A PHYLOGENY AND CLASSIFICATION OF THE HIGHER CATEGORIES OF TURTLES EUGENE S. GAFFNEY BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY VOLUME 155.: ARTICLE 5 NEW YORK : 1975 __ - --- -. -"I-.,'11, ., '_., I,.. II""",.-.,,,.__.;;_:.e.;i:,;, ",.i,,;,,._ I .r....I!. ," _A !I. "' v--..1! -I "I I!,.;* I,. .....-""," Ml;,_. ,4%'. -,",.-,.A-r--'...-;,',,_-, ",,ill;,',:,. .,,,,- I.,- ," .. ., , 1, "_,,1,.,..,.'-"" _.,,... ". .1._.; ,,",".B..`."17!1_1'1,.`,if',1- ..,,w ._ ....,.",--I 11 lel- 4, 1_1.'11, I,;., ._k_;v.--,-,,.SftI_-1. I.,.Z,.-w-% .141 ", "k: ,l., I,. ,-I.,Iiii.Mll..-,.V_-_II,..',,-1 ,"'.."4..",_-",,.,AIIiti., 47.,,.`_.'..-.-i.--.,,'..,,,.,."',-:.', ..,,,'.,",'..,",-.,:",-,-"--.,",,, -.;,,, '.'.!",I..-, - ..,,9I.-, Pr-,.`!,,.,,ll,...*;-,..,.`_._-.iI., i..,,., ..,,-- ,-;_"".'_iW.J'14'11; ..,..I_, ., .", " ", 1,iw.`-i.11., 0 ,,%._.-iftcx,,, ik,.kN 1,. ,_,_'.'.. .;,. .",.I- - _.,-.;`Ml -I'_'-,.-' 4 ,_.,--.-.., .,,,, --I,"!.;,..,',-,.A4- ._,Av- -,, ll. I..;.e,, ".1I"_i. -.1 ,.. ....._.-1.11 -,".,'.;.-4.4, -, --, ,''N,"II`"_I"".§11 ,.. ,RvF., ", .,,.i., ..,t ;I. ,17.- ,,,..',' .-f _,.., ._-,,-,...,"...,.'-.' ," '--..-_".-RK." ,..wtlil_. .I.'',-"I "'.'.1-1, ..,,.;.7,. ,"'.,, .-,-%".`,.,.Iklr..I,_."'i,11-.,.11....-I-."..:,.....,,,.,,,-_ ,.' ,i.,a7 1, I.,,.-.., -, -,..1,,. ,,:- " rIrrvk"'1.1.1...,, ,,. 4.4,.. jj!kv.,"I'l--1..._-,-.jt .,,',., 1,";.,,. ., 1"',- _-,"'n ,,, "'! ,..-.,.1.-.N.:"A ,,, "'.." ._.,".1. ,--. 'A"1 ''.1 -'. .' .,: '.I-,_..,,.1,I,.4, .;' <-. "-'r ,- -, !!, I_`;-,i-i.,-, , -1II..,.,I..,l.-,,1,.--..: .....". ,-,.f,-,.,`.'..,. 11,-1-.-,.r..,, i. i`_I_" .l -- .- ,.,..,;i-._1, .,.-'1111_1;. 4..,I...I- N,.',,^,4i,"lm'.1.11 .".. ,..`.''_ 1-1114_111..,I .",.,.- I-, -,1.-1.,-,.,.7,,.., .; A,,.,_I .-., ,.. 1! _111, I-,,'.. .. .;..?- ,,,..,,,...11.,I.;.l 1,;. ",,.. ._,.lz,. A!Pi ..; `FW11.., --I ` .',,7,.----,-.;Z,q4S,,..'i,-,71. .",' .."`.....'.l,..z ,-II1Ii,Z..rI._.-.i--.-"I'l,"j".",I,.li,.,.,,..l Ili 1; 1.1.11. -t-,_.11,' I .. 11W -rl-.lc_X..4),.0',,,,,`- 11 1.11.",..,I,.,-, ,,. ,.,.,. A. ..,'..l-,,.,,..i.l-"..:,:.ll..t-I I-- -_.--,-,. .,-.--- , -4_z.-,IV h. -:.lz .4&#-'kf "-,-, `.-,.,I.i',,,; b. .,,,-'-',. ",, .""'.1..O.-..,.-1,. i, -1 .,-- ., ;, ,,V-., ..,j',"' -.-.,,._6X..'l..,..:,, ,Jr -1-I,I"." .,.- -.., "' """-,3'...,,",,,""- N,.'f,.;---...;,",,-""%__.---1-,,,"'`.7,I",,,,,..l., .. .....-.- -f.".. -1 -1 `31 _7 ,:--.,,,.,;,I-l ","w,.j.,- ,t. I,-,Z- .N,"....,,, ',,.r,,.%,t.. fg. -',t .-,. J-P;i.., ."-4"i,I,-1-", ,"pZ:71 1;I.,li,I.I_,&`:, -..,I.l,,&_I_1.,.I!,., .,-1-,",-,.", - ., 1. ..",-,.. -.. ,.;I,,--.: ,,-,-, .l ., --,,,.%-,,.. ,.. .,Y?l.-;ll-'l-'ln,t,,",.6..t',4'r. ,;,., ,i.!." .4'. ..III..,_.- ,-,. ,.'..O'w ,"4,-A ,.,,,,,, _,P:.-. 4.1 ,.4.I-i-I,,,_4 ,.,,.,.,.,,-..--. .I ., "',1111 -.- _i. ., ,,',-...- ..! ..,.l-A .1,,',.., Z"', __k.ll. ',.,,. ..J.'i,_-.-,-.' i"I,", ,,!.I..'t..,..': 1:,,,",.,,.-2,-,A,,-,".I..-.1" ."..-I---,I-_.'4..-.",,"',,.,..,..I-1"4.,,.i1-`,-,..",..-.,",,, %, ,..T.1;, ., ..":,I.1;,i.--,,.I.i,..., ;,_k,. .,,.. .. ;I f, .o -",, "I..;. ... ,# r.... ",_.: -.- .f ._..'3i;!_--,,.VW*..- ;,".,*_ T`.1_1'11 ,4"-_-J. .,?.J...`.`-.-.`,,,,",,i- - ... .._,.,.-....I_.-:--:01" -.-.,":'.., illtl..111.'.`_..:.,._,;.,,4..,i,! lf, tit,n .F_,"..I,1:-.. -`j. :.-,i,--.,_-,-,-,."', 11.1-1,.1"7, "I .',,,,-.L-,,, .,cN-. v, .-4 "' ., '__, ,_v,.,,.. 4.11111' -, r",`-, .,.:-...,,-,",k.-;...1, "' - - " ., _ llf., .l,I,,..-4,.t,.,,..., -.j,. ,, .,.,.,-,,." .,,4, ,":. --.1,I'lliI.,..-, " .,_ i.. ,,,, '" ON. I.-li,,11 ,:. ''. 'y ., ,-F,,.-0%_.".I-I.",,,.."I-,-,-ILA,,,m,gN "I-,.l -..-".I.,R.,.t,.,;-.-,-:;. - ,_,,-,5"'.3," ,,-:, ",.,'..,..1114 1..-":C""' JPT..":.",."j,.k,-," _ .,,,,-,".,),-1, ,,.,..-..,._i.t .."1'.. -, .1. .-. ,.'t ";II;, .!. .,,.-" -- .,; -.. 1. -:.;!-,_."'I_-,-1, . ,,, ,%;..w.,4.-..,I....,I".,I.."."_4.I.4,. Ell_. ., :,7', ..-,:'' _-.. 4'_,t,.- -t- -- O-_14,'.-'.A., 1--.'' .4. .",14 A ._-.-..".1,,,..,.,-..",-..,.-'r,-,.,...,,..'-,-?.,,_ A, ,,,.A ,., i;'44A, .;!!.-,..,,-,,t.If;,,t.",.. _. ,,.""'.1".. V?4,`,.;-,t..11, --,,--,-.-.,."-I` -iI ,_,%A..,-.z,,.t_-.I"-..,.::,"..",".0 '.'..f.- ..-4'11..,.1.11,...'I'.,.'.."".,".-"'-I, I,-,-,-,.:w ,*A, ,iW..:.-` 11,,-_7i',-.i.-w-'-;,1)".. _-,.;;..,,, ;,- "6;,i" ,,!',.4, -1-it'., *,..11;'1% ,,". -, -kII"I. -,.,P,7'w'I.- _1,I.1I.o ..,;,,.,, J`-,i4 ]1-1,W.,.7,::,.-fk.S:..._ll ,, 1.,. .; ,A. ,4, .,.,. ,.. I_. .._ " ` -,-,'5 ,."'11,.-."'',-yl'.."I.%.,.-`- ., `pf'.',- ,;S"-..-:.,,;,,_.,_,-,., IIA .I."i.';9,.C"',* 1..0.4,,10- ,- .i ",-''*.io. I"II---.:. If %TA i, oe..A,w. 'Z.,, .1I,1.7;.....i,,,-'t0"... ,.lk -,,).,,,,-,,0l,.,,,._....'.1.7;4-"",71,. 'T.",It --,l,,l,-,l 1, "* ,11-1-11.11 R I...,;,j .,.,,.. k-, .'4-,,,,,--,'%,. 4c2l`.,xz",,-.,.11.,,.I,,1- 4, t-i -1171.,,-.1i..-.`.;,.A-llfl--,-..,,--,-1,1- ,.-, -., 1Y, li,_I1-.h,?..'.1.-_,; 11 U,,. .X-,.0, _x .- -1 4 i,:.'l"-,!'",..'..,I,..'r I1.-.ii.-- ,vik. '.I""60-.vI--.-..,",7 "' ,,,,,,,..,.,..".'.,.el.,...."E.., ,.,.,,-..-,.', :,,..'-l't. .,. 1, -.:...,. ,V,,,- ., ._11-.',I.",'..... .,,vl.M,.,,-.1,-., ",:' ,,,,' "" ". -.` I'.,'s,,,A, ,"x?_,"".--. .,; -,- .., _'.'O,_-,.. -,1._X, I,.j;.,...- 4- :_ "o. .. ':"-7. ," .4. ...-._.,I,i..,,.1--7-.,.--.,,.,W,,...,.,..'74",_ . ..: "' ,-'N-,:... -11,,1. .,'.',,;,.,..,.,7 .,: ,,:`..'--.r'-.N"":',-'-,.'' .i t, ..W";0i.I,,,, ,. 4.. _,. ,..','.. -'_ ....,.Zlit -,,, ,,_",-,", ,.`j,.C,;,,_I .; ,;,., _,4 ..t..,?.l"IN,,,.1,.,,.i'l._..t...,.-. " ,.";.fi.* 4..,,-,,.,, .,-,-.__ -.:.,:- _-.I,, -1--.I'll. '*'..., 4.,.,'A',,-,,.,.,:-f*-.-'-,'NI...,'.4,,-,I,i:zl.R 1_t .,,..!,,.,'. "'. 'l- F".,., -U;,. " ., "fI..__,,,:. .v"-1 W, .,4 ,,,,_," ,11 ",,.'.-,-A,4,,.*!!l.,':_.i'_ .-..11,_..,-IP1..,,,,, W.. .,,- ,, -i .. ..t7 Jw",I,-,..-F*.',,....,,:;p,.;.-, ,,,.' '.Y,'.".1 ..:.;, ,.__i:,,._'!, -. .", ,e -..,.,-_..,.,- -,,,I .-I.I.-N,,1114I.-I'iII'll``^.-., ` -,',Ckv-,A'......,..'4 ., i", 1....:,I;-,Ilg.,...,.r.I,..,.-j-...,I..4I*I-,.11i-.,.I.,:,.. ..",J.. -.' ".-",,, ., ,'.'ll .1' q-,"t.-_ ,r.%.Ii.'I, . 1.,,,V, -,.,.-,,, " C., ;Yl"_-Is -1,_,I""!iv, ,'.O,.,r."" ` _,,,.,_1:., I,1.It,...Wl,-- .- .11,.,. -1,.,'i-4s,.-,11, 1.1 -k.i".1 `.-%','A.;.-.' .'. _-- ..-...4.'_,,._-,N, -,--,".,.-!. -.-,..., ., -_,.1, -I -,f,:.,-_ ,,..I1. Z -2- ,I-,,I.q'. I.;,I_'T.. 1. .,-,,,,-;, -_.'':,".", .:,a I-,I. .N.--,.id-.2a.,-4.4.Aj"I"',"'.. 0.-*',A',,',..:.-,.N.,..,,W_,%'--',,:._';l.:,", ., ? "", ,.,. 10" A.;- .. I 7 R, X., .I,;,4,,,- .." ..:;,.,I""I'll,.I..,..liI.;:,-. A$1- ",4"t,', ., II -.;-..-.r--,,- -..,;., I"II...,1. "Z", -`,,4. -- ,-.. _-,., ,.`1Z-A1'i -_.-.,-1I---.-,!.-.,,., .,,`..-,. ,,_ ,",i,,.o.,.. -, Z,, .,'f..l""..','.,,."e!4i....,.:,I.'.,; 1( ._ .lI.i I., - -, .1-.:' -. ,I.. &,_,.v.I",I,".I1, .I,..," "',,-', ,11. .,'.,.. ,- I'll '. I-1 "_!.."';.,.,'*.,, ."'I., -.,. 1". ..,,II.11-li'l ,,.._,.-"Ii,.";.."4".".I..."v.-.".,-..,_,.'L". -".Cl..";,-,:, ."_-":,,.,. ,,,±, ., 1, ,. ,-.. ,4- *,I.I-.,..I.,.,,,-.,7. .', V , ,C,.., !"".. :',.' i.'q.-,.1-.."-, .. .'-,,;-.,k.,r,.. V.,--, 310- t,:Ak-.!. -- "":.,-.---..'_'- :4,,-I` -, ,,, - 1-11" 11 ..I ,,,, ,---. .,;,41.-..!, I.-, .!,"A.A.4 lF .-I11 M-,IV.t.., k-.1 ..,., ---,...,e. "..,*'-..`.'.-. ,-`.,` 4F..,I-..-,",, ol.II--., I, -,,,.1-..,,;*e. ,-..,. ..,,,._.t---li ..* .',.",P,,,". 4.,ft I,",.,,.1 I,-"'i-lal-I-1.-t.. wr" "I I.,-..-ot,., lvll I.: ,1: 'r, "-.'t .w'-, ,,,-F; .,.. ,.-_,.,.i,"-.".,.".,....-,: cA",..,--.,--;-'-;...'., ,, #...,"..1."..,- _.1 11*I..'kN.I."I,-,,I,,.i. ".11,Ii.U.,r,-,."771%kA.1-l"_N.j", _,.,.,.:_.',,";Q.".2,..-'' -A, ,._., .I1, I,..., .,J, .,_.;k2l _I.4 .. .,! ";,..__ .. ---.:., _14`_ _'_.._..OT,.-1111". V.,:.,, 1,41 -.1%, 7,-.I.., '6.".._ J--,,.,.,..1 4, .l-4.1 ..""., 1,I.n.,'i,vl.l, ,.,.l ._ -41 .-- I..,-`-`,_7""T,- ,.` .- X.F1*7,,lK.- -I;41IA.;, F, .-,.. Z"...`.,-,-.I. I_,",-, .,.-' v.-I:.--,I- ,.W,,,-_....,""Z..,-,.z...t,i....'..'.'.',..,,Pl.,.4',-'4.-.l,.i,..._lll ,7, x. I--,lllll..1-11.7111,.,-,I,'.l.". ,.tl "., .,. 'j, i.,i,.,..,," ., -, 1,-I,.,-;..IIk..,...-I.G ''-"'j.I.,.,'_..,-_71, .,.1-4"..%.-4--1'1. --Ikl- '!., ,.- .,_,',.7,,..., .: _,.-; -..- -1 1:..i., 31'r-,..--..-. II I- ,.4,..,.Zf, -AT."N' 1. 11 11 1,)0._I.;-*-_. li-1, ,.I .,,t,."", I, ..-7111-_.,.,.."Z' AA1,, '.VI,"4"'j, 4, .`;"!",g, -.,-,I,,,.1, ._` "t1 .1-:,,,,, ..'.I. .< LW,-.i0'lti-.. .;-_,.,-,_%...,..,",.:-,,'?$ .- 1.'.'.. "', T", .11. I;. -:Z.-*'.,Z, M--,.A...I.,,-,- -, - . 41_.l1,,iil t-I ...,t. -_ ,.- 1. l,I....-,., ,..-.,,--,,.,,,,,:.,I..I.I-4." -,,,-;- ,..4.',`.%'_ ". e',,.7-,..I":,,'',,,-'-.,.,I-I',---, '40-, -5" ,4.4 .r.,,,. ,,:; ,.N 7y. ".,.. -.,,'4,_`.*A,..:-..-I,t_.",a'..4 ",.,.,I:-i ._- 11 . ., .,.Ttz AM-..m P'll., '". ".;I.,I-I", ?W .."_,.:..-'. ". -`,'N", .", ,'.'!,., ,' ,-",.*- " -,.,.F,,..11- ..;,"""',- ".,.,., .-lj.. ,., ,,I,.-1 ;i%.,'k"._'.,4 " -,. ,,.-.F .* .,-_I._,.,.11--...,.."r --,'',-. ,-,,.,,I-4,7-,,-, n"""I,,. ,I. ,.7V, `Z'-.ll,.-'4,.,-,..,.,,.. e..II._,.....,".,.,,,.,-lt.:, _P'.,,,.'_,,-. _. _-.,,'7Zf "41 ,_',,'."!.:. f,*;'.f,.q ,II '- -,,,; 1.1WA-17-1. .1, ._lk.,..,.;11,,-,,-.x--- --,.._.4 _.. -, ,, I.-1.,",__..g.,II_,., -I'-, ,,'.;-.-'-',,i;k,:,0i. e.-,.--_!...., -, I""i..-..-.-,.,f,..,.;.,..,i:.,,..",..,.,..- .,f,,I%;-01,,"...,.4"'.-Ijp.',-.,.,---1_910, j.--,;----,.,-$04-."--ATZA,.. 1_"..tlil-11-0,...Z,-,-; .,.,t. ..,-,t_,.,,I...,14.."",. .41".. 5,.,-,,,.,,,"" A",-411`,.e","I., 1..x'',.I,,,..,.,,,,,r,.:..*,,,-,.1.A-`.:;.7'-,-'.--`K'-,V.-'-.:V",."--,__.-..R$-,,..,1,-,,,.',1,...,,.-4._,.,.,.,.-.,-1,"'.',-',1._,,.,.1-"__,til,14.L A PHYLOGENY AND CLASSIFICATION OF THE HIGHER CATEGORIES OF TURTLES EUGENE S. GAFFNEY Assistant Curator, Department of Vertebrate Paleontology The American Museum of Natural History Adjunct Professor, Department of Geological Sciences