BulletinofGunmaMuseumofNammlHistory(8):1-20,2004 1

Ori9inalArticle一一一一一一

SystematicsandEvolutionofDromaeosauridae (Dinosauria,Theropoda)

SENTERPhil',BARsBoLDRinche㎡,BRITTBrooksB.3,andBuRNHAMDavidA.』

'DMs/o"QfMtJrlie'"α""""Sc/e"",Lα"'αrSY"ECo//ege-O""ge,4/0舟℃"57．，

0F"Tge, 7Ex"77630US4

』J"s""eo/G"/･部A化"go"α"14"""z)'Q/此だ""s,E"紬TZzかα"jGI狂わ"I//, I/ﾉα〃

B"0r2/035ﾉ，ル/b"9り"α

｣D""""e"/q/.G"/0gllB噸/r"")f)""gUWA'e""P"",U/αル84602,LEA

.MJ""α/〃碗onjMWsE""1α"dBiDJ〃E蹄jり'R"Eqﾉ℃ルC"rgJ;""JLJJ'〃αwRBﾊ'乱，

U"八'ersfりノザKα"s",L"we"",K"7J･""043,UM

Abstract:Wepresentaphylogeneticanalysisofcoelurosauriantaxatoelucidaterelationshipswithin

Dromaeosauridae.TheresultsoftheanalysisindicatethatDromacosauridae (definedasall taxaphylo-

geneticallybracketedby蛇わarqp/"andDr･""""r"TJs) includes amonotypicVelocriapto'inae

(consistingonlyof姥んc"･"ml･"I｡"9･"e"J･jJ) andaspecioseDromaeosaurinae (inwhich"""z"郡，

鼬z"ひJ鞭"方0/Esr",DeiJToM)'C〃ｲs,4cﾉ""06"oJ3andUJq"mpIoﾉ､aresuccessivelymorecloselyrelatedto

DFり""E"""ws).Microraptona, thesistertaxontoDromaeosau'idae, includesMrcmFqp"J･口"a｡”"“

(=11Micmr"".gz"'!=nOplpml'0/q"""J"'),NGMC91,S/"｡"J"ﾙ"""･",andBa版6"np"r.Wealso

findthatS""1,e""0J･ isabasaldeinonychosaur,Aves is sister taxontoS/"ove""DF+(Microrapto-

ria+Dromaeosauridae),andtroodontidsaremorecloselyrelatedtoomithomimidsthantodeinonycho-

saurs. Thedistributionofcharacterstatesrelatedtoarborealitvandaerial locomotionsuggeststhat~ヴ ーー

thesebehavioral traitsaroseindependentlyinAvcsandMicroraptoria.

芦ロー』

KeyWords;Dinosaulia,Theropoda,Dromaecsauridae,Deinonychosauria,Phylogeny

|ntroduction ducedwithinDromaeosauridae (WeishampelandJianu,1996).

Inotheranalyses,S/""""ﾙ"""wsand〃『cmr"rcrareplaced

outsideanunresoIvedcladethat includes'4c""o6"",""""-

〃ｨ3,DEj"0"'ﾉcﾙ".F,DJ℃""""""Js,Sql"℃r""師/"r",U"""-

9",(/m"qpror,andlGMIOO/1015 (Xu,Wang,andWu,1999;

Xueta1.,2000;Norelletal.,2001;Clarketal~,2002;Xueta1.,

2002;Hwangeta1.,2002).

Here,anewphylogeneticanalysisofDromaeosauridaeispre-

sented, Thegoalsoftheanalysisare toresolvephylogenywithinDromaeosaulidaeandtonoteanvevolutionalvtrendsre-

ご ~戸~

latedtosizeortocharactersrelevanttoarborealitVandaerial lo-

comotion.

ThetheropoddinosaurfamilyDromacosauridaeincludessev-

era1 ''sickle-clawed'! generainwhichthesecondpedaldigit is

hyPerextensibleanditsullgual isenlarged. Theseanimalsare

ofparticular interest due totheircloserelationshiptobirds

(Gauthier,1986;Holtz,1994,2000,2001ラForsteretal.,1998;

MakovickyandSues,1998;Sereno,1999;Xu,Wang,andWu,

1999;Xueta1.,2001;Norell eta1．, 2001;Xu,Cheng, eta1,,

2002;Xu,Norell,eta1.,2002) andtheirpossessionoffeathers

(Xuetal.,2001;Czerkasetal.,2002;Xuetal.,2003).ColbertandRussell (1969) erectedDromaeosauridaefbrre-

ceptionofthesickle-clawedCretaceoustheropodsDJ℃"、“”"‐

繩sヲ姥わc"qp/Dﾉ-, andDe/"｡郷℃/"ｲJY, Sincethen, severalmore

genera (Sues,1978;Osmolska,1982;Barsbold,1983;Kirkiand

etal.,1993;NovasandPuerta,1997;LeLouefrandBouffetaut,

1998;Perleeta1., 1999;Xu,Wang,andWu， 1999;Allainand

Thquet,2000;Bumhametal,,2000;Xueta1.,2000) andun-

namedspecimens (Camenter,1982;Buffetautetal.,1986;An-

mnesandSigogneau,1992;RauhutandWemenl995;Csiki

andGrigorescu, 1998;Cifellieta1.,1999;Eaton,Cifelli,etal.,

1999;Eaton,Diem,eta1., 1999;Kirklandeta1., 1999;Pamsh,

1999) havebeenassignedtothefamily. Relationshipswithin

Dromaeosauridae, however,haveremainedlargelyunresoIved,

despiteseveralphylogeneticanalysesthatincludemultipledro-

maeosauridspecies･ Inonesuchanalysis,noresolutionispro-

Abbreviations

AMNH,AmericanMuseumofNamralHistory,NewYork

City,NewYbrk. BMNH,MuseumofNamralHistory,London,

UnitedKingdom. IGM,MongolianlnstimteofGeology,Ulaan

Bator,Mongolia. IVPRInstimteofVertebratePaleontology

andPaleoanthropologyjBeUing,China.LH,UniversidadAut6-

nomadeMadrid,Madrid,Spain.MCEMuseoCannenFunes,

PlazaHuincul,Argentina. NGMC,NationalGeologicalMu-

seumofChina,BeUing,China_ UA,Universited'Analanana-

Iivo,Antananalivo,Madagascar.YPM,YalePeabodyMuseum,

NewHavenConnecticut,USA.

Reccivcdl4Janualy2004;Acccptedl2Febnlaly2004

、／］ SENTERPhil,BARsBoLDRinchen,BRITTBrooksB.，andBuRNHAMDavidA

FirstphylogeneticAnalySiS:Methods

pmxP/り′/oge"e"c,fl"4's/s

AdatamatrixoflO1osteologicalcharactersfbr32coeluro-

sauriantaxawasconstructed,with』ﾉﾉ“α"jws/i""11asout-

group (Appendiceslandll) ． ThecharacterlistwasmodifiedffomthatusedbvSenter (2003).Thelatterlistwasusedasa

ご

basebecauseitwasconstructedlargelywithoutreferencetopre-

viouslypublishedphylogeneticstudies,soastoavoidbiaSofre-

sultstowardthoseofprevious smdies. However， it included

manycharactersiITelevanttocoelurosaurphylogenyandlacked

manycharacters relevant tophylogenywithinDromaeosauri-

dae. Characters irrelevant tocoelurosaurphylogenywerede-

leted, and characters relevant to phylogeny within

Dromaeosauridaewereadded. Allcharacterswerelefiunor-

dered. OnlyonesupraspecifIctaxon (Pro"ﾉ℃ﾙαgOpだﾉ)'Lhr)wasusedintheanalysis.

PhylogeneticanalysiswaspelfbrmedwithPAUP4.0bsoff-

ware (SwofTbrd,2002),usingbothabranch-and-boundsearch

andahlllheuristicsearch. ABremeranalysisandabootstrap

analysis (100replicatesinbranch-and-boundmode,withlO

random-additionsequencesperreplicate)wereperfbnnedwiththesamesoftware.

I

．I

｢I

篭 く亘二Fi9ure l Similarites between premaxillae (top) and dentaries

(bottom) oftheholotypesofP""℃/7口E“だF"(len,drawnfifom

photographsofNGMC2125) and ::I"cic八℃sα"Fws" (right,modifled

fmmXu,Chengetal.,2002). i=incisifbmtoothcrown,d=

denta『yｩpmx＝premaxilla.ri＝『ootofincisifbrmtooth．

Theungual exhibitsanextremelyprominentnexormbercle,

andtheungualisdorsaledgearcheshighovertheproximalar-

ticularsurface. Amongdromaeosaurids, thesetwotraitsare

fbund inmanualungualsbutnot inthesecol1dpedal ungual

(Ostrom, 1969, figs､ 63,71;NorellandMakovicky,1997,fig.

6;NorellandMakovicky,1999,fr9.8;Hwangeta1.,2002, fig,

30) . Theelementisconsideredtobelongtomanualdigitl,asit

lackstheproximodorsal"lip'! fbundindromaeosauridmanual

ungualsllandIII.

Here, thesecondpedalungualofJc〃oわr"耐p灯加aisconsid-

PDre""α/4'Co""ひveバノα/ScDJ･"唖

Theholotypeof"c""06"("毎gjg"""sincludesanelement

thatwasdescribedasasecondPedal ungual (Perleet al.,

1999) .However, it ishereconsideredafIrstmanualungual.

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68 １’85,2

81,284.2 １

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型レドー」

98,且57,1＝＝ 二＝＝＝＝＝凸

88，2 Dromaeosaurinae:63,1MiCrOraptOna56_l7-

Troodon-

tidae:73,2

mfctometa-

tarsalia:83,2

Dromaeosauridae:87,･

1 75,･IDein･nychosaurial

Para…74､｡IManiraptora,64,･

Maniraptorifbrmes:69,･’’ 64,｡

｣91,･’

Fi9ure2Strictconsensustreeoffirstphylogeneticanalysis.Numbersateachnodeindicatebootstrap (plaintext) andBremer (underlined)values.

1SystematicsandEvoILltiol]oI､Dl･omaeosauridae (Dinosauria,Theropoda)

eredunenlarged, contraZhoLlandZhang (2002) . 1nJ.βﾉ･ﾉ"7α，

thesecondpedalungualisonlyl.3xthelengthofpedalphalanxⅡ-2． a l-atiosimilal･ tothat inotherMcsozoicbirdswithunen-

largedsecondpedalunguals, e.9.4ﾉ℃ﾙαeOp花ﾉ]猟／"ﾙogﾉ．“ﾙﾉcα

(BMNH37001) , ノルeﾉひ"7“oﾉﾂ7耐ﾉU"7匂．α〃 (LH22) ,Co'7co"7/.''

/qc,"IF"肥､ (LH2814) , andS肋o/77/s""/e"s/s (Sere,,oet a1.,

2002, fi9.8.6) ． 111Rq〃o"αWs""()"7/, troodontidS,anddeinony-

chosaursothcr than"ctYsα"/WS， thesecondpedal ungual isat

least twicethe lenglhofpcdalphalanxll-2 (notcoLIntingthe

postcl.oventl･al "lip"ofphalanxll-2) . This istl･ueevenfbl,dei-

nonychosaurs inwhichpedalphalanxll-2iselongated, sLIchas

M/c/･oﾉ傘qp/oﾉ" (Hwangeta1.,2002,f19. 30) andNGMC91 (pCl･-sonalobsel･vationbyP.S.)Here,C"ﾙ“oﾉwzsljα"f"c"isconsid-

eredajulliol･syllo11ymofS"7or"/s""re"s/s,aspel･Se!℃11oetal.

(2002) ~

Here､ﾉ"cﾊﾊﾉo"'"･"s isconsideredajuniorsynonymofPﾉ｡o‐

/“℃ﾙaeOpreﾉ:Mx. Data fbrPﾉ-Orαﾉ℃ﾙqeOpre/yxwel℃collected

廿omtheholotypeofR/･o伽sm(NGMC2125) andthepub-

lisheddescriptionofRg""〃e"(Xu,Chen9, eta1.,2002) . P

g""ﾙﾉeノゾwasol･iginallydescribedasthetypespeciesofanew

genLIs､ /"cパパo:"""/､T (Xu､Chen9,etal.､2002)However, the

twospccieSshOuldbetl-eatcdascongeners、astheil･ovcl･lapping

elementSal-c identicalexcept ibrtoothcount. Thesctwospc-

ciesL111iquelyshareaverybizarredentition､whichhasanLImbel･

ofLInusLlal attl･ibutes: (1) asinglepairofenlarged,anteroPostc-

l･iorlycomprcssedteethat thcfi･ontoftheprelnaxilla, (2) di-

mlnutlve、 pcgliketeeth immediatelypostcl･ior tothccnlal･ged

pail･ofteeth, (3) diminutive, lanceolateposteriol･ teeth,and (4)absenceofteethat thetipofthedentarv (NGMC2125;Xu，

ユ ~‐

Chen9,eta1 .,2002) (Fig. 1 ) . AIso,deSpitethecl･LIshingofthc

skullinNGMC2125,whichissoextensivethatfbl-mostuppel･

teethonecannol tellwhetherthevarelefiol- l-ight teeth、 it isジ ー

clear thatNGMC2125 sharcswith "I"cﾊﾊﾉo”"ﾉw3" (1) a

sho1-t,highskull， (2) tallpremaxilla,and(3) denta'ywithante-

l-iormarginstl-onglybeveledsothat thedorsalmargintermi-

nates ina50｡ -60｡ POint (NGMC2125;XLI,Cheng, eta1.,2002) (Fig. 1 ) . Weconsidel･ theseSimilaritics, csDcciallvthe

ー ﾀ 八 学

dental charactcrs， sLIfficient todemonstrategenel-ic identity. A

signiliCantdill℃renceintoothcount (Jieta1. 1998:XLl.Chcn2、ｰ ー〃

etal,2002) jLIstifiesmaintenanceofseparationofthetwotaxa

attheSpecies Icvc1 ． Howcver, thiSdifferencedoesnotcompro-

misetheutilitvofthegenusasanoperational taxonomicunit lllヴ ー ュ

thepl･esentanalysis.

PLlblisheddescl･iptionsstatcthatSerratio11sal-eabsentonthe

teethof "/"c/sﾊﾉ“α"ﾉWS" (Xu､Cheng,eta1.,2002) butprescnt

onthoseofNGMC2125 (Jieta1., 1998) . Howevel･,examina-

tlonofNGMC2125byP.S.、usingazoomlcnsasamakeshifi

microscope,revealedthattheapparentserrationsontheonepl-e-

servedenlargedpremaxillarytoothcrownareplaceswhcrethe

edgesofthetcethal･cchipped. Noserrationscouldbcsecnby

P.S･ onthepostel･ioI･ tceth、 11orareanyevident inthepublished

photosofthesetccth (Jieta1. 1998, f19． 1) .

treeS ispresentedinFig.2． Fol- synapomol-phies, seeAppendix

lllandFig.3. Bootstrapvalues showmoderatesupport fbr

mostnodes,withveryhigh(>95) sLIpportfbrcladcswithinMi-

Cl･oraptorini (Fig. 2).Brcme'､valuesarehigh (>5) 1bl･Dro-

maeosauridae, DromaeosaLll･idae +Microraptorini,

S//7oﾉ･""ﾙo"z"ws+ (NGMC91十ﾉWcﾉ℃ﾉ℃〃o｝･) ,Pa,･avcs,Mani-

raptora,Maniraptorifbrmes,S/""""･O/)/e/WW+Maniraptorifbl--

mes, and O/77"ﾙo/a""+ (S/"os"""/eﾊ概十

Maniraptorilbrmes) , indicatinghighsupporl lt)1､ thecomposi-tionsofthesecladesasibundbvthIsanalvsis.

Subsequentphylo9eneticAnalysesMethodsandResults

Usingthesamcmethodas inthefil･stphylogencticanalysis,a

fEwsubsequentphylogeneticanalyseswererun・ Inoneanaly-

sis,S//701,e""oﾉ‘ cﾙα"g//was excluded, to seewhether this

wouldinnucncetreetopology. ltdidnot.

Ineachof4othel･analyses,anewingl･oupwasadded, tosee

whereit fbllwithinthetree. Thesenewingl･oLlps (U"e"/qg/q

CO"7αﾙ"e"s/.F,〃"/.s'""p"ﾒ〕eﾉﾂe/,2y/ひﾉ'"/0ﾉ．0{w"ﾉ〕"/'F,and肋ﾉｿｰ

ﾉ"(")/Oﾉ･ ﾉ""/7//7oﾉ"""7) hadbecnexcludcdll･omthefirstanalvsis

duetoalargeamountofmissingdataineach. ltwassusDcctcdｰ ー 人

thatsuchlargcamountsofmissingdatawouldIowcl･ resolution

inthetreeThissuspicionwasjustilicd, aseachnewingroLIp

ibrmedpartofapolytomy (Fig.4) . Thcseanalysesconlirmed

dcinonychosaL11･ianstatLIsfbr[ﾉ"e/7/q9/",〃"As'qﾉｳﾌGs,andPwo-

ﾉ"qp/Oﾉ･,butcOuldneitherconfirmllo1･ ltllsih/deillo11ychosaurian

statusfbl- ItW『叩/Oﾉ" (Fig.4) .

Extra-StepAnalyses:MethodsandResults

Scveral extra-stepanalyscswc1℃pc1･1brmedbyopenillgthe

strlct consellsus tree廿omthe first phylogeneticanalysis, in

MacClade3.08a (MaddisonandMaddison,1999).Taxaofin-

terestwerethenmovedtovarioLIspositions,andthechangein

thenumbel-ofstepsl℃cordcdfbl･eachnewtrec (Fig.5)

Theseanalysesshowthat>12cxtl･astcPSal･Cl℃quiredtopail･

S"7()1ﾉe"q/o/"withT1･oodontidaeo1･ topail･Troodontidacwith

11011-avianparavians, andthat>9extl-astcpsarerequiredto

placeDromaeosauridaewithinAvcs・ ThehypothescsthatS/"o-

vE"α/oﾉ" iSabaSal troodontid (Xu,Nol℃Ⅱ、 etal.、2002) 、Troo-

dontidae iscIoselyrclatedtoDromaeosaLl'･idae (Ostrom， 1969;

Sues, 1978;Currie、 1987;Gauthiel-、 1986) , andD1･omaeosauri-

daefallswithinAves (Czerkaseta1.． 2002: Paul. 2002) al･e

therefbl℃notsupportedbvthedataselusedhcl･c.人~ユ ヅ

Discussion

P/IV/oge"e"cPos///o"Q/､S/"OVe""rOﾉ・

ThemostcontentioLIs I･csultofoLII･ phylogenetlcanalyscs is

probablythepositionofS/"oIJe""oﾉ･. Hcrc,wefindthatS/"ove-

"q/Oﾉ・ iscIoselyrelatcdtoDI･omacosauridae, on thebasisof

threeunambiguous synapomorphies: vcntrally-ftlcingantel･ior

articulal･ sLIriacesofcervical centl･a， apostc,･ovc,,tral (､lip'' on

pedalphalanxII-2, andanenlargedsccondpedalungual (Ap-

pendixllnAsthclattertwochal･actcl･ statesa1℃sharedwith

TroodOntidae,support ibrthisphylOgenelicpositionfbl-S"701>e-

〃α/o/､ islow. l-Iowever,Smo1ﾉe""o/､alsocxhibitsI-etl-opubyand

Fir"PhylogeneticAnalysis: Results

Phylogcneticl-csLIIIswerethesamefbrbranch-and-bOundand

heuristicscal･chcs. Theanalysisrecovercd27most-parsimoni-

oLIstreesof227steps.Fol･ thesetl-ees,CI=0.4912,RI=0.7972,RC=0.3915、 andHI=0.5088． Thestrictconscnsusofthesc

SENTERPhil，BARsBoLDRinchen､BRITTB1･ooksB. ､andBuRNHAMDavidA4

talyteeth, (7) slendel･sccondandrobust ibul･thmctatal･sal, (8)

maxillaanddentarywithmoderatenumbel･ofsmall tceth (25

30 indcntal･y) , (9) anteriordol･sal transversepl･occsscs short,

wide,alldonlyslightlyinclined， ( 10)nat inte'･narialbar.

ThePairingofS/"Clﾉe"WOﾉ〃withTroodontidacbv I)1-evioLIs

analvses isduepartlvtouseofadifferentsetofcharacters. Of鼻~ 一

thepLltativcS/"ove""oﾉ今-Troodontidae synapomol-phies listed

above,all but thescventhandeighthl-epl℃sentcharactersthat

werenot LIsedinthepresentanalVses.

Howevel-, the fifihthroughtcnthpLltativc synapomorphies

above l-epl･esentmiscodings andotherproblems･ The fifth

throughscventhputativesynapomorphiesrepresentmiscodillgs

adistallyginglymoidthirdmetatal-sal,whichareparaviansyna-

pomorphlesthatare lackedbytroodontids (AppendixIII) .

P1-cvioLIsanalyses thathaveplacedS//701ﾉe/7"Oﾉ" asabasal

troodontid (Clarketa1..2002;Hwangeta1.,2002;XL1,Chen9,

eta1 .,2002;Xu,Norclleta1., 2002) all LIsedval･iatio11sonthe

samcdatamatl･ix,eachsubsequentanalysiSaddingcharactersortaxa. Intheseanalvses, thefbllowing lOcharactel-statesunited

ず グ ー

S"701ﾉeﾉ7"/Oﾉ･withTl･oodontidae: (1) absenceofbasisphenoidre-

ccss､2) oval fbl･amenmagnum, tallel･ thanwide、3)nut,･ient

fbraminaonlatcralsurfaceofdental･yinadeepgroove, (4)hol-lowquadratewithDosteriol･ depression、 (5) thin-walledbonv

~エ エ ュ ー

rimal･oundantorbital fbssa、6) anteriordentalytecthsmallcr,

morenumerous,andmorecIoselyappl-cssedthanposteriol･den-

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｛。SystcmaticsandEvolutiollofDl･omaeosaLll･idae (Dinosauria､Theropoda

ofchal-acterstatesofS"70'ﾉe"〔"o/･cﾙα"g"・ Pcl-sonal observa-

tions ofIVPPV12615 andIVPPV12583 (S//7ovE"c//oﾉ．

cﾙα"g") byP. S. rcvcal theibllowing． S. cﾙα"g//has llomol℃

ofabonyrimal･ound itsantorbital fbssa thandodromaco-

saurids~e.9. Jt/ocﾉﾉ."roﾉ･〃7oﾉ噌o//e"s/s (AMNH6515).Ante-

riorandposteriOl･dentaryteethofS. cﾙα"g//areroughlyequal

insizc.numbel-､anddensitv･ Thesecondandfbul-thmctatal-salsゴ

ofS.cﾙα"g//donotdiifel･appreciablyindiameter.

ThccighthputativcsynapoInolphyaboverepresentsmiscod-

ingsoftl-oodontidtaxa. Presel-vational al-tifaCtSpl-cventadefi-

nite toothcount in the troodontidsBW℃"o‘”"ﾉ･"‘，ノ‘"を/ and

S/"oﾉ･""ﾙo/""'oz"79/,butpersonalobservationsbyP.S･ ofthefbnner (IGMlOO/983) alldofacastofthelattel･ (unnumbered

IVPPcast) reveal that thedentary toothcountwas certainly

higherthall40,giventhesizesofpreservedteethandthelengthofthedentaries. Tootll countsexceed30 in the tl-oodontids

Saz"℃"7"ﾙo/北s/""/oﾉ" (Bal･sbold, 1974) and刀ひoch)/7/b""“"s

(figured inCurl･ie、 1987) ． TToodontidsshal-e therefbreshareー

thecharactel･ state "teethrelativelvsmall andnumeroLIs (more

thall30indentaly)" (Clarketa1.,2002,p50)withthebasal

ornithomimosaul･Pe/ecα"/"""I"〃ル(W",/ (personal observatio'1

byP.S.ofLH7777:PルQ)'"/) . Thispl･ovidesadditional sup-

port fbl- thecloserelationshipbetweenTroodontldaealldOr-

nithomimosauriafbundbVtheprescntanalVscs.

TheninthputativcsynapomorphyabovcmaybcductotheLIseof $G?"fbrtherelevantcharactcr inccrtaintaxa・ Forexam-

ple,Clarketal . (20()2) codethischaractel･state :c?" 1b1･4/℃ﾙae－

”だﾉfyx/"力ogﾉ痩叩ﾙ/cα、Raﾙo"αWso3"o"7ﾉ、 andovirapto1-osaurs

otherthanM7cJ"ove"α/o7･ce/e'" (fbrwhichthecharactel･ iscoded

G;l'',as inS/"ovE""oﾉ"andtroodontids) . Personalobservatiolls

byP. S・ ofspecimensofj. ／"ﾙogJ謬卯ﾙﾉc" (BMNH37001),R.

OS"℃"7/ (UA8656) and theoviraptorosaurkﾙαα〃〃7歌e""αノ(IGMlOO/1002､ IGMlOO/1127) revealthat thesetaxacxhibit

state1 ． Thischal･actel･state､ therefbre､coLlldbeinterpretedasa

maniraptoransynapomorphythat is l･evel-sedinDromaeosauri-

dae+MicrOraptoria.

ThetcnthputativcsynapomolphybetweenS/"oww"oﾉ､ andTl･oodontidae is･ shal･edbvomithomimosaurs (Clarket al.、

ソ

2002) . ltcouldjllstasparsimoniouslybe interpretedasan

omithomimosaul･-tl-oodontid synapomorphywithconvel-gence

inS加ove""oJ'asaS加ove""o7"-troodontidsvnapomomhvwith雪r ユ ~ユご

convergenceinOmithomimosauria.

s叩ﾉ割age"e"c/Vo"7e"c/"z"で

Supragenericnomenclatureofthe sickle-clawedtheropods

hasalongandconvolutedhistoly.MatthewandBrown (1922)

el-ectedthesubfamilyDromaeosaurinactoemphasize thedis-

tinctncssofDJ"o班αeos""wswithinthellow-defilnctitlmilyDei-

nodontidae. Later, Colbert andRussell ( 1969) rasied

Dl･omaeosaurinae tofamilystatus (asDromaeosauridae) and

addcdDe/"oﾉwcﾙ"､F and It/oc""p/Dノ" tothe ftlmily. Ostl-om

(1969) addedSqz//oﾉ”"ﾙo/cたs', STe"oMyc/7os[""ws, andC/7/ﾉ℃‘y‐

だ"o/"toDI･omacosauridae; however,Cﾙﾉﾉ℃,We"o/"waSsubse-

quCntly reたrl･ed toOviraptorosauria (Currie andRLIssell,

Fi9ure3 ScIcctcdsyl1apomorphiesoI､dcinonychosaurianclades. A. Pcdal digit llofM/cMJ蕊叩/oﾉｰzﾙaojα""|3 (mOdifiCdfi･OmHwangCta1.,

2002) 、 showingsynapomorphicsofDcillo11ychosauria:postel･oventl･al ! 'lip'!onpcdal l]llalanxll-2; cnlal･gementofpedalungual ll. B.Teethof

S""℃I．""ﾙo/"/"/α"9,‘/oﾉ7/ (modi llcdn･omCun･iect il1., 1990) ,dentaryofit/o

SENTERPhil，BARsBoLDRinchcll,BR['[･･lBrooksB.、alldBuR1､《HAMDavidA６

high, relativelylargeSkull"、heassignedDﾉひ"7qeo."Z"･".5',De/-

〃o"1ﾉcﾙ"‘『， and4dt/s"""s_ 1bthe latter, consistingof $Gsmall

dromaeosauridswitha low, relativelysmall skull'' ,heassignedIf/oc"wﾌ/oJ雫．

』

Tworecentpapers (Sereno,1998; Padiancta1.,1999) have

attemptcdtostabilizesupragenericnomellclaturefbl･ theropod

taxa, usingtheprillciplesofphylOgenetictaxollomy (DeQuci-rozandGauthier， 1990) . 111 thenrst, Serello ( 1998) defines

Deinonychosauriaas al l taxaphylogenCticallybraCketedby刀CO”〃and陸/oc"""Zo/~. l-ledefYnesTroodontidacasalldei-

エ

11onychosaursmoreclosclyrclatedto7)"oo""thantolt/oc〃叩一

/Oﾉ割, andDI-omaeosauridaeasalldeinonychosaursmorcclosely

l･elatedtoIt/oc""/o/･ thanto7)"oocわ". HedelinesDromaeo-

saLII-inaeasalldromacosauridsmoI-ecloselyl-clatcdtoDﾉ℃"7α“－

s“"･"s than to Ft/oc"･叩/oF, andVelociraptol･inae as all

dromaeosauridsmorecloselyrelatedto"/oc'"")/oﾉ･ thanto

DI℃"7""""ws. Inasubsequentpapcr,Padianetal. (1999)

de61]eDeinonychosaul-iaasall lnanil-aptol･ansmol-ecloselyre-latedtoDe//70ﾉ”cﾙ"sthantoNeomithes.ThevdefincDI-omaeo-

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Fi9ure4 St'･ictco''sensL's t'℃csofphylogeneticanalyses incILlding

[ﾉ"e/7/Qg/αし'o"7"/1"e/7,$/.5･,〃"伽"加露peﾉ･/e/,角‘ﾉr〕ﾉ･"/oﾉ．o伽"/)"ﾊ､ ,and

ﾙ加･"･〔ｳ刀oﾉ・ 〃ﾉ“.ﾙﾙ7oﾉ･""7. A. Strict collsensus treeofphylogenctic

analysis incILlding(/"eノ7/Q9/Qcoﾉ77α加e"sﾉs. B. Stl･ictconscnsLIs tl･cc

ofphylogcnctic analysis incllldillg""/""p"peﾉ･/e/・ C. Sll･ict

conscnsus trcc oI､ phylogenctic analysis includingRWY)ﾉwp/("‘

oか""刀"s'. D. Stl･ict conscnsLIstreeofphylogeneticanalysis illcILld-

illgif〃〃α/〃o/･〃7(Jc/7//70/･""7．

12 12

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91988),andS[""℃ﾉ･""ﾙo化陀『andSfe"o"VcIzos"z//w'Fwereplaced

with刀ひo伽〃inthcftlmilyTroodontidae (Russell, 1969; SLIcs，

1978).Barsbold (1974) erectedtheflmilvSauromithoididac

fbl-S""．o〃7"伽/ct,FandSfe"oﾉりﾉcIIos""･z/s, toemphasize thcil･

distinctncss fi-om刀ひ“んﾉ7& however、 Currie (1987) subse-

quentlyarguedthat"e/7o"ycﾙCs‘""筆"s isajuniorsynonymof

7)"ooC/()", and thatSauromithoididae is ajuniol･ synonymofT1-oodontidae.

Colbcrt andRLIssell ( 1969) erectedthein6･aordel･Deillo11v-

chosauriafbl-DI-omaeosaul･idac, tocmphasizelhe lattel-' sdis-tinctness h･ommembers of ill廿aordcl･s Coelurosaul･ia and

Camosauria・ Troodo11tidaewassubsequentlyaddcdtoDcinony-

chosallriabysomcauthol･s (e.9. SLles, 1978;Gauthic1･， 1986:

Curric, 1987) ,butnotbyothel･s (e.9.Barsbold､ 1983) .BarsbOld (1983)dividedD1･omaeosauridaeintotheSubitlmi-

l iesDromacosaul-inaeandVCIociraptorinae. Tothefbrmersub-

family， consistingof"modcratelylargedromaeosauridswitha

’

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24

Fi9ure5RcsLlltsofextra-stepanalyscs inwhichS/ﾉ701'eﾉ7moﾉ･.Troo-dontidac,andOrnithuraearel･cpositiollcd.Numbcl･ inmiddlcofeach

cladogl･amindicatesthenumbcl･ofextl･astepsncccssal-ytoproduce

thc cladogram's topology. Arch, 』ﾉ℃伽eO/J/2/yx. D, Dl･omaeo-

saul･idac. M, Microraptol･ia. 0m、 Omithomimosauria. 01･th,

Ornithul･aC.Ov,OviraptorosaLll･ia. Sv,S/"oIﾉ“α/oﾉ．.T,Troodontidae

(withOUlS/ﾉ7Cl，e"moJ･) .

７SystematicsandEvoILItionofDromaeosaul･idae (Dinosaul･ia,Thcl･opoda)

saLII-idae as all taxa phvlogeneticallv bracketcd bv入－ ｰ ‐ ヴ

Dﾉ℃"7α“”"ﾉ"z/sand It/oc""/o/"・ Theil- definitiollsofDro-

maeosaul･inae andVelociraptOrinaematch those ofSereno(1998) ．

UnderthephylOgenyrecoveredhere, Scl･cno, s ( 1998)Dei-

11011ychosaLIria is a juniol･ synonymofManiraptorilbrmes

(Holtz, 1996)Thenext-publishedphylogcncticdeiinitionof

Deinonychosauria, thatgivenbyPadianetal. ( 1999) , thel･efbre

haspriority. AIso、underthephylogcnyrecovel-edhere,Deino-

11vchosaul-iaasdefincdbvPadianetal． (1999) cxcludeS7)･oo-ご ｰ

c/b/7, which renders Sercno's (1998) definitions of

T1-oodontidaeandDI-omaeosauridaenonsensical. Thcnext-pub-

lisedphylogeneticdefinitionsofTroodontidae (publishedhel･e,

inAppendixlll) andDl･omaeosauridae (thedefinilionpub-

lishcdbyPadianctal.,1999) 、 therefbrehavepriority (note,

however, that Lllldel･ aphylogeny inwhichTroodontidaeand

Dromaeosauridaearesistel- taxa,Sereno,sphylogeneticdefini-

tionsarcnot llonscnsical,andtherefbrehavepriority) .

Thesistcl･cladetoDromaeosauridaesE"s"s〃/c/oIsspecIosc

enoughtowal-rant itsowntaxonomicname･Wetherelbl℃intro-

duccthenameMicrol-aptoria,definedasthccladcoftaxathat

aremorecIoselvrelatedtoA"7cﾉ響07W"/Oﾉ" thantoIt/oc/ﾉて"ﾌ/O/･Cl・ゴ エ 』

Dm"7"e"'""〃パ. Wchavcdeliberatelvavoidedthcsufflxes "-

idae'' and"-inae'' fbl･ thisname. Underaphylogeny inwhich

TroodOntidaeandDromaeosauridaeal℃siste,･ taxa,andSerends

(1998) nomcnclaturchaspriorty, "Micl･01･aptoridae"wouldoc-

curwithillDI-omaeosaul-idac、 engellderingconlilsionbyyield-

ing theappeal･anceofa familywithina family. Undel- thc

phylogenyl･ecoveredhere､ ;Microraptorinae"wouldoccurout-

sideanytaxonwithanameclldingin"-idae'',engenderingcoll-

1iIsioll byyielding theappearanceofa sLlbfamilywithout aflmilv.

Fi9ure6 SilhouettcsoI､coelul･osaul･sandatroglodyte, toscale, to

showsizetrendswithinCocILll-osaul･ia,withcmphasisonDcinolly-

chosauria. CIockwise fi･omlowcl･ lcli: Oﾉw"ﾙo/"/", S""""ひ"‐

たﾉｯ猟,S加oJw"ﾙo/火.『、P/℃/“℃ﾙ“oﾉﾌﾞ/"", 望ｲﾉ℃〃α“ﾉﾌﾞ/eﾉﾂⅨ,Ra加"αw‘§'，

ﾉe力o/oﾉ鋤/"s, S/"CVGﾉ7“0ﾉ．,&"77〃ﾉ.(")/oﾉ･, S/"0ﾉ．""ﾙ““"･"s, 11Cﾉ坪"o‐

”/α"s" (=/Wcm'･qp/oﾉ鍔; sectcxl) , レセ/oc//"q/J/o/-､故わ”z"曇"s,Sα"ﾉ.oﾉ･‐

""ﾙo/"/"、De"7o"1)c方"‘↑,小'/7///(ﾉ伽/ひﾉ｡.D(""“Csα"ﾉ""4F,U/αん･fip/Oﾉ・・

Scalebar=2m.

(Czerkaseta1., 2002) ,al･ealsofbund in"Mgz//'! (Xuetal,

2003) ． SpecimensofMzﾙαoﾉ〔"7"､9withunfilsedstemalplates

(Hwangeta1.,2002) al･csmallerthallthoseof!!Mgz//'1and"C

pα"〃! thatexhibit filsedstcmal plates, soontogenycanllotbe

1-uledoLltasanexplanationlbl- thedilTbrence.

Characters that diagllose ' 'M． g"/' ! includeabowed tibia,

curvedpubis,sho1-tcl･manualdigitl,andprominentbicepstube-

rosityontheradius (Xueta1.,2003)However,Mzﾙαo/α""s

and ! IC.pα"〃' exhibitbowcdtibiae (Hwangeta1., 2002, fi9.

28;Czel-kasetal.,2002､hgll).AIso,thelengthl･atioofmcta-

carpal l+1nanualphalanxl-l tohumerus isthesame (0.46) in

Mz/rqo/α""s, "Mgz//1 1,and! IC.βα"//! ! (Hwangeta1.,2002, ta-

blel;Czerkasetal.､2002,69. 17;Xuetal,2003;6g. 1)"C

pα"〃' andSﾉ"oﾉ･""/7“α"ﾉ･"3〃""e""exhibitbentpubes (Xu,

Wang, andWU、 1999;CzerkaSeta1., 2002, f19. 2) 、 andde-

scribedspecimensofMごﾙα

8 SIzNTl｣RPI1il.BARsBoLDRillchen､BR[TTBrooksB.、andBuRNl.IAMDavidA

D城ﾉ･肋""0〃0/､〃ﾙoﾉでα//〃α"cMeﾉｿαﾉLoco"10//o"//7Mn"〃α"/o/･α

~~ ~冬

SomeresearchershavehvDothesizedthatmembersofDI-o-ジユ

maeosaul､idaese/7sos/ﾉ･ﾉ'c/oaresecondarilyllightlessandterres-

trialbirds､ descendedfi･omHving, arborealancestors (Czerkasジ －〆

etal.、 2002; Paul， 2002) . Thephylogeneticresultsreportedhel-eindicatethatdromaeosaul-idsal-enotmembcrsofAves､but

thisalonedoesnotpl･ecludethepossibilitythat theirancestors

werevolantol､arbol･eal. Toaddressthishypothesis, itwouldbe

usefill toexamine the l)hvIoueneticdistributionofchal･actersム ダ －

1･clevant tonightandarborcality. Thefbllowingcharactersare

ofparticular interest: fbathel-vanesymmetry, themolphology

ofmallualphalanxII-1,humel-aldiameter, thcpositiollofmeta-

tal･sal l、andpcdalphalangealproportions.

Vanedfeathcl･switharachisarcknowninoviraptorosaurs (Ji

eta1 ., 1998) ,deinonychosaurs (Czerkaseta1. ,2002) ,andbirds,

butnotamongthe intcgumental-ystructurcsofnon-manirapto-

rallthcl･opods (Chencta1., 1998;XLI,Tan9,andWang, 1999) ;

vanedfeathersarethel･efbl℃likelyasyllapomolphyofManil-ap-

tora. FeatherswithaSymmctl-icalvanes,ahallmarkofvolant

birds、arekllowninbasalbil-dsandinjiuibtangmicroI-aptoriansー 上

(Czerkaseta1.,2002) . lt ispossiblethatasymmetricalvanes

areasynapomol-phyofAvesandtheJuiibtangmicoraptorians.

Ifso, then this charactel･ state--and. bv implication， aerialF ジ ユ

locomotion--wasprescnt in theparavianancestol-・ However,

cleal-ly-preservedfbathervanesareLInkllownintaxaPhylOge-

neticallybrackctcdbyAvesandJiufbtangmicl･oraptorians (S/-

"OVe""rOﾉ･, Dromaeosauridae, and basal microraptorines) .

Symmetricalvanes inanysLIchtaxawouldcastdoubtuponthe

hvDothesisofcommon inheritanccofvaneasvmmetrvinAves〃ユ ヴ ~－

andtheJiufbtangmicroraptorialls、whereasasymmetricalvanes

insuchtaxawoLlldsuppol･t thehypothesis. lt istherefbrebest

toreservejlldgementuntilmorelindsarelllade.

AposterioI- flangeonmanualphalanxll-1 , increasingitsdi-

ameter, ispresent inOI-nithuraeand inmicroraptoriansothel-thallβα"76"｡α"/Oﾉ急.At lcastonel･esea1℃herhashvDothesizedthat

璽 ゴュ

thisnangeisrclatedtolbathel･attachmcntalldtoflight,andthat

bil-dSandmicl-oraptorians inherited it竹omacommonancestor

(Paul,2002) . Howcvcr, theflallgeisabsent in4′℃ﾙαeOp舵ﾉﾂx，

DromaeosaLll･idac、andβαﾉ"ﾙかf")/0ﾉ感､whicharephylogenetically

bl･acketedbyO1-nithuracandthellange-exhibitingmicroraptori-

ans. lt ismol･eparsimonious toinfbrthat thenangeevolved

twice (a2-stepscena,･io) thantoinfb,､ that itappearedonceand

was then lost thl℃c timcs (a4-stepsccnario) . It is therefbremore likelvthat theflangeevoIvedindenendentlvinbirdsand

ｰ ~ ユ ヴ

mlcl.oraptorlanS.

Humeral diametel・ is incl-eased inOmithuracandJiufbtang

micl･oraptorians, appl-oachingol･ excccdingfemoral diameter.

This inconsistentwithincreasedrelianceuponthefbrelimbfbr

bodilysupport,e.g・duringacrial locomotionbyibrelimbwings

inthesetaxa. l Iowever、 thesetaxaphylogeneticallybracket5

taxa inwhichhumeral diametel･ isnot increased (』ﾉ℃hqeOp-

reﾉ忽", Dromaeosauridac, βα"76"."Ioﾉ･, S//7oﾉ”"ﾙos""""&9, and

NGMC91).ltismorepal･simonioustoinferthattheincrease

inhumeral diametcl･evoIved indcpcndentlyinOrnithuraeand

Jiufbtangmicroraptorians (a2-stepscenario) thantoinfel･ that

theincreaseinhumeraldiameterwaspresent intheparavianan-

cestorandthen lost independentlyinthc5bl･acketedtaxa (a6-

stepscc,'ario) .

Distaldisplaccmcntofmctatarsal l,present illAvesandMi-

cl.01.aptorla、 suggestsal･boreality, as itwouldhave facilitated

perchingbyallowingthefirstpedal ungual togripabranch.

Metatarsal l isnotdistallydisplaced inDI-omacosauridac、 and

thestateofthischaracterisunknowninS"7ol）e"“oﾉ｡.AhypOthe-

sis that distal displacement ofmetatarsal l occulTed in the

paravianancestol･andwas Iost inD1･omaeosaul-idacl-equil-estwo

evolutionarysteps, asdoesahypothesisthatmetatarsal lwas

displacedindependentlyinAvesalldMicroraptoria・ TheknOwn

distl､ibutionofstatesfbl･ thischal･actel･ isthel､efbreequallycon-

sistentwithbothhypotheseS.

InAvesand inmicroraptoI-iansothel- thallBα"76"qp/oﾉ急,pe-

nultimatephalangesonpedaldigitsIIandlVarelongel･ than

morepl-oximalphalanges. TheseproportiollssuggestarboI-eal-

ity (Hopso11,2001 ;ZhoLlandFal･low,2()01 ) . Thesetaxaphylo-

genetically bracket βα"7〃/･"/oﾉ・ andD1-omacosauridae、 In

whichall penultimatCpedal phalangcsa1℃shortel･ thanmore

proximal phalanges. lt ismol℃pal･simonioLIs to inibl･ that

lengthenedpenultimate phalanges evolved independently in

Avcsandmicl･01･aptorians (a2-stepsccnal･io) thantoinferthat

penultimatephalangeswere lcngthened intheparavianances-

tor, thcnshortened inDromacosauridacandβα加6/mp/oﾉ" (a3-

stepscenario) .

lll summary, thephylogeneticdistribLltionoftheflangeon

manualphalanxll-1,humeraldiameter,andlengtheningofpc-

nultimatepedalphalanges,all support thehypothesisthatbirds

andmicrol･aptol･iansacqLliredaerial locomotionandarboreality

independently, 1-atherthanfi･omacommonancestor. Thephylo-

geneticdistl-ibutionsoflもathel-vaneasymmetryanddistaldis-

placcmentofmetatarsalldonotprovideenoughinfbrmationto

supporteithel･hypothesis.

It isworthnoting that the fbl･elimbcharacterstates listedー

aboveneednotrelatetoacrial locomotion. Theydosuggestin-

creasedl-elianccontheibrellmbsfbl･acrodvllamic locomotion，草

butsuchcoLlldhavebecllwing-assistedinclincrunning,ashy-

pothesized lbl･ maniraptorans byDial (2003) andHoltz(2003） ．

Size7)"e"応

BasalmembersoftheoLltgroups toDeinonychosauriawere

rathel･ Small (Fig. 6) . An inc1℃aSe inbOdySize tookplace

withinthecladeMicrOraptOl-ia+Dromacosaul･idae､withasubse-

quentsizedccrease inM/c,ﾉ℃ﾉ輿"/oﾉ･ andahuge leapupwal･din

sizeill4cﾙ///o6(//oﾉ-andぃαﾙﾉ･叩加ﾉ．、 Thelattel･ twotaxaphylo-geneticallvbracket thesmall-sizedtaxonDﾉ℃〃αe““〃"s. Un-~ ごtil thediscoveryofmol･c taxaphylogeneticallybracketedbyJIcﾙ///o6"oﾉ･, [//αﾙﾉ･"/oﾉ",andDﾉひ"7αe“α"/w鵬『, it is impossibletodeterminewhethel･ lal-gesizewasancestral fbl･ theSethreeｰtaxaandlost inD/℃"7αe“α"ﾉ･",$orwasattainedindependentlyin4cﾙ///()6qroノ- and (〃α力/."p/Oﾉ･. Bothscenarios involvetwoevolutionalySteps,andareequallyparsimonious.

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〃e"cβ“/"o"Q/小ﾉ".Ph.D.dissertation,Northemll linois

UI1ivcrsity,259pp.

Sereno、P.C( 1997) :Theoriginandevolutionofdillosaurs.

』"""α/Rewewq/､Eα"ルα"JP/α"e/qrySc/e"c",25: 435-489．

Serello, P.C( 1998) :Arationalefbrphylogcncticdefinitions,

withapplicationtothehigher-Icvel taxonomyofDinosau-

ria.Neues JahrbuchftirGeologieundPalaontoIogieAb-

halldlullgell.210:41-83.シ ダ

Sereno,P.C. (1999) :Theevolutionofdinosaul･sSc/e/7ce,284:

2137-2147.

Sercllo, P.CCRao, andJ. Li (2002) : S//7oﾉ"パ‘､α"/e/7Sﾉ‘F

(Avcs:Enantiomithes) fi･omtheEarIVC1･etaccoLIsofnorth-

eastcl-nChina・ InLM.ChiappeandL.M.W1tmel･ (eds.) ,

Mｾ."zo/cB/ﾉﾎ."加''e//7e"""Q/､//7eD//7o""ﾊ，､UnivCl･-

sityofCalifbrniaPress,Berkeley, 184-208.

Sues,H-D. (1978) :Anewsmall theropoddinosaurhomthe

JudithRivel､Formation (Campanian) ofAlberta,Canada.

Zoo/0g/cw/JOz"~"α/o/.//7e〃""eα〃Soc/e",62: 381-400.Sues.H.-D. ( 1997) :OnChirostenotes. aLateCI･etaceousovi-

raptol･osaL11･ (Dinosaul･ia:Theropoda) fi･omwestemNol･th

America.Jbz"･ﾉ7(7/Q/陸ﾉ･花か"GPq/eo"/o/ogV, 17:698-716.

Swoffbrd,D. L. (2002) :P/ly/oge"e"c'""α"/sUr/"gP"/w-

〃7o/駆陀バ/o"4.0h/0/t""Mqc/"/"ﾙ. Sinauel-Associates,

Inc.,Sunderland,Massachusetts.

Wcishampel,D.B.andC-MJianu. (1996) :NewtheroPoddi-

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aOntologicAbhandlungen，200: 387-404.

Xu,X. ,YCheng,X.Wang,andC.Chang. (2002) :Anunusual

oviraptol-osauriandinosaul-hOmChina・Na"""e,419: 291-

293．

Xu,X,M.A､Norell,X.Wang,P. J.Makovicky, andX.Wu.

(2002) :Abasal tl･oodontidfmmtheEarlyC1･etacousof

China.A/""〃e,415: 780-784.

Xu,X.,Z.Tan9, andX.-L.Wang. (1999) :Atherizillosauroid

dinosaurwith integLImentarystl-uctures廿omChina. ノV"－

〃"で,399： 350-354．

Xu,X. andXWang. (2000) :Troodontid-likepes inthedro-

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Xu,X.,X.Wang,andXWLl. (1999) :Adromaeosauriddino-

saurwithafilamentous intcgumentfromtheYixianForma-

tionofChina・MJ""で,401: 262-266.

Xu,X.,ZZhoLI,andR.OPrum. (2001) :Branchedintegumcn-

tal stlucmres inS//70ﾉ”"〃0m"ﾉ･"sandthcoriginoffbath-

ers.MY""で,410:200-203.

Xu,X.,Z.Zhou, andX.Wang． (2000) :Thesmallestkllown

non-aviantheropoddinosaur. /Vt"z"で,408: 705-708.

Xu,X.,Z.Zhou,X.Wang,X.Kuang, F.Zhang, andX.Du.

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335－340．

Zhou,Z. andJOFarlow. (2001) : Flightcapabilityandhabits

ofCo)1/i/c"/so/･"/s. In ~lGauthicl･ andL. F.Gall (eds.) ,

ノVEwPeﾉW)ec/n》eSO"/ﾙeOﾉfg/〃α"cIE"'･IvEvo/""o/7Q/

B/ﾉ称.PeabodyMLIscumofNaturalHistoly,NcwHaven,

Connecticut,237-254.

Zhou,Z.,X.Wang,F.Zhang,andX.Xu､2000) : Importantfba-

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SI]NTl]RPhil，BARSBoLDRinChCn、BRITTBrooksB.、andBLIRNl.lAMDavidA'2

Appendix l anteriol･teeth, (1 ) presentonall teeth. (Houeta1., 1996)22.Numberofdentarvteeth (0)30. (similar

charactel- inNorcll etal.､2001)

23．DenticIcsondistalkeclsofteeth-(0) present (at least inL

posteriol･ teeth) , gently l･oLInded, (1) absent (all teeth) ,

(2) prese'lt (at least inposteriol･ tceth) , apicallypointed.

(Norelletal . ､2001)

24.Dentary-(O) toothed,(1) edentulous.(Sues,1997)

25.Dentaly-(0) notboweddownward, (1) boweddownwal･d

(concavedorsally) .

26. Posteriol･ surangulal･ fbramen-(0) Small, (1) enlarged.

(Holtz, 1994)

27. Articulal･ glenoid-(O) 11ot vclltrallydiSplaCcd､1) ven-

trallydisplaced. (MaryanSkaeta1.,2002)

28.Mid-length incrcase inhcightofdental･y-(0) absent， (1)

pl.csellt.

29． Stronganglingofantcriorflccofcervical centra､ toface

largelyventrally (O) absent, (1) p'℃sent，

30. Lengthofmid-cel･vlcal cel1tl･a-(0) sLlbeqLlal to lengthofdOrSal cCntra、 (1)markcdlv Ioll2el･ thandorsal centra.

－ －

(Evans,1988)

31.Lengthofmid-cel･vicalcentra-(0)

SystematicsandEvoILItionofDrolnaeosaLIridac (Dinosalll･ia,Theropoda) 13

faceslaterally. (NovaSandPuerta, 1997)

48.Humerallength-(O) femoral

length. (Evans, 1988)

49.Lengthofhumeralshafibetweendeltopectoralcrestanddis-talcondvles (0)3xdiame-

tel･ofshaft.

50.LengthofradiLIs (0)femoral length. (Senter,2003)51 .Radialdiameter-(O)>05xulnardiametel･， ( 1 )

SIzNTI.]

SystematicsandEvoILltioI]ofDromaeosauridae (Dinosalll､ia,Theropoda) 15

ＶｍＩⅢⅢ川小》"

(~~' HH

克動 一恵。1 1

Eせぞ」 房こぜ＞DD EE

国司

圏FF

匡－1コ

e，＜

GG

showingstateO (11euralspinctall) .Right,posterioI･doTsalvcl-tcbraofﾙ"cﾉw~qp/oﾉ･z/7[Jo/f""（(aiterHwangeta1.,2002),showingstatcl (''cu'･al

spillcshort) .MChal･actc1･44: anglcbctwccnacl･omionpl･occssandscapulal･blade.Lch, scal)Lllaof,4//o|"""パルJg///s (211Tcl･Madscn, 1976) ,

showingstateO (al]glestrong) .Right, scapLllaofRaﾙ()"αw､§' (）‘W(ﾉ"〃 (drawnifomYPMcaSiofUA8656),showingstatcl (angleweak)N.Chal･actc1･45: overhangofacl･omionpl･occss. Lefi, scapLllaol､ ｣4//"[""･".『"g///s (anel･Madsen， 1976),showingstateO (ovcThangabscnt) .

Right､ScapulaofRqﾙｲ)"αwざ“"℃"7/ (drawn廿omYPMcasto｢UA8656) , showingstatc l (ovcrhallgpresent) _0.Characte1･46:pl･oximal coll-

strictionofcoracoid.Top,coracoidofDe//7om'('ﾙ"sα""ﾉ･ﾉ”"4, (afierOstrom, 1969) , showingstateO (collstrictionabscll(IBottom､col･acoidof

S/"oﾉ･""力“〔""･"s〃7ノル""〃(aficrXLIetal.1999),showingstalel (constrictionprescnt) ．P.Charactcr54:lengthofmetacarpalll.Top,manusofDe/ﾉ70ﾉ”c/7"sc"7"ﾉ･ﾉひp"s (aitcl･Ostrom, 1969) , showingstatc() (mctaca叩alllshortel･ thanmctacarpall+mamlalphalanxl-1).Middle,manLIs

ofNGMC91,showingstatcl (lengthofmetacall)allIsLlbequaltothatofmetacarpall+manualphalanxI-l).Botlom,manLIsof脳"o/"ﾉJ〃7〃‐

""/ (IVPPV12558),sI'owingstalc2 (metacarpallllongcl･ thanlcngthofmetacarpall+manufllphalanxl-liQChal･actc1.55: lel1gthofman-Llal l)11alallx II-2.Top,mallLISoI､NGMC91,showingstatel (phalanxII-2

SENTIzRPhil、BARsBoLDRinchen,BRITTB1.ooksB.,andBuRNHAMDavidA16

Appendixl l

Charactcl--taxonmatrixusedibrphylogencticanalysis

K〃αα〃〃7cAe""m

l lOlOO1 10？ ？2001 101－－

0101 1 12000 0?01 101010

000()000000 1001000000

1000000101 0

Pﾉ"O/α庇'〃αgOp/e/"yx

l lO]OOl lO？ ？2001 1001 1

01011?2000 0?0??O?010

000100???0 100100???0

1000?0?101 0

O〃7"ﾙo/e,sr"ルピノ曇"?α"ﾉ7ノ

1001000??? 01101100??

0001110000 0??????010

?00??000?0 00000?0000

l?0????0?1 ?

AFCﾙaeO/)花ﾉ:)w/"加gﾉ･"/7Icq021 1001 101 0200010011

1 101 1 1 101 1 ？01 1 101 1 10

000101 1110 1011001111

1000001 101 0

c/e/7o/oﾉ･ﾉ応pﾉ．/"7αI？1 1？？1 1？？ ？？？00？00？‐

1 1?1??00] 1 ??llllll ll

O1 1 1001 1 1？ 1 1？lOOl l()1

?()()0???101 0

)t"7oﾉwパ〃7α〃"〃

021 1??1 1?? ??000?101 1

l l???????? ?11??ll lll

l l l lOO?1 l? ????O??l??

1000101 101 1

S/"or/7ﾉ‘y”ﾉ7/e"sJs

021 1??1 1?? ??000?lOl l

？？？’？？-‐1 1 ？l l ll l？1 1 1

01 1 1001 1 1- 01--0011 1 1

1000001 101 1

R(ﾉﾙo"αw‘ｻ0，W℃"〃

？「>ウワ‘)ワワワワヮ ワウワワワワワヮワワ

??01 11??1 1 00?11?l??1

？？？？？？l l l l llllOOl l l l

lOl l？l？101 O

Sﾉ"Clﾉe"α/Oﾉｰcﾙα"g〃

?00?001?01 0200????11

01 1 1 1 1???? -011?01???

0??10?1 1 10 10110?01?l

?01 1???1()1 0

M/cﾉ℃ﾉ鋼叩/Oﾉ"2ﾙαo/α""s

01???????? ??1?????1 1

？？l l lOOl l l －0111？lOlO

l?1 1001 1 1 1 101 1??0101

001 1 1 1 11 1 1 0

'!Mcﾉ℃ﾉ忽"/o/"g"/'!

ワワワワワワワ？” ？？？？？10？？？

？0？’ ］？01 1 1 ？？111？lOlO

ll l lOO？’？1 1？lll l？’？？

？？？’？l l l l l O

1!CﾉW)/01ﾉo/f"ﾊﾉﾌα"〃！0????????? ?????????l

??????01 1 1 ???1??1010

l l l lOO？1 1？ 1？？11101？？

---1001001

000()000111

1 110000100

4//"f""WSβ蕊αg"/s0000000000 0000000000

0000000000 0000000000

0000000000 0000000000

0000000000 0

7W卸α"ﾉ7““"WSノゼ･x

0000000011 0001100000

0000000000 0000000000

0000000000 0000000000

0000000000 0

S/""(""可)/e/yX/刀w"α

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0111110010 -000000000

0000000000 1000000000

1000000101 0

S""｡o"7"ﾙo/"s/z""oﾉ〃

?01 101 1 101 1 100210002

???????000 ???????ワ”

ワワワ‘フワワワ‘)？ワ ヮワ？ワワワ77ワワ

?????????1 ?

S""汐ﾉw"ﾙo/"s"7o"go//e"s/'s'

?01 101 1 1?? ??0021??02

??llll??O? -0????????

??？?？？？0?0 ？0？0？？？？？？

1 1 1 1？1010？ ‘）

S"70ﾉ･"〃ﾙo/d@s)ﾉo""gI

lOl lOl l??? ???02?0?l l

l l l l??1001 -??0001010

10?100?0?0 ?0?0??????

l l l lOOl l？1 0

乃ooclb""〃ﾉ綱"70.W"s

101 (ﾉO1?l l l l l?????002

?1 1 11??0?? -?????????

??0??????? ?01?00????

01 1 1000??1 0

Pe/ecα"/"7/"7"s' JIov"/

?00101 1 1 1 1 ??0021 101 1

1 111?????? -??0001???

00?100???? ??????????

ウワワワ‘)7‘)ワウワ ワ

SY/w/ﾙ/o"""7zパα""‘f

l lO101 1 1 1 1 1 10021 10--

l l l l l l lOOO -000001010

0001000000 0000000000

1 100000101 0

Bw℃"“〔"""sﾉ‘〃で／?Ol?O1 1??l ?2102???ll

ワワワワワワワワワワ ワワワヮワワヮワワワ

‘77‘>9？ワワワワワ ワワヮワワワ‘)？‘)？

?11??????? ?

Cα"C/y)/e}:'wzo'〃

1 1010()1 1？？ ？？001 1 1 1 1‐

0101 1 12000 0001101010

()001000000 1001000000

1()00000101 0

0()0000000(）

0000000000

00000000?0

lOlOO？lOOO

0000110111

lllO?00?00

0000010000

000011-000

00001001？0

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00??0?001?

()?0?000??0

()0000()0000

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0010000100

lO10001001

010111011 1

OlllOOlll2

1 1200?00??

？?‘)ワワ‘”ワワワ

?？？？1？？？？？

10-0001001

0102011011

01?IO???12

l?200?O???

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OllllOO101

lOlOOOlOO？

OlO201-011

？1 1 10？’？l(〕

1?200?0001

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?1 1 1?00101

lOlOO??0?1

000201-011

011 1011-12

1 1200?000?

?????????1

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‘)‘〉？？77‘ﾌワワゥ

0？？？？？？？？？

()l l lOl l？10

lllOOO1001

O？101 1()000

｢>ワワr>り‘)r)‘>ワワ

10100??010

?????1????

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0010110000

001？1001？1

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1001011111

0011011002

lllOO??0??

‘)‘)‘7777‘〉？‘)‘）

‘)〈)‘>ツワr)‘)ワ‘)‘）

??？？？？？??0

1001011111

0?1l???00?

1--1001000

0000010010

1ll000()100

0()20?????？

100101111 1

??l l???00?

17SystematicsandEvolutionofDromaeosaul･idac (Dinosauria､Theropoda)

？0？1 1？l l l I O

S"70ﾉ･""ﾙ“αz"w‘§‘〃""E"〃

0101 101 1？？ 0？lOOlOOl2

？？？？？？？’？？ ？01 1 1 1 1010

1 10100?l 1 1 101 1 1 1????

001 101 1101 0

NGMC91

01011011？？ ？？lOOlOOl l

？？？？？？1 1？？ ？01 1 1 1 1010

110100???? ??????????

001 101 110？ O

Bα"76"〃/O『､た加beﾉg/0101 101102 0210010002

0000001 1 1 1 001 1 1 1 1010

1001001110 101 1000101

01 1 1？lOlOl O

"/o['//W/)/oJ-"70"go//e"~,'八、

0101 100012 01 1()010002

0001001111 001 1001010

0001 1 1 1 120 1001000101

01 1 1010101 0

此""．oﾉ‘""ﾙo/"/cs/α"gWo"ノ?？？？？？？0？？ ？??????002

0000?0???? 0????????0

????111??? ??????0101

??11?????0 0

，ﾉ．o"7αe““"WSα/6e/"/e/7s/4｢

??????0012 01?10?0000

ワワ‘フワ‘)「)ワワワ‘フ ワ‘)？ワヮワ‘)ワワワ

ワ‘>ワワワワヮ‘ﾌワワ ‘フワワワ‘>‘)ワワワワ

011??????? ?

批此ﾉ”"ﾉWS〃わ"go〃e"』‘耐??????O?1? ?????100??

00?1???l?? ?O????l???

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[//αﾙﾉ忽"/oﾉ･“"℃"7"7α”ノ?1???????? ????????00

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??0100???? 00?1 1?1?l?

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？？？？？？？10？ ？

〃"航""“DCﾉ･/e/よ ユ

ワワワワワワワヮワワ ワヮヮワ《ﾌワワワ？？

ワワワワワワワワ‘フワ ワワワワワ‘ﾌワウワワ

？？‘>‘)？‘)77‘)【） ‘>77‘)‘)ｳ‘)‘)‘)‘)

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Appendixl l l

SynapomorphyliSt

Theverbaldescriptiollofeachsynapomolphyispl･ecededby

thellumberbvwhichthechal･actel- iscataloguedinthechal-actel･シ －

list (Appendixl) 、 fbllowedillparenthesesbythenumbel･repre-

sentingthccharactel-state. Fol､example, 1 16(0今l)"refb,･stoachange instate ii-omOto l fbrcharactel- 6. InACCTRAN

ｰ

mode,PAUP4.0bassumes that fbragivenclade,achanged

charactcl･ stateisasynapomoI-phyevenifitcanbeconilrmedin

onlyonetcrmillal taxoll. SLIchdubioLIs ! 'synapomorphicsI1 are

omitted ill lheACCTRANlistshere. Salientsynapomorphies

ofDcinonychosauriaallditssub-cladesareillustratedinFig.3.ツ

nameanditsdefInitionarealsogiven.Here,namingoftaxafbl-

Iows the rules ofphylogenetic taxonomy (DeQueiroz and

GaLlthier, 1990) , e.9 (1) oI11ycladeS aregiven taxonomlc

names､2) thecarlieStpllblishcdphylogeneticdefinitio11ofa

taxonOmicnamehaspriol･ityovel･ subscqucntdefinitionS, (3) aphvlogeneticdefinitionmavbe stem-based (all taxamol･e且 ‐ ｰ ツ

cIoselyl･elatedtoAthantoB) ol･ 11ode-based (all taxaphyloge-

neticallybracketedbyAandB; that is,AandB, theircommon

anccstor､andallofitsdescendants).Mel･elistsofincludedtaxa

arcnotcOnsideredvalidphylogencticdefinitio11s.

Coclurosau1･ia:allneotetanuransmorecIoselvrelatedtoNeomi-ジ

thesthantoAIIosaurus (Sereno,1998)

ACCTRANandDELTRAN: 10(0今l) :Caudaltympanicre-

cesspresent. 88(0->1) :MetatarsalVl) : Paroccipitalprocesses not

strollglydowlltumed. 15(0->l) :Lengthofprcorbital l･cgionof

craniLIm

l ) :Manualphalanxll-2Zl3xlcngthofmanualphalanxll-l.

56(0->l) :Manualunguallshortcl･thanmanualphalanxl-l.

O『""ﾙo/ey/es+(S/""""雪o"/erMx+Maniraptorifbnnes) .ユ ヴ ム

ACCTRANandDELTRAN: 4(0->1) : lntemalan(ol･bital

fbnestl･asmallcrthanol･bit. 12(0->1 ) :Transversewidthoffbra-

menmagnumsubequaltothatofoccipitalcondylcl6(0->1) :

FIolltai processofpostorbital verticallyoricntcd. 34(0->1) :

Lcngthofdorsalcentrasubcqualtool･>height.35(()->1) :Pos-

tcriol･ dorsal neural spincs< 1 .5xtallerthananteroposteriorly

lon9. 36(0->1) : Postcrio,･ dol･sal parapophyscsnot Seton

stalks (nonpedunculate).91(0->1) : Tbtallengthofpedalpha-

lanxII-IZ3xlengthofdistalcondylal､eminence. 100(0->l) :

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SIzNTERPhil,BARsBoLDRinchen，BRITTBrooksB.,andBuRNI.IAMDavidA'8

PI-omincntlateralflangeatproximalendofmetatal･salIVabsent.

ACCTRANollly: 1 (0今1) :C]･anial length≦0.8xfemoral

length. 8(0-~l) : Postorbitalproccssoffi･ontalnotsharplyde-

ma1℃ated. 19(0->1 ) :DenticlesabSelltonpl･emaxillalytceth.

20(0->l) :Denticlesabsentonmesialkeelsofposteriorteeth.

49(0->l ) :Lengthofhulneral shaiibetweendeltopectoralcrestanddistalcondvIcs>3xshalidiamcter.

ジ

S/"“α"m/)/eﾉ訳x+ManiraptOrifbrmes.

ACCTRANandDELTRAN: 7(0->1) :Lengthofmaxillary

processofjugal>4xheight.32(0->1) :Cervicalneuralspincslow、 83(0->l) :LengthofmetatarsL1s>4xpl･oximal transverse

ｰ △

width. 98(0->1 ) :Tibialongel･ thanlbmur.ｰ

ACCTRANonly: 15(1->0) :Lengthofprcol･bitalregionof

craniumZ2xheightatantcriol･cdgeofprcorbitalbar.33(0->1)

: P11eumatoporesabsentondorsalvertcbl･ae.71(0->1) :Ischial

length1) :Postorbitalproccssoffontalnot

sharplydemarcated、 19(0->1) :Dcnticlesabsentonpremaxil-

lalyteeth. 20(0->1 ) :Denticlesabsentonmesialkeelsofposte-

l･iOr teeth 55(0->1) :ManualphalanxlI-2Z13xlengthof

phalanxII-1.

Mallil･aPtorifbrmes: all taxaphylogeneticallybracketedbyOﾉ制一

ﾉ7"ﾙo"7"""sandbirdS (Holtz, 1996) .

ACCTRANandDELTRAN: 21(0->1) :Teethconstrictedbe-

tweenl･ootandcrown. 23(0->l) :Dcllticlesabsentondistal

kcelsofposteriol･teeth. 37(0->1 ) : Lengthofbo11ytaillengthofpresacralseries. 64(0->1) :

Lengthofmanualphalanxl-l>5xdiameter.

ACCTRAN: 27(0->1) : Al･ticLllal･ glenoidventrallydis-

placed. 47(0->1) :Pectoralglenoidfilces Iaterally. 60(0->1) :

Distalcarpals land2with scmilunate shape. 82(0->1) :Greatel･andanteriortrochantersfilsedintotl-ochantericcrest、 84

(0->1) :Metatarsalldistallydisplaced.DELTRAN: 49(0->l) :Lengthofhumclal shafibetweendel-

ｰ

topectoralcrestanddistalcondylesZ3xshaltdiameter. 56(0->

I ) :Manualunguallshortel･ thanmanualphalanxl-1.

AI℃tometatarsalia: all taxamol･e closelv related toOﾉ･‐ご

""ﾙo"7ﾉ"7//sthantobirds (Holtz､ 1996) .

Bullatosauria: all taxa phylogenetically bracketed byOﾉ輿一

ﾉ7"ﾙo"""7"sand7)･oo"'7 (Holtz, 1996) .

ACCTRANandDELTRAN: 6(0->1) :Maxillaryfenestraen-

largedandanteroposteriorlyelongatell(0->1) :Parasphenoid

basebulbous. 15(0->2) : Lengthofprcorbitalregionofcra-nium>3xhcightatanteriorbordel･ofPl･corbitalbar. 22(0->1)~ ユ:Dentarywith>30tCcth. 30(0->1 ) :CcTvical centraSignill-cantlylongcrthandorsalcentra. 31(0->l) :CervicalcentraZ3xlongel､ thantal1. 85(0->l) :Anteropl･oximalcontactbetween111etatarsalsI1andlV90(0->l) : LengthofpedalphalanxlI-2=06xlengthofpedalphalanxll-l.92(0->1) :TotallengthofDcdal l)halanxll-21) :Craniallength1) : CraniOmandibularjointsubol･bita1. 53(0->1) :Metacal･pal l>0.8xlcngthofmeta-carpalll.ACCTRANonly: 60(l->0) :Distalcarpalsland2notsemi-

lunal･ illshape.

DELTRANonly: 9(0->l) : Pal･occipital proccsscs not

stl･onglydowntumed. 27(0->1 ) :Articulal･glenOidventrallydis-

placed.

Troodontidae (newdelinition; seeDiscLIssion) : all taxamore

cIoselyrelatedto刀℃o""thantoOﾉw"ﾙo"7"""s，Mb"o"yA"s,

7ｿ7e"z/"o.9""･"s, P@/oc,〃叩rOﾉ･,0Wﾉてゆ/Oﾉ",01･Avcs.

ACCTRANandDELTRAN: 3(0->1) :Maxillacontactsex-

tcmalnaris. 93(0->1) :Posteroventral "lip! 'onpedalphalallxll-旬ノー●

ACCTRANonly: 27(1->0) :Articulal･glenoidnOtventrally

displaced. 73(0->l ) : Lossofelongatccxtensionofischial

shafidistal toobtL11･atorproccss. 94(()->1 ) :Pedalungualllen-

larged.

s/"o"7"ﾙo/d@s+ (刀℃o""+SL""ひ"7"ﾙOﾉ火s"7o"go"e"s/IF+

助"ﾉ℃"7"ﾙo/ctJs/""/o『) ．

ACCTRANandDELTRAN: 23(1->2) :Denticlespresenton

distalkeelsofpostel-iol- teeth,apicallypointed.

DELTRANonly: 60(0->1) :Distalcarpals land2with

semilunateshape. 82(0->1) :GI℃aterandanteriortrochantel-sfilsed into trochantericcl･est. 84(0->1) :Metatarsal ldistallv

displaccd.94(0->1) :Pedalungualllcnlarged.

7)Y)o(ﾉt)"+Sm""oﾉw"〃oただw77o"go//e"&,､/､w+Sα"ﾉ℃/w"ﾙojdbs｡/""/oﾉ銅．

ACCTRANandDELTRAN: 19(1->0) :DcnticlespreScnton

prCmaxillaryteeth、 20(1->2) : Denticles present onmesial

keelsofposteriorteeth,withmoredenticlespel･5mmthanon

distalkeels.

Maniraptora: allcoelurosaursmol-ecIoselyrelatcdtobirdsthan

toO1･nithomimidae (Gauthier, 1986) .

ACCTRANandDELﾉTRAN: 12(1->2) :T1･ansverSewidthof

lbramcnmagnum>thatofoccipitalcondyle.44(0->1) :Anglebetweenacromionprocessandscapularblade>160｡.45(0今

△ ユ

1 ) : Overhangofacromion antcriol･ to anteriol･mal･ginof

scapLIIo-coracoidal suture 58(0->1 ) : Proxilnodorsal "1ip"011

manualungual l1. 59(0->1) :Tipofmanualungual levelwith

Hexol･ tubel℃le. 74(0->1) :ObturatoI･processdistallydisplaced.

ACCTRANonly: 2(0->l) :AnteriOrbordel･ ofpremaxilla

slantcdposteriorlyat50｡ -70｡ . 9(1->0) : Paroccipitalproc-

essesstl･onglydowntumed. 33(1->0) : PneumatoporespresentondOrSalvertcbrae.

DELTRANonly:60(0->1 ) :Distalcarpalslalld2withsemi-

lunateshape.71(0->1) :Ischiallength1) :Lengthofpreorbital

regionofcranium2) :Lengthofbonytail1) : IschialshafIstl･onglyconcaveposteriorly.ACCTRANonlv: 84(l->0) :Metatal･sallllotdistallvdis-

一 ジ~

placed.

DELTRANonly: 1(O今1) :Cranial length1) :Anteriol･bordel･ofpl℃maxillaslantedposteri-

orlyatSO｡ -70｡ . 27(0->1 ) :Articulal･glenoidventrallydis-placed.エ~

SystematicsandEvolutiollofDromaeosauridae (DinosaLIria,Thcropoda) 19

C"/c//p/e/γx+K伽α"･

ACCTRANandDELTRAN: 18(0今I ) : Parietals longerthanfi･ontals.24(0->1) :DentaryedentuloLIs.55(1->0) :Man-

ualphalanxll-2l) :Anterior hemal

arches1) :Anteri-

ormostdistally-expalldedhemalarchanteriol･tocaudal9. 43(0

->l) :Humerus longel･ thanscapLlla. 67(0->1 ) :Antel･oventl･al

comerofiliacbladeposteriorlydisplaced. 68(0->1) :Retro-

pubypresenL69(0->1) :Anteriorportionofpubicbootabsent.73(0->1 ) :Lossofelo112atcextensionofischial shafidistal to

-

obturatol･ pl･ocess 78(0->1 ) : Posteriol、 endof iliacblade

pointed80(0-W) : Ischialpeduncleofiliumrcducedinheightrclativetopubicpedullclc. 87(0->1) :Metatarsalllldistallv

△ 一

ginQIvmoid.ｰ ｰダ

ACCTRANolllv: 1 ( I->0) :C1･anial length>0.9xfemol･alご ー

length.54(0->1) :Metacarpalllsubequalinlengthtometacar-

pall+manLlalphalanxI-l.

Aves: all taxaphvIogeneticallvbl･acketedbvAﾉでﾙα“､花ﾉ"wand人ゴ ー ヴ ヴ ユ該

Omithurae (Se'･eno、 1997) .

ACCTRANandDELTRAN: 2(0,1->2) :Anteriorbordel､of

premaxillaslantedposteriorlyat≦43.3(0->1) :Maxillacon-

tacts external llal･iS 30(0->1 ) : Cervical vertebracmarktlV

lOngel･ thandorsals. 31(0->1) : CervicalcentraZ3xlongel･

thantall. 48(0->1) :HLImel･LIs Iongel･ thanfemur. 52(0->1) :

LengthofmanusZl.25femorallength77(0->1) :Lengthof

iliumanteriol- tomid-actctabulumlnarkedlylongerthanlength

ofiliumpostcriol･tomid-acetabLllull1. 79(0->1) : Iliacbladean-ユ

teriorlyroLI11ded. Hallux1･ctroverted. 90(0今2) :Pedalphalanx

II-2ZlellgthofpedalphalanxII-l. 97(0->1) : Pedalphalanx

IV-41ongcl･ thanpedalphalanxIV-3.

DEIJTRANonlv: 27(0->1 ) :Articulal･glenoidventrallvdis-学 ~ 亭placed. 84(0今l) :MetatarSalldiStallydisplaced.Omithul-ae: all taxamorecloselvrelatedtoextantbirdsthanto‐4/℃ﾙqeOp/e/"(Gauthier, 1986)ACCTRANandDELTRAN: 50(0->1) : RadiusmarkedlvlOngel･thanfemu1-． 46(0->1) ;Coracoidproximallyconstricted.54(0,1->2) :MetacarpallllongerthanmetacalpaII+manualphalanxl-l. 57(0->1) :Manualphalanxlll-20) : Proximodorsal !!lip"absentonmanualungual ll62(0->1 ) :Diametel･ofmanualphalanxll-lenlargedbyposteriOl･ flangc63(0->l) :Humeraldiametel･Zfemoral diamcte1･． 72(0今l ) : Ischiumwithstrongproximal,posterodorsalproccss. 86(0->l) :Metatarsal lldistallygingly-G1molq．

surftlcesofcervical centraface la唱elyventrally. 93(0->1) :

Posteroventral "lip"onpedalphalanxll-2. 94(0今l) :Pedalun-

gual2enlarged.

ACCTRAN: 27"( 1-,0) :AITicularglenoidnotventrallydis-

placed. 38(0->l) : Distal caudal zygapophyses longel･ thana

centrum. 91(0-,l) :Totallengthofpedalphalanxll-ll) :MaxillaryfeneStra

dorsallydisplaccd・ 10(I-,2) : Caudal tympanicrecessdis-

placedontoparoccipitalpl･occss. 20(1->2) :Denticlespresenl

onmesialkeelsofposteriorteeth,withmoredenticlespermm

thanondistalkecls 21(()->1) : Lossofconstrictionbetween

toothcrownandroot,at leastfbrantel･iorteeth. 23( 1->2) :Den-

ticlesprcsentondistalkeelsofpostel-iorteeth,apicallypointed.

Dentarybowed (collcavedorsally).32(1->0) :Cervicalneural

spineshigh. 36(1->0) : Posteriol･ dorsalparapophysesseton

stalkS86(0->1 ) :Metatarsal lldistallyginglymoid. 92(0->1 )

:Total lengthofpcdalphalal]xll-2=2xlengthofdistalcondy-lal･eminencc.

ACCTRANonly: 13(()->1) :Nasalsanteriorlyuptumed.19

(1->0) :Denticlespresentonpremaxillaryteeth. 35(1->0) :

PosteriordorsalneuralspinesZl.5xhigherthananteroposteri-

orlylOng.

DELTRANonly: 2(0->l) :Anteriorbol･del･ ofpremaxilla

slantedpostcl･iol･lyat50｡ -70｡ . 38(0->1) :Distalcaudalpl･ezy-

gapophysesIongel･ thanacentrum96(0->l) ;Pedalphalanxll-

l shortel･ thanpedalphalanxlV-l _

MiCroraptol･ia (11ewtaxon) : all taxamol･ecIoselyrelatedtoMi-

croraptorthantolt/oc/ﾉ")/o/-OI-D/･0"7qe"'""ws.

ACCTRANandDEIJTRAN: 46(0->1 ) : Coracoidproxi-

mallyconstl･icted. 57(0今l) :Manualphalanxlll-1Z2xlength

ofphalanx II1-2. 61 (0->1) :Manualphalallxl-lbowed (coll-

caveventrally).88(l->0) :MetatarsalVZ0.5xlengthofmeta-tal･sal lV.

DELTRAN: 13(0->1) :NasalsanteriOrlyuptul･ned. 54(0->

l ) :MetacarpalIIsubequalinlengthtometacamall+manual

phalanxl-l.84(0->l) :Metatarsalldistallydisplaced.

S"7oﾉ""ﾙo"zI/ws+ (NGMC91+ (A"a℃ﾉ雪"/oﾉ･Zﾙαo""7"s＋

1 1/Wcmmp/oﾉ.gz"Ⅷ+' !CﾉW)rolﾉo/q"s1 1) ) ,ACCTRANandDELTRAN: 51 (0->l ) :Radialdiametel･<

0.5xulnal･diamcte1･. 55(1->0) :Manualphalanx ll-2l) :Diameterofmanualphalanx

II-lincrcascdbyposteriorflangc. 70(0->l) :Tipofobmrator

process fbrmsastrollglyacuteangla 75(0->l) : PLlbicshafi

kinkedpostel･iorlyatmid-length. 76(0->l) : Posteriorpartof

pubicboot lost. 90(0->2) :Pedalphalanxll-2≧lcngthofpha-

lanxlI-l. 97(0->l) : PedalphalanxlV-41ongel･ thanphalanxIV-3.

ACCTRANonly: 19(0->1) :Lossofdenticlcsonpremaxil-

laryteeth. 35(0->1 ) :DoI･salncuralspines0) : Totallcngthofpedalpha-lanxlI-2>2xlengthofdistalcondvlareminence.

NGMC91 + (M/cﾉ℃/｡q/)/oﾉ･亘ﾙαo/α""‘『+ ! IM7cﾉ．oﾉ.cmroﾉ"gz//' 1+

"C心仰ovo/αﾉﾊ'')ACCTRANandDELTRAN: 20(0->l) :Denticlesabsenton

Omithothoraces: all taxaphylogeneticallybracketedbyS加oﾉwﾉs

andNeornithcs (SerellO, 1998) .

ACCTRANandDELTRAN: 17(0->1 ) :Craniomandiblllal~

jointsLlborbital.71(1->0) :Ischium>0.7xpubiclength.101(0->1) :Fibuladocsnotreachtarsus.

DELTRANonly: 42(0今l ) : Sacrumwith>8vcrtebrae.

Deillo11ychosaul･ia:all taxamol･ecIosclyrclatedtoDe/"oﾉWcﾙ"s､

thantoNeomithes (Padianetal.,1999) .

ACCTRANandDELTRAN: 29(0今l) :Anteriorarticula'

SENTERPhil，BARsB()LDRillchen,BRITTBrooksB.,andBIJRj,IHAMDavidA20

I]-2=2xlengthofdistalcondylareminencc.

Dromaeosaurinae: alldl-omacosauridsmorecIoselvrelatedto一

DFC"?αe“αz"調"sthantoIt/oc""/oﾉ" (SCrcllO, 1998) .ACCTRANandDELTRAN: 100(0->l) :Metatal･sallVwith

pl･ominent latel-alnangcatproximalextremity.

ACCTRANolllv: 54(1->0) :Lell2thofmanualphalanxl-1ジ ー ‘

0) :Dentarystraight.

DELﾉTRANonly: 64(1->0) :Lengthofmanualphalanxl-l<5xdiameter.

DromaeosaLll･idac: all taxaphylogeneticallybracketedbyDﾉ､o－

"7α””""耐andlt/Dc""