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群馬県立自然史博物館研究報告(5) : 1-8, 2001 1
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OriginaIArticleZ~〆、ゴ、ノーゾLダヘゾー、ゾヘダー、ノー、ゴーL/、
PreliminarynoteontheMiocenenightlessswanfromtheHaraichiFormation,
TomiokaGroupofAnnaka,Gunma,Japan.
MATSUOKAHiroshige',NAKAJIMAHajime2,TAKAKuwAYujisandHAsEGAwAYoshikazu3
Ⅲ、幼α”〃E"jq/Geo/ogya蛇騨j梅"E池jqgMG"""EScノz“jQ/Scjg押蝿幻“0酌z”E芯地' fKy“0,“6-節“ノ""z
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3Gz"z糀α暇"se""zQ/Mzj"趣ノHMO7y: Z6再一Z,K""娩郡”釦α,""2"",G"""ea,37D-a345jノ”α刀
Abstract
Nearlycompleteskeletonofaflightlessswan,namedthe"AnnakaShort-wingedSwan",
wasfoundfromtheupperMiddleMioceneHaraichiFormation(11-12Ma),Tomioka
GroupinAnnakaCity,GunmaPrefecture,Japan. Thewingelementsarereduceddistal-
ly; onthecontrarythehindlegchangedlittlefromthevolantancestor, Theskeletonand
livingpostureof 、$AnnakaShort-wingedSwan"areproposedinthisreportthroughthe
preliminaryinvestigation.
KeyWords:GunmaPrefecture,Annaka,TomiokaGroup,Miocene,Flightlessbird,Cygnini
Startlingly, the fossil bird isa flightless species
belongingtoTribeCygnini,especiallyswans. Aswell
astheplotopteridswhichprosperedintheEoceneand
Oligocene (e9.OlsonandHasegawa, 1979, 1996), the
occurrenceofsuchanobviouslyflightlesscoastalbird
fromtheMiocenesignifies thetransitionofmarine
zoogeography.AIso,thefactthattheskeletonisnear-
lycompleteissignificantforstudyingavianflightless-
ness, afrequentpatternofevolution.
Adetailedosteologicaldescriptionandtaxonomical
conclusionswillbepresentedinfuturepublications,as
thespecimenneedsfartherpreparation. Herewepro-
videapreliminaryreportonthefossilflightlessswan.
INTRODUCTION
OnJanuaryl, 2000, thefirstdayofthenewmillen-
nium, juniorauthorHajimeNakajimadiscovereda
fossil vertebrate skeletonexposed on theMiddle
MioceneriverflooroftheUsuiRivernearHaraichi,
AnnakaCity,GunmaPrefecture(Fig.1).Forreinforce-
mentaflat resincoatingwasappliedfor removal.
Afterthat, theslabwaspreparedfromthebottomside
verycarefullyintoarelief-likespecimenbyHN. Sub-
sequently, itwasfoundtobeanalmostcompleteskele-
tonofa largebird, oneofthemostcompleteavian
fossilseverfoundinJapan.
Thefindingof this fossil hadverygoodtiming,
becausel49thRegularMeetingofthePalaeontological
SocietyofJapan (June24-252000) organizedbythe
GumnaMuselnnofNaturalHistorvdirectedbvY.
Hasegawawasabout ready toopen.Theauthors
decidedtopresentatalkonthisfossilatthismeeting.
Weinvestigatedinahurry, andtheperspectivewas
read. Thisisanoteonthespecimenthatwasreadin
themeeting.
SYSTEMATICS
ClassAves
OrderAnseriformes
FamilyAnatidae
SubfamilyAnserinae
TribeCygnini
aff.Mega/odytessp.
受付苫2001年1月10日,受理:2001年3月16日
MAT、suoKAHiroshige,NAKAJIMAHajime,TAKAKuwAYujiandHAsEGAwAYoshikazu2
SPecimen. ---Skeletalelementsinrougharticulation,
scatteredhorizontallyacrossaslabof70cm×maxi-
mum40cm(Fig.2, P1. 1).Clearlyrepresentingone
animal. Theelementspreservedare: cranium,man-
dible, incompletesternum, 9ribs, 16cervicalandthor-
acicvertebrae,pelvislackingonlypubisbranches,2
caudalvertebrae, leftscapula, rightandlefthumeri,
rightandleftulnae, right(?)radius, leftcarpometacar-
pus,rightandleftfemora,rightandlefttibiotarsi,right
andleftfibulae, lefttarsometatarsus, andsomeminor
bones.
Becauseit isdifficulttoassigneachvertebratothe
cervicalorthoracicinthepresentconditionofspecimen,
wehavenotreachedafinalassignment. At least,no
distalcervicalvertebraeareincluded. Possibly,more
thanseventhoracicvertebraearepreservedinthefossil.
That is, however, contrarytothefact thatCygnini
membershaveseventhoracicvertebraeallteriortothe
synsacrum. The right tibiotarsus isbroken in the
middleoftheshaft,andismissingthedistalhalf.
Local岬---NearHaraichi, AnnakaCity, Gumna
Prefecture,Japan; riverflooroftheUsuiRiver,about
600mdowntheriverfromtheNakahasiBridge.
Discoveryb ---ByH.NakajimaonJanuaryl, 2000.
HoriZonand theage. --・Massivesiltstoneof the
HaraichiFormation,TomiokaGroup. ThelateMiddle
Miocene (approximately ll-12Ma). Benthic for-
aminifera(KanekoandNomura, 1998)andlithofacies
indicatethepeiagicdepositionalenvironmentforHarai-
chiFormation. AgedeterminationfollowedNomura
andOhira(1998).
Measurements. ---Becausehalfofbonesarecovered
bymatrix, itisimpossibletogivefinalmeasurementsof
manybones.Asafirstapproximation,werecordedthe
Figurel. LocalitymapsfortheMiocenefossilbirdfromAnnaka,Gunma,basedonthel :25,00Oscaletopographic
mapoftheGeographicalSurveyInstituteofJapan, "Tomioka"and"Matsuida".
PreliminarynoteontheMioceneflightlessswanfromtheHaraichiFormation,TomiokaGroupofAnnaka,Gunma, Japan 1
謬曾汀1um
=
‘ andi
クⅦ″八〃咄棚〃
『
馴、~~
③Cd、V夕
10Cm
Figure2. Identificationsofeachskeletalelementofthefossil (ontherightslab),andthepositioninthebirdskeleton(inthe
leftbird,blackedbones). Abbreviationsare; d,distal ; l, left;m,middle; p,proximal ; r,right/CdV,caudalvertebra;Cm,
carpometacarpus;CV,vervicalvertebra;Fb,fibula;Fm,femur;H,humerus;R, radius;Tm,tarsometatarsus;Tt,tibitar-
sus;TV, thoracicvertebra;U,ulna;V,vertebra; cervicalorthoracicunknown.
maximumlengthofthemainelements. Themeasure-
mentsareasfollows.
Cranium,maximumlengthofthepreservedpart: 95
nlln・
Mandible,greatestlength: 105mm.
Pelvis,greatestlength(withoutpubis) : 200mm.
Pelvis, lengthalongthecenter: 170mm.
Sternum,maximumlengthofthepreservedpart : 105
1,,1.
Scapula(left),GreateSt length: 108mm.
Humerus(right),greatestlength: 220mm.
Humerus(left),greatest length: 218mm.
UIna(right),greatestlength: 125mm.
Ulna(left),greatest length: 124mm.
Radius(right?),greatestlength: 115mm.
Carpometacarpus(left),greatestlength: 66mm.
Femur(right),greatest length: 97mm.
Femur(left),greatestlength: 94mm.
Tibiotarsus(left),greatestlength: 200mm.
Tibiotarsus(left),axial length: 190mm.
Tarsometatarsus(left),greatestlength: 110mm.
humeruslength(P1.2 3). Volantbirds, ingeneral,have
ulnaewhicharealmostequivalenttoorlongerthanthe
humeri,andshorteneddistalelementsarearemarkable
character inflightlessspecies. Theothercharacters
suchas; inhumerus,muchsmallerdistalendthanthe
proximalend, flattenedproximalend,andstraightand
slendershaft (Pl.2-2,3) ; Iowcarinaof sternum(Pl.
2-6) ; shortenedandslendercarpometacarpus(P1. 2 3) ;
narrowandsmallscapulawithreducedacromion(P1.
2 7),etc・ alsoindicatetheflightlessnessinthisfossil
bird.
Thepelvis(P1.2 8)mayalsoshowsaneotenicstateas
wellasthewell-knownexamplesinupperhalfofthe
body. Firstofall, themorphologyofthepelvisofthis
birdisatypical. Judgingfromthesizeofthefemur,
whichindicatesthetrunksizereasonablvwell,wewould
expectamuchlongerpelvisforthisbird. Theother
pointwefindpuzzlingisthenumberofproximalcer-
vical/thoracicidentificationofvertebrae. Itseemsthe
fossil includesmorethanseventhoracicvertebrae、but
theresult iscontrarvtotheseventhoracicvertebrae
foundinCygninibirdS.
Toexplainthetwoenigmasclear,wehypothesized
thefossilpelvisasa@Gmaturedbutunfused"one, i.e.,the
shortpelviswasperhapsprimitivethanthelongerpelvis
whichconsistsofmorevertebrae, thoracicvertebrae
anteriorandcaudalvertebraeposterior, but is inour
fossil lackingsomeunfusedonesonbothends,beinga
Strikingfeaturefortheecologyoffossi l bird
Themostconspicuousfeatureonthisfossilbirdisthe
evidenceofflightlessness, showingextremeadaptation
foraquaticmodeof life,withnolongdistancemigra-
tion. Forexample, lengthofulna isonly55%of
MATsuoKAHiroshige,NAKAJIMAHajime,TAKAKuwAYujiandHAsEGAwAYoshikazu4
tibiotarsus,andafemurandproximalpartsofhumerus
havebeenreferredtothisspecies. However,according
toHoward(1988),theholotypictibiotarsusisl51.9mm
Iong in axial length. The :!Annaka Short-winged
Swan'',whichhasl90mmaxiallengthoftibiotarsus, is
much larger than". wzo7Zio"", andbelongingtoa
different category. Our investigation is tooprelimi-
narytoassignthe"AnnakaShort-wingedSwan" to
particulargenus.Wehavenotexaminedthespecimens
of〃“z/0の花s”0〃o""jatanyrate,andstillneedto
examineotherflightlessanseriformes.
neoteniccondition. Thishypothesisexplainsthenot
chedcaudalmarginoffossilpelviswithshortsynsacral
caudalvertebrae・ Forfurther investigations,wemust
preparetheventral surfaceoffossilpelvis. AIsofar
therpreparationofthevertebraeisneeded.
ldentificationandSystematics
Thisfossil isthatofalargeanseriform・ Further-
more, inadditiontothecharacteristicproportionsof
widewings(eventhoughit isreduced,thelonghumerus
suggests increasedwingspan), skeletal elementshave
characteristicswhichallowustoidentifythefossilbird
asaswan. Wingelementsandpelvisshowunusual
morphologiesashavebeenmentionedabove,butsuch
modificationsareprobablvneotenicandfunctionaldif
ferencesreflectingaflightlessmodeoflife.
Herewewould liketonamethis fossil swan the
"AnnakaShort-wingedSwan''.
WeknOw"egIz/0の彫s加0画0ノz"jHowardl988wasa
flightlessCygnini,reportedfromtheMiddleMioceneof
California. The holotype is an almost complete
RECONSTRUCTIONOFTHE
$ $ANNAKASHORT-WINGEDSWAN' '
Osteology
Thenearlycompleteskeletoninthefossilallowsa
reconstructionofthebody(Fig. 3).
ConsultingClie"がes (Mergini), anexemplumof
flightlessanseriform, inwhichfemurchangedlittle, in
、譲舞,
k ロ 一℃‐
1夢=琴豐讓蚤誤ジル.』
VolantCygnus
(C.(Oノo〃oノoり
FlightlessCygninispeciesfromtheMiocene,
Annaka,Gunma,Japan.
Figure3. Skeletalreconstructionof"A'makaShort-wingedSwan'' (right),comparedwiththeskeletonofvolant
swan,Q智"〃so/"(left).
PreliminarynoteontheMioceneflightlessswanfromtheHaraichiFormation,TomiokaGroupofAnnaka,Gunma, Japan 5
spiteoftheextremereductionofwingelements (Liv-
ezey, 1993),the&[AnnakaShort-wingedSwan"mayhave
evolvedfromaspecies thatwasthesizeofQ唇""S
m/郡沈6"""s. Thus,overallproportions,exceptforthe
pectoral andwingregions, resemblethoseofQg""sプ ~7D
COJ〃”Z“α邦況S・
However, thewings, especiallythedistal elements,
areremarkablvreducedinbothsizeandbulk. The
ulnais55%thelengthofthehumerus. Elbowjointof
the"AnnakaShort-wingedSwan"mightnotreachthe
coxawhenfurled; onthecontrarvvolantswanssettleit
inthebackofcoxa.
Theincompletesternumistheonlvremainsofthe△
pectoralgirdle,butitstillshowsflightlessadaptationin
thesmallsizeandlowcarina. Nocoracoidsorfurcula
arepresent.
Theshortpelvisrelativetothebodysize isalsoa
remarkablecharacterof the "AnnakaShort-winged
Swan''. However, the total lengthof thevertebral
columnincludingpelvismightnotbechangedfromits
ancestorbecausetheenigmaticpelviscanbeunderstood
asaneotenicconditionasdiscussedabove.
Thetrunkmighthavehadaslightlyhigherangled
posturethanitsancestorwhenstanding, likethegeneral
trendofflightlessbirds.
Thehindlegchangedlittlefromthevolantancestor.
Butphalangesarenotwellpreserved
Thefaceisimpossibletoknowbecausethecranium
showsonlythebasalviewandtherostrumismissing.
Themandibleisbentandhasnarrowsvmphvsialarea.
Thislooksstrangebecauseanseriformesusuallyhave
straitbillandanextensivesvmphvsisinthemandible.
Furtherpreparationisneededtofindthestateofthebill
of "AnnakaShort-wingedSwan",whether thefossil
indicatesaspecialadaptationforitsdietaryhabitator
merelydiageneticdeformation.
ACKNOWLEDGMENTS
TheauthorsthankmostsincerelvPatriciaVickers-
RichofMonashScienceCentre,MonashUniversityand
HollisW.ButtsofMuseumlmport,Tokyowhohelped
kindlytocompletemanuscript. Weare indebtedto
MakotoManabeofNationalScienceMuseum,Tokyo
forhishelpinexaminingtheosteologicalspecimensof
recentbirdshousedinNSM.
REFERENCES
Feduccia,A.andTordoff,H.B(1979) : FeathersofA7c"""-
彫γyr: asymmetricvanesindicateaerodynamicfunction.
Sc""", 203: 1021 1022.
Howard,H(1988) :NewrecordsofMiddleMioceneanserifor-
mbirdsfromKernCounty,California. 1n,K.E.Camp-
bellJr.ed.,Rzpe窓加A"jα〃Rz/""加/Ogy,230236,Natural
HistoryMuseumofLosAngelesCounty.
Kaneko,M.andNomura,M. (1998) :Fossil foraminiferaand
PaleoenvironmentofHaraichi formatiOnandlowestpart
of ltahana formation inTomiokaandAnnakaarea,
Gunmaprefecture, Japan. B""節加Q/G""脚α〃"s鋤加
Q/rMz畑mJHISZD剛, 2: 57-66.
Livezey, B.C. (1993) :Morphologyofflightlessness inCノzg"‐
の彫s, fossil seaducks (Anatidae:Mergini) of coastal
California. ノリ"7w(z/ qf睦吋妨緬彫Hzノ"""/cgy, 13 (2) :
185-199.
Nomura,M, andOhira,H(1998) :Fission trackagesof
MiocenetuffsinTomiokaGroup,Tomiokaarea,Gunma
Prefecture. B"""加QfG""”α〃"s""Q/ JV鹿加"/
賊Sm籾, 2: 35-42.
OIson,SL. (1973) :EvolutionoftherailsoftheSouthAtlantic
islands (Aves:Rallidae). S柳"向so"""Co""""0"s /0
Zt)0/qgy, 152: 1-53.
OIson,S、L・ andHasegawa,Y. (1979) : Fossilcounterpartsof
giantpenguins fromtheNorthPacific・ Sc/"z", 206:
633635.
OIson,S.L・andHasegawa,Y(1996) :Anewgenusandtwo
newspeciesofgiganticPlotopteridaefromJapan(Aves:
Pelecaniformes). ノリ〃γ"α/ q/W減酌""Rz"0れゎjOgy, 16
(4) : 742-751.
MATsuoKAHiroshige,NAKAJIMAHajime,TAKAKuwAYujiandHAsEGAwAYoshikazu6
和文要旨
群馬県安中市の富岡層群原市層から発見された
中新世無飛翔性ハクチョウについて
松岡廣繁】 ・中島 -2 .高栞祐司3 .長谷川善和3
!京都大学大学院理学研究科・地質学鉱物学教室:〒606-8502京都市左京区北白川追分町2〒379-0116群馬県安中市安中1 9-15
3群馬県立自然史博物館:〒370-2345群馬県富岡市上黒岩1674 1
要』日
群馬県安中市の碓井川河床に分布する中部中新統上部の富岡層群原市層から発見された,全
身の骨格要素がほぼ完全に保存された烏類化石は,ハクチョウ族に含められるが著しい無飛翔
性の特徴を示す種であることが判明したので, ここに初期的に報告する.
化石は,尺骨の全長が上腕骨の55%にすぎないなど,翼の要素は遠位ほど退縮している.一
方下肢の要素は飛翔性のハクチョウ類と大きな違いはない.翼の退化した本化石をここでは,
"AnnakaShort-wingedSwan"アンナカコバネハクチョウと呼称することとした詳細な分類
学的検討はさらに剖出作業を進めてあらためて行うこととし,本報告では,骨格を中心に,飛
翔性ハクチョウ類とアンナカコバネハクチョウとの相違を示す.
キーワード:群馬県,安中市,富岡層群,中新世,無飛翔性鳥類,ハクチョウ族
PreliminarynoteontheMioceneflightlessswanfromtheHaraichiFormation,TomiokaGroupofAnnaka,Gunma,Japan 7
Platel.Theslabincludingskeletalelementsofonefossilbird,collectedfromtheMiddleMiocene,ofAnnaka,Gunma,Japan
MATsuoKAHiroshige,NAKAJIMAHajime,TAKAKuwAYujiandHAsEGAwAYoshikazu8
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Plate2. Closeupfiguresofdistributionofbonesontheslab. 1.basalviewofcranium(leftbottom)andmandible(right) ;2.anconalviewofproximalpal-toflefthumerus; 3.wingelememts; 4. legelements; 5,posteriorviewoflefttarsometatar-
sus; 6, right lateralviewof incompletesternum; 7. dorsalviewof leftscapula; 8.dorsalviewofpelvisandtwucaudalvertebrae. Allscalebarsare5cmlong.