOftaltitudesituatedaboutlOOmileswest−nOrth−WeStOfTai− YuenrFu”and“remainedinthisplaceti11December6th,1907”(Thomas,1908b). Therefore,Ijudge“100miles N.W.of Tai−Yuen−Fu”to be“Chao−Cheng−
Shan”.
Accordingto Anderson(1920),“Chao−Cheng−Shang”is a group ofthree summits,thehighestofwhichiscalled“Mo−erLShan”.Thedescriptioncanbe confirmedfromhispicture(Anderson,1920).Theexactlocation of“Chao− Cheng−Shan”or“MoMerLShan”,however,hasneverbeenrestricteduntilnow,
becat!SeCurrentChinesemapsandatlasesdonotreferto theselocalities・
Clark and Sowerby(1912)collected six specimens of Cmseonws shanseius (USNM155050−55),andnotedthatspecimenNo.41(=nOWUSNM155055)was caughtonthesummit ofMo−erh−Shanand thetype specimenwas collected from the same district.On the skinlabelof the specimen(USNM155055), thereisanoteinpencilas“Essentiallyatopotype”.Thus,IfoundMt・Mo− erh−Shan(about37055′N,111030’E;alt.9296ft=2789m)ontheroutemapofthe Clarkexpedition(Clark&Sowerby,1912).
Next,Iidentified the current place name of Mt.Mo−er(h)−Shan as Mt. GuandiShanorMt.Nanyanshan(37054′N,111030′E,alt.2831m;AltasPublica− tionCompany,1979;Zhuang,1983).Theirequivalenceaccordswiththelati− tude,longitudeandaltitudeofthemountain,therunningdirectionoftheriver, andthenameofnearbytowns. 2.CmseonwsaquilusAllen,1912(MCZ7190;Locality34) Thetypelocalityhasbeengivenas“Showlungtan,Hupeh,China”(Allen, 1912,1940;Hinton,1926;Ellerman&Morrison−Scott,1951).“Showlungtan”
wasmarkedonahandwrittenmaphousedintheMCZ,Whichwasdrawnby
Walter R.Zappey,the collecter of this holotype.Current Chinese atlases, however,donotreferto“Showlungtan”or“Haso−lung−tang”,Whichwasnoted byWilson(1913)whowentto HupehwithMr.Zappey.TheLocationofMt. WantiaoShan(=Mt.Wansonshan:Allen,19120rMt.Wanchaoshan:Wilson, 1913)andtherunningdirectionoftheJiudaolingHeRiver,OneOfthebranches oftheHanShuiRiver,aCCOrdonbothZappey’smapandtheHupehProvince map(scale=1:1,100,000;Altas Publication Company,1982).On the other hand,thelocation of Hsienshan Hsien(=Ⅹingshan Xian)and the running
directionoftheChangJiangandXiangXiRiversaredifferentonbothmaps.
BecausethelocationofZappey’s“Showlungtan”coincidesapproximatelywith thatofHongpingontheProvincemap,Irestrict“Showlungtan”to approxi− mately31036′NandllOOO6′EusingthelocationofHongplngglVenbytheAtlas