Martinez 61..88

‚ivist— st—li—n— di €—leontologi— e ƒtr—tigr—fi— volume IIH supplement ppF TIEVV tuly PHHR

„ri ve‚qi pyƒƒsv wewwevƒ p‚yw f se @i‚s„‚ieAX ƒ‰ƒ„iwe„sgƒD

fsygr‚yxyvyq‰ exh €eviyix†s‚yxwix„ƒ

I P Q Â

fsix†ixshy we‚„sxiExe†e‚‚y D vy‚ixy ‚yyu D ewere ƒiqsh D

R P R

hiƒƒevi ‰yƒsip D we‚gy €F pi‚‚i„„s D tiriƒuiv ƒryƒrexs D

S Q

„i‡ivhi wF „igvi D ‰yƒsip vsfƒiuev

‚e™eived w—r™h PPD PHHRY —™™epted w—y PHD PHHR

uey wordsX w—mm—l f—un—Y ƒystem—ti™sY fio™hronologyY p—un—l —sso™i—zioni f—unisti™he efri™—ne ˜en ™onos™iuteD —ttr—verso un interv—lE

lo tempor—le d—l €lio™ene ƒuperiore sino —l €leisto™ene wedioF ‚esem˜l—n™e sndexY i—rly €leisto™eneY iritre—Y efri™—F

e˜str—™tF „his p—per offers — prelimin—ry overview on the l—rge

fossil m—mm—ls @€rim—tesD ry—enid—eD €ro˜os™ide—D ‚hino™erotid—eD

sntrodu™tion

iquid—eD rippopot—mid—eD ƒuid—eD qir—ffid—eD —nd fovid—eA from the

i—rly €leisto™ene sediment—ry su™™ession of the h—ndiero @fui—A f—sin

„he fossiliferous —re— of fui— @IHH um south of

@h—n—kil hepressionD iritre—AF „he IWWSEIWWU iritre—nEst—li—n fossil

w—ss—w—D northern h—n—kil hepressionD iritre—A w—s

™olle™tion h—s ˜een revised —nd studiedF e few signifi™—nt rem—ins ™olE

dis™overed in IWWS during — prelimin—ry geologi™—l surE le™ted during the PHHPEPHHRfield —™tivities h—ve —lso ˜een in™ludedF

vey under the —egis of — ™oll—˜or—tive rese—r™h progr—m iighteen spe™ies of m—mm—ls h—ve ˜een identifiedD des™ri˜ed —nd illuE

str—ted in the prin™ip—l spe™imensF „he system—ti™ study —llows gener—l ˜etween the hep—rtment of winesD iritre—n winistry

™on™lusions on the ˜io™hronologi™ —nd p—leoenvironment—l signifi™—n™e for inergy —nd ‡—ter ‚esour™es —nd the hep—rtment

of the f—un—l —ssem˜l—geF „he study is ™ompleted ˜y —n —n—lysis of

of i—rth ƒ™ien™esD niversity of ploren™eF purther stuE

––qenus p—un—l ‚esem˜l—n™e sndex99 @qp‚sA with some well known

dies were ™—rried out in the —re— during sever—l field f—un—l —ssem˜l—ges of efri™— —™ross the time sp—n v—te €lio™ene ± widE

se—sons in IWWSEIWWUD —lso involving rese—r™h te—ms dle €leisto™eneF

from the x—tion—l wuseum of iritre—D the x—tur—l risE

‚i—ssuntoF †iene present—to uno studio prelimin—re dei gr—ndi tory wuseum @qeology —nd €—l—eontology se™tionA of

m—mmiferi @€rim—tesD ry—enid—eD ‚hino™erotid—eD iquid—eD rippopoE

the niversity of ploren™eD —s well —s other st—li—n —nd

t—mid—eD ƒuid—eD qir—ffid—eD e fovid—eA provenienti d—ll— su™™essione

iurope—n institutionsF sediment—ri— del €leisto™ene snferiore del f—™ino del h—ndiero @fui—D

„hese —™tivities led to the dis™overy of — new imE hepressione h—n™—l—D iritre—AF v— m—ggior p—rte dello studio si ˜—s—

port—nt p—l—eo—nthropologi™—l site —t fui— @e˜˜—te et —lF sull— ™ollezione delle r—™™olte effettu—te dur—nte le ™—mp—gne IWWSE

IWWVAD the re™overy of —n —˜und—nt fossil verte˜r—te ™olE IWWUD i ™ui m—teri—li sono st—ti rest—ur—tiD prep—r—ti ed —n—lizz—ti nei

lo™—li del nuovo l—˜or—torio geoEp—leontologi™o —llestito presso il ––x—E le™tion —nd the identifi™—tion of — huge num˜er of —rE

tion—l wuseum of iritre—99 di esm—r—F ƒono st—ti in™lusi nello studio

™h—eologi™—l sites with extr—ordin—rily —˜und—nt —nd

—n™he —l™uni resti di p—rti™ol—re signifi™—to r—™™olti dur—nte le ™—mp—gne

m—gnifi™ently preserved wode I @yldow—nA —nd wode

PHHPEPHHRF ve IV spe™ie di m—mmiferi identifi™—te sono des™ritte ed

P @e™heule—nA tool industries @w—rtini et —lF PHHRAF

illustr—te per i reperti piuÁ signifi™—tiviF v9—n—lisi t—ssonomi™— eÁ —ffi—n™—t—

sn IWWVE IWWW field rese—r™h w—s interrupted due d— ™onsider—zioni rigu—rd—nti l— ˜io™ronologi— ed il p—leo—m˜ienteF vo

studio eÁ ™omplet—to d— un— —n—lisi sugli indi™i di simil—rit—Á @qp‚sA tr— to the renewed ithiopi—nEiritre—n ™onfli™tD ˜ut in e—rly

Á Á re— de €rehistori— ± nit—t esso™i—d— —l gƒsg E niversit—d ‚ovir— i †irgiliD €l—™Ë— smperi—l „—rr—™o ID I sg‚ieD qrup d9eutoe™ologi— rum—n—D e

RQHHS E„—rr—gon—D ƒp—inF

Á P hip—rtimento di ƒ™ienze dell— „err— e wuseo di ƒtori— x—tur—le @ƒezione qeologi— e €—leontologi—AD niversit— di pirenzeD †i— v— €ir— RD sESHIPI

ploren™eD st—lyF gorresponding —uthorX lrookdgeoFunifiFit

Q x—tion—l wuseum of iritre—D niversity of esm—r—D €FyF fox SPVRD esm—r—D iritre—F

R hep—rtment of fiologyD niversity of esm—r—D €FyF fox IPPHD esm—r—D iritre— S hep—rtment of winesD €FyF fox PIPD esm—r—D iritre—

TP fF w—rtõnezEx—v—rroD vF ‚ook et —lF Â

@ƒmithD IVQVAD †—r—nus niloti™us @vinn—eusD IUTTAD —nd PHHH the ™oll—˜or—tive st—loEiritre—n rese—r™h proje™t

w—s revived upon —pprov—l from the niversity of esE €ython ex grF €F se˜—e @qmelinD IUVWAF

m—r— ‚ese—r™h gommittee @‚ook et —lF PHHPAF

e˜˜revi—tions

„he first step of this revived proje™t w—s the esE

snstitutions

t—˜lishment of — l—˜or—tory for keeping fossil ™olle™E

hƒ„pX i—rth ƒ™ien™es heptD niversity of ploren™eD st—lyY

tions —nd for fossil ™le—ning —nd prep—r—tion —t x—tion—l

hwiX hep—rtment of winesD iritre—n winistry for inergy —nd ‡—ter

wuseum of iritre—F ell the fossil m—teri—l ™olle™ted durE ‚esour™esD iritre—Y xwiX x—tion—l wuseum of iritre—Y wq€pX x—tE

ing the IWWSEIWWU se—sons w—s in f—™t tempor—ry stored ur—l ristory wuseum @qeology —nd €—l—eontology se™tionAD niversity

of ploren™eD st—lyF —t the hep—rtment of wines in esm—r—D w—iting the

golle™tionsF eX h—ndiero f—sinD ‡—di e—l—d ™olle™tion —re—

™re—tion of proper sp—™e in the x—tion—l wuseum veE

@north of ‡—di h—ndieroAD x—tion—l wuseum of iritre—Y hexX h—nE

nuesF i—rly in PHHID the new geoEp—leontologi™—l l—˜or—E

diero f—sinD hioli ™olle™tion —re— @south of ‡—di h—ndieroD north of

tory w—s ™ompletedF „he fossil m—teri—l w—s tr—nsferred

‡—di w—e˜eleAD x—tion—l wuseum of iritre—Y wrfX h—ndiero f—sinD

there —nd the long —nd deli™—te ™le—ningGrestor—tion ‡—di w—e˜ele ™olle™tion —re— @south of ‡—di w—e˜eleD ‡—di qhers—E

work ˜eg—nF „his —llowed us to further study —nd deE loit—AD x—tion—l wuseum of iritre—Y i‚X i—st ‚udolfD x—tion—l wuE

seum of ueny— ™olle™tionsY ‡„X ‡est „urk—n—D x—tion—l wuseum of s™ri˜e in some det—il the f—un—l —ssem˜l—ge of fui—D

ueny— ™olle™tionsY ywX ysteology hep—rtmentD x—tion—l wuseum of whose f—un—l list w—s prelimin—rily pu˜lished in e˜˜—te

ueny— ™olle™tionsF et —lF @IWWVAF

we—surementsF vX lengthY ‡X widthY ‡evX width —t the —nteE

rior lo˜eY ‡€vX width —t the posterior lo˜eY „rigX trigonidY o™ —pX

„he fossils ™—me from sediments ˜elonging to the

o™™lus—l —nteroEposterior lengthY o™ trX o™™lus—l tr—nsvers—l lengthY

h—ndiero qroup @formerly ––upper h—n—kil porm—E pt™X proto™one lengthY €sX €roto™on—l sndexY l pfX postfossette lengthY

tion99Y ƒ—gri et —lF IWWVY e˜˜—te et —lF PHHRAF wost of psX €ostflexidi™ sndexY —p extX length @—nteroEposteriorA on the extern—l

sideY —p intX length @—nteroEposteriorA on the intern—l sideY protX width —t the ™olle™ting sites —re ™on™entr—ted in the —re— from

proto™oneY metX width —t met—™oneY ivX extern—l lengthY svX intern—l

‡—di e—l—d @to the northAD —™ross the h—ndiero river

lengthY €„hX proxim—l tr—nsverse di—meterY €e€hX proxim—l —nteroE

to ‡—di w—e˜ele —nd ‡—di qhers—loit— @to the southAF

posterior di—meterY w„hX tr—nsverse di—meter —t the di—phisisY we€hX

snterpret—tions of the f—un—l —ssem˜l—ge —nd the m—gneE

—nteroEposterior di—meter —t the di—phisisY h„hX dist—l tr—nsverse di—E

tostr—tigr—phi™ d—t— —llowed e˜˜—te et —lF @IWWVA to refer meterY he€hX dist—l —nteroEposterior di—meterF ell me—surements —re

the romoE˜e—ring sediments to the top of the t—r—millo given in millimetersF

p—leom—gneti™ event @su˜Eghron gIrFInAD —˜out one

million ye—rs fF€F @for further d—t— on ™hronology see

—lso el˜i—nelli 8 x—poleone PHHRY fig—zzi et —lF PHHRAF ƒystem—ti™ €—leontology

„his ™ontri˜ution represents the first study on the

yrder €rim—tes vinn—eusD IUSV whole l—rge m—mm—l —sso™i—tion from the h—ndiero

ƒu˜order g—t—rrhinii @ƒimpsonD IWRSA @fui—A f—sin @the IWWSEIWWU ™olle™tionD integr—ted with

p—mily rominid—e qr—yD IVPS notes on some of the PHHQEPHHRfindingsAF por system—E

ti™s —˜ove spe™ies level we follow w™uenn— 8 fell qenus romo vinn—eusD IUSV

@IWWVAF „he only m—mm—li—n group th—t h—s ˜een thus

romo ––ere™tusElike99 f—r studied in gre—ter det—il is the pro˜os™ide—ns @perE

retti et —lF PHHQAF elsoD — des™ription of the fossil reptiles yne mostly ™omplete ™r—nium @ eEQIAD one left

P

from the h—ndiero f—sin is given ˜y helfino et —lF —nd one left s @ eEPPP —nd eEQTWD respe™tivelyAD cs I

@PHHRAF —nd one p—rti—l right hipE˜one @ eEIUQ C eERHSAD —nd

„he prelimin—ry verte˜r—te f—un—l list pu˜lished — left pu˜i™ symphysis @ eERTTAD h—ve ˜een re™overed

˜y e˜˜—te et —lF @IWWVA is upd—ted —s followsX from the so ™—lled romo site in the ‡—di e—l—d —re—

romo ––ere™tusElike99D „heropithe™us ™fF „F osw—ldi @ erƒAF €relimin—ry morphologi™—l —nd size des™ripE

@endrewsD IWITAD ™fF gro™ut— ™ro™ut— @irxle˜enD IUUUAD tion of the e QI ™r—nium —nd —ssessment of its 4ere™E

ileph—s re™ki hietri™hD IWITD ger—totherium simum

tusElike4 st—tus h—ve ˜een ˜—sed on the —n—lysis of the

@fur™hellD IVIUAD iquus ™fF iF grevyi yust—letD IVVPD

origin—l unrestored @e˜˜—te et —lF IWWVY ‚ook et —l IWWWA

rippopot—mus gorgops hietri™hD IWPTD rex—protodon

—nd of the p—rti—lly restored spe™imen @w—™™hi—relli et

spFD uolpo™hoerus olduv—iensis @ve—keyD IWRPAD uolpoE

—lF PHHPD PHHRAF f—sed on the ™urrently —v—il—˜le eviE

™hoerus m—jus @ropwoodD IWQRAD wetridio™hoerus —ffF

den™eD — revisionD refinementD —nd integr—tion of our

wF modestus @†—n roepen —nd †—n roepenD IWQPAD qirE

previous —n—lyti™—l work is in progressF

—ff— ™fF qF jum—e ve—keyD IWTSD „r—gel—phus ™fF „F spekei

€FvF ƒ™l—terD IVTQD €elorovis oldow—yensis ‚e™kD IWPVD

p—mily ger™opithe™id—e qr—yD IVPI

uo˜us ™fF uF ellipsiprymnus @ygil˜yD IVQQAD rippotr—gus

qenus „heropithe™us qeoffroy ƒ—intErill—ireD IVRQ gig—s ve—keyD IWTSD q—zell— spFD g—prini indetFD gro™oE

dylus niloti™us v—urentiD IUTVD €elusios ™fF €F sinu—tus „heropithe™us ™fF „F osw—ldi @endrewsD IWITA

possil v—rge w—mm—ls from fui— TQ

e l—rge ™er™opithe™id ™r—nium fr—gment @ eERTQA

w—s ™olle™ted in xovem˜er PHHRin the —re— of ‡—di

e—l—dD —t — site identified —s uHHQF st preserves the fronE

t—l —nd the —nterior region of the p—riet—ls —ndD in —

prelimin—ry ™omp—risonD m—t™hes the —n—tomy of „herE

opithe™us osw—ldiD the l—rgest v—te €lio™eneD i—rly €leisE

to™ene —nd lowermost widdle €leisto™ene efri™—n ™erE

™opithe™idF „his spe™ies is found —t — num˜er of iurE

—si—n lo™—lities in™ludingX the i—rly €leisto™ene site of

– ˜eidiy—D ssr—el @felm—ker PHHPAY wirz—purD sndi—

@helson IWWQAD guev— †i™tori—D ƒp—in @qi˜ert et —lF

IWWSA —nd €irro xordD st—ly @‚ook et —lF in pressAF

yrder g—rnivor— fowdi™hD IVPI

ƒu˜order pissiped— ƒimpsonD IWRS

p—mily ry—enid—e @qr—yD IVTWA

qenus ™fF gro™ut— u—upD IVPV

™fF gro™ut— ™ro™ut— @irxle˜enD IUUUA

xo skelet—l rem—ins were found —s™ri˜ed to —

pigF I E ™fF gro™ut— ™ro™ut—F eEIIRD goproliteF f—r s™—le in ™enE

l—rge hy—enid nor other ™—rnivoresD ˜ut three ™oprolites

timetresF

of — l—rge hyen— h—ve ˜een identified in the fossil ™olle™E

tion from fui—F „he dimensions @in millimetresA of these

ment the o™™urren™e —t fui— of — very l—rge formD ™h—rE fossil ex™rements —reX eEIIR@pigF IAX STFH x PUFI x

—™terized ˜y extremely hypsodont mol—rs —nd thinD

PQFTY eEIWHX PQFQ x QTFH x QSFSY eEQWPX RUFI x RQFH

highly folded en—mel @pigF PAF „he mol—r fe—tures ™loE x QTFHF „he size —nd morphology of these ™oprolites

sely ™omp—re to those des™ri˜ed ˜y feden @IWVUA for the m—t™hes those of the l—rge efri™—n hy—enid gro™ut— ™roE

su˜spe™ies iF re™ki ileretensis @™omp—r—˜le to iF re™ki

™ut—F e se™ond ––indire™t99 eviden™e of the o™™urren™e of

st—ge Q of w—glio IWUQA from uoo˜i por— —nd sleretD this l—rge pred—tor is found on — uolpo™hoerus ™r—nium

ueny—F xeverthelessD spe™imens showing more derived @ eEPHAF „he l—tter h—s gn—w m—rks on the n—s—ls —nd

fe—turesD simil—r to th—t of iF rF re™ki @st—ge Rof w—glio the p—l—teD whi™h ˜y their size —re suggestive of gn—wing

IWUQA from feds sss —nd s† —t ylduv—iD —lso o™™ur withE ˜y gro™ut— ™ro™ut—F

in the s—mpleF ‚—ther th—n representing two different gro™ut— ™ro™ut— is the l—rgest hyen— re™orded in

t—x—D we ˜elieve the fui— s—mple to represents —n iF e—stern efri™— —fter the extin™tion in the —re— of €—™hyE

re™ki popul—tion whose evolution—ry st—ge is intermediE ™ro™ut— —round PFS w— @‡erdelin IWWWAD —lthough ™fF

—te ˜etween iF rF ileretensis —nd iFrF re™kiF @perretti et —lF €—™hy™ro™ut— ˜revirostris is ™ited —t the site of h—k—D

PHHQAF „his interpret—tion is ™onsistent with the l—te ithiopi—D —round IFH w— @esf—w et —lF PHHPAF

i—rly €leisto™ene —ge inferred from the m—gnetostr—tiE

yrder €ro˜os™ide— slligerD IVII gr—phi™ —n—lysis of the fui— su™™essionF ileph—s re™ki is

p—mily ileph—ntid—e qr—yD IVPI very ™ommon in the €lioE€leisto™ene of i—st efri™—F

qenus ileph—s vinn—eusD IUSV „he spe™ies is ˜est known from deposits in ithiopi—D

ueny— —nd „—nz—ni— @w—glio IWUQY feden IWVUY u—l˜

ileph—s re™ki hietri™hD IWIT

8 we˜r—te IWWRY „odd IWWUAD ˜ut h—s ˜een —lso reported

w—teri—l from hji˜outi @gh—v—illon et —lF IWWHAD gh—dD —nd

ƒee perretti et —lF @PHHQAY no new eleph—nt m—teri—l w—s ™olle™ted

g—nd— @w—glio IWUQAF „he iritre—n finding to d—te

during the PHHPEPHHRfield workF

represents the northernmost o™™urren™e of iF re™kiF

„he mol—rs of the fui— iF re™ki show spe™ifi™ —d—pt—tion

his™ussion

to gr—zingD whi™h suggests this eleph—nt preferred open

„he fui— eleph—nts were des™ri˜ed in — previous

environmentsF

work @perretti et —lF PHHQAF xew field surveys in PHHPE

PHHRdid not ™ontr—di™t the ™on™lusion th—t only one

yrder €erissod—™tyl— ywenD IVRV

eleph—nt spe™iesD n—mely —n —dv—n™ed form of ileph—s

p—mily ‚hino™erotid—e qr—yD IVPI

re™kiD is represented in the h—ndiero f—sinF „he spe™iE

qenus ger—totherium qr—yD IVTU

mens ™olle™ted —t sever—l sites in the —re—s of hioli @—™E

ger—totherium simum @fur™hellD IVIUA ronym hexA —nd ‡—di e—l—d @—™ronym eA do™uE

TR fF w—rtõnezEx—v—rroD vF ‚ook et —lF Â

w—teri—l

Q Q R Q Ew —nd left € Ew yne mostly ™omplete ™r—nium with right €

@hexEISHAD Q in™omplete lower mol—rs @hexEVID hexEIVHD hexE

PIRAD I —tl—s fr—gment @ eEQPVAD I ™ervi™—l verte˜r— @ eEPWWAD P huE

merus dist—l end fr—gments @ eETQD eEIPTAD I ti˜i— dist—l end @hexE

RWAD I t—lus fr—gment @ eERUAD I ™—l™—neum @ eERSVAD I dist—l wtER

@hexEIUHAF

hes™ription

et the momentD only one spe™ies of rhino™eros is

re™orded in the fui— sitesD the white rhino ger—totherE

ium simumD whi™h is entirely gr—zer @poster IWTUY

qroves IWUPAF st is represented ˜y — mostly ™omplete

™r—nium @hexEISHA @pigF QA —nd few poorly preserved

teeth —nd post™r—ni—l elementsF ger—totherium simum is

the only spe™ies of white rhino present in efri™— —fter Q

w—F st evolved from the i—rly €lio™ene —n™estor gF

pr—e™oxF „he spe™ies gF simum is represented ˜y two

su™™essive su˜spe™iesD gF simum germ—no—fri™—num

from Q to P w—D —nd gF simum simum from P w— until

presentF „his line—ge is ™h—r—™terised ˜y — progressive

de™re—se in size @querin IWVSAF

„he ™r—nium hexEISH @pigF QA preserves — ™omE

plete neuro™r—niumD p—rt of the muzzle @espe™i—lly the

Q Q

right sideAD —nd most of the p—l—te with right € Ew —nd

R Q left € Ew F e™™ording to the tooth we—r ™l—sses proE

posed ˜y rillm—nEƒmith et —lF @IWVTAD hexEISH repreE

sent — juvenile individu—l refer—˜le to the we—r ™l—ss ––ˆ99D

™orresponding to —n individu—l —ge of VEII ye—rsF „he

™r—nium is doli™ho™r—ni—lD with the o™™ipit—l ™rest proE

je™ting posteriorw—rdF „he ™heek teeth —re hypsodont

with mu™h ™ement on the ™rownD —nd protoloph —nd

met—loph —re ™urved posteriorw—rdF gomp—rison of

the fui— spe™imen with gF simum fossil ™r—ni—l rem—ins

from i—st „urk—n— @i‚ QPVgD i‚ QPWD i‚ PQPHA —nd

with one re™ent spe™imen @ywEPIVRAD shows th—t the

iritre—n spe™imen metri™s f—ll within the norm—l r—nge

Q pigF P E ileph—s re™ki @—dv—n™ed formAF eEfX hexEPRQD right w in

of v—ri—˜ility of ger—totherium simumD thoughD in geE l—ter—l @eD o™™lus—l surf—™e —t ˜ottomA —nd o™™lus—l @fD

ner—lD the v—lues —re ne—r the upper end of the r—nge —nterior to the leftA viewsY gX eERID right w in o™™lus—l Q

view @—nterior to the rightAF f—r s™—le represents IH ™mF @„—˜le IAF DAN-150 ER-328C ER-329 ER-2320 OM-2184 „—˜F I E we—surements of the fui—

Maximum length *740,0 800,0 -635,0 -500,0 825,0

Minimum length from nasal boss to nuchal crest *610,0 733,0 633,0 756,0 ger—totherium simum

Length from nasal boss to external auditory meatus *530,0 652,0 540,0 459,0 hexEISHD ™omp—red to

Basilar length *570,0 752,0 -635,0 713,0 uoo˜i por— —nd ext—nt speE

Length from palatonarial border to occipital condyles 365,0 431,0 400,0 -308,0 427,0 ™imens @d—t— from r—rris Width at nasal boss 193,0 182,0 161,0 82,0 194,0 IWVQ—AF Maximum width of cranium at orbits 328,0 302,0 -240,0 166,0 180,0 Minimum width of cranial vault at temporal fossa 127,0 120,0 115,0 103,0 112,0 Width of dorsal edge nuchal crest 320,0 275,0 -208,0 216,0 Maximum width of the occiput 263,0 283,0 216,0 Width of occipital condyles 165,0 151,0 -158,0 154,0 Depth foramen magnum to nuchal crest 169,0 156,0 -98,0 162,0 Maximum width cranium at zygoma 390,0 373,0 -320,0 -286,0 335,0 Width of the palate at M1 98,0 65,0 106,0 69,0 100,0 Width of the palate at M3 118,0 82,0 113,0 * until P3

possil v—rge w—mm—ls from fui— TS

pigF Q E ger—totherium simumF hexEISHD ™r—nium in l—ter—l @eAD dors—l @fA —nd ventr—l @gA viewsF f—r s™—le represents IH ™entimetersF

TT fF w—rtõnezEx—v—rroD vF ‚ook et —lF Â

„he teeth @„—˜le PA —nd the s™—nty post™r—ni—l elements

f—ll within the norm—l v—ri—˜ility of ger—totherium siE

mumF

p—mily iquid—e qr—yD IVPI

qenus iquus vinn—eusD IUSV

iquus ™fF iF grevyi yust—letD IVVP

w—teri—l

R I @ eEQHIAD I w @hexEPQHAD I right m—ndi˜ul—r fr—gE yne €

@hexEIRHAD P w @hexEISQD eEIQAD P € @ eEPSRD ment with € R P P

hexEIVSAD I ™ervi™—l verte˜r— @ eERSSAD I lum˜—r verte˜r— @ eEIIWAD

I humerus dist—l end @hexEIPRAD I ti˜i— dist—l end @ eEWTAD I right

m—gnum @ eEIRAD I pisiform @ eERSSAD I t—lus @ eEPUUAD I medi—l

ph—l—nx @ eEQQQAF

hes™ription

elthough s™—ntyD the study of the equid m—teri—l

in the ™olle™tion —llow us to ex™lude the o™™urren™e of

the genus ripp—rion @the genus w—s reported erroE

neously in the IWWV f—un—l listA —nd to —ttri˜ute —ll the

equid rem—ins to the genus iquus @pigF RAF „hey pro˜E

—˜ly ™orrespond to — spe™ies simil—r to the l—rge ze˜r—

inh—˜it—nt of the semideserti™ —re—sD iquus grevyiF „he

fui— dent—l rem—ins in™lude l—rge mol—rs with moder—te

proto™on—l indi™es in the upper seriesD —nd show stenoE

nine morphology in the lower oneF ynly IS spe™imens

h—ve ˜een —s™ri˜ed to this form of equidD in™luding two

m—xill—ry —nd five m—ndi˜ul—r ™heek teethD —lthough

oneD the lower w hexEISQD is not me—sur—˜le @„—˜le

P pigF RE iquus ™fF iF grevyiF eX eEIRD right m—gnum in proxim—l

QAF sn the upper seriesD —s in the uoo˜i por— spe™imens

in o™™lus—l @fAD ˜u™™—l @gA —nd viewY fEgX eEIQD right w P

R lingu—l @hA viewsF f—r s™—le in ™entimetresF —s™ri˜ed to iquus ™fF iF grevyi @iisenm—nn IWVQAD the €

eEQHI shows — developed ™—˜—lline foldD —nd the proE

to™on—l indi™es @€sA is shorter in this spe™imen th—n in

I the w hexEPQHF @QSFH —nd RTFID respe™tivelyAD suggestE in the lower series it h—s sm—ller l—˜ioElingu—l di—meters

ing —n in™re—se in the €s from the premol—rs to the —nd sm—ller v—lues in the postflexidi™ indi™es @psAF „his

mol—rsF „he lower ™heek teeth show deep lingu—l groove —nd differen™e in the ps is prominent on the lower € R

in the premol—rs —nd very deep l—˜i—l groove in the only D where the m—ximum v—lues for – ˜eidiy— —re RVFH w P

one mol—r of the ™olle™tion @w eEIQD pigFRAF „heir —nd QRFTD respe™tivelyD —nd for the spe™imens from fui— P

me—surements f—ll in the v—ri—˜ility of those of iF ™fF iF —re TPFV —nd TSFRD respe™tively @„—˜le QAF „hese morphoE

grevyi from uoo˜i por—F metri™—l d—t— suggest th—t the fui— spe™imens —re not

„hese spe™imens h—ve —lso ˜een ™omp—red to refer—˜le to iquus t—˜etiD —nd their —n—tomy —nd me—E

iquus t—˜eti from uoo˜i por— @iisenm—nn IWVQA —nd surements —re ™lose to iquus grevyiF

– ˜eidiy—D ssr—el @iisenm—nn IWVTAF sn gener—lD the teeth

yrder ertiod—™tyl— ywenD IVRV —n—tomy of iF t—˜eti shows some simil—rities to the fui—

ƒu˜order ƒuin— qr—yD IVPI spe™imensD espe™i—lly in the upper ™heek teeth seriesD ˜ut

P3/ P4/ M1/ M2/ M3/ L ap ext L ap int W prot W met L ap ext L ap int W prot W met L ap ext L ap int W prot W met L ap ext L ap int W prot W met L ap ext L ap int W tr DAN-150 Rt 48,8 38,9 47,3 47,1 49,6 40,2 48,6 46,8 54,8 47,8 56,7 54,4 63,9 47,7 50,0 46,1 56,1 55,9 46,4 DAN-150 Lt 50,5 38,0 45,3 48,8 57,8 48,3 58,9 53,4 64,0 48,4 55,4 40,6 51,3 50,3 46,4 ER-1190 49,4 46,4 51,5 48,8 ER-328 54,5 41,1 63,6 56,5 50,0 39,8 70,0 64,5 64,0 56,0 77,2 64,8 64,4 61,3 59,8 ER-1189 46,1 43,4 59,2 55,3 77,4 69,4 54,7 ER-329 71,0 48,2 57,1 33,8 ER-1191 58,4 54,8 67,6 54,9 ER-659 68,2 65,6 61,2 OM-2184 46,0 46,4 51,9 47,0 55,2 51,2 51,2 44,6 50,5 52,5 58,0 55,1 73,2 60,2 51,6 45,7

OM-2216 38,0 30,4 34,7 31,0 44,6 39,0 39,0 35,8 46,1 37,9 42,6 36,2 46,4 43,8 51,0 40,5 58,3 46,3 42,3

„—˜F P E we—surements of the ger—totherium simum teeth @hexEISHAD ™omp—red uoo˜i por— —nd ext—nt spe™imens @d—t— from r—rrisD IWVQ—AF

possil v—rge w—mm—ls from fui— TU

upper teeth UA-301 DAN-230 P4/ M1/ oc ap 30,0 27,1 oc tr 30,8 31,4 ptc 10,5 12,5 PI 35,0 46,1 lower teeth UA-185 UA-254 DAN-140 UA-13 P/2 P/2 P/4 M/2 oc ap 34,9 33,9 31,2 28,3 oc tr 20,0 16,8 19,6 18,5 l pf 14,8 17,3 14,9 10,5

FI 42,4 49,6 62,8 65,4

„—˜F Q E we—surements of the upper —nd lower ™heek teeth of

iquus ™fF iF grevyi from fui— pigF S E rippopot—mus gorgopsF eEPPQD right m—xill—ry fr—gment

I P Q with w @deeply wornA w —nd w D in o™™lus—l viewF ƒ™—le ˜—r in ™entimetresF

47,0 45,0 Turkana basin 45,0 Buia basin 40,0 Ubeidiya 43,0 H. amphibius 35,0 41,0 30,0 39,0 M1/ anterior width M/1 anterior width 25,0 37,0

35,0 20,0 30,0 35,0 40,0 45,0 50,0 55,0 60,0 30,0 40,0 50,0 60,0 70,0 M1/ length M/1 length

65,0 55,0

60,0 50,0

55,0 45,0

50,0 40,0 M/2 anterior width M2/ anterior width 45,0 35,0

40,0 30,0 40,0 45,0 50,0 55,0 60,0 65,0 70,0 40,0 45,0 50,0 55,0 60,0 65,0 70,0 M2/ length M/2 length

70,0 55,0

65,0 50,0

60,0 45,0

55,0 40,0 50,0 M/3 anterior width M3/ anterior width 35,0 45,0

40,0 30,0 40,0 45,0 50,0 55,0 60,0 65,0 70,0 75,0 60,0 65,0 70,0 75,0 80,0 85,0 90,0 95,0

M3/ length M/3 length

pigF T E fiv—ri—te plots of rippopot—mus upper @left ™olumnA —nd lower @right ™olumnA mol—rs dimensions @see „—˜les R—nd S for d—t— sour™eAF

TV fF w—rtõnezEx—v—rroD vF ‚ook et —lF Â

p—mily rippopot—mid—e qr—yD IVPI h—s ˜een —s™ri˜ed to rippopot—mus gorgopsF „he ™olle™E

qenus rippopot—mus vinn—eusD IUSV tion is represented ˜y dent—l —nd post™r—ni—l rem—ins

˜ut unfortun—tely no ™omplete skull h—s ˜een do™umenE

rippopot—mus gorgops hietri™hD IWPT

ted to d—teF

@—dv—n™ed formA

hentition ± rippopot—mus gorgops from fui—

shows ro˜ust —nd hypsodont teeth with thi™k en—melD w—teri—l

pour upper ™—nines @hexEIISD hexEITPD eEPPD eEITQAD Q —nd ro˜ust in™isors —nd ™—ninesF ynly one upper preE

I P @hexESHD hexEPHHD eESEUAD P s @hexESHD hexEITIAD I p—l—te s

mol—r is present in the ™olle™tion @hexEIISAF „he

I Q P Q with right w Ew —nd left w Ew @ eEPPQAD I left p—l—te fr—gment with

upper mol—rs —re tetr—™uspidD —nd show — l—rge ™inguE

P Q I Q Q Ew @ eESRAD I left p—l—te with w Ew @hexEUVAD I € @hexEIISAD w

lum on the mesi—l —nd dist—l sidesD whi™h is —lso present

I P Q Iw @hexEPIWAD P w @hexEWID hexEPIVAD Q w @hexEPPPD hexE

in the lingu—l —nd l—˜i—l f—™esF „he in™re—se of size from Eg @ eEWPD eEIVRAD Q IWQD hexESQAD P m—ndi˜ul—r fr—gments with s I

I P Q

s @hexEPHD eEPRD eEPTQAD R s @hexEIITD hexEIIUD eEWPD I P to w to w is very prominent @pigsF SD TY „—˜le RAF w

eEQRSAD R lower ™—nines @hexEISD hexEISRD hexEITTD eEIAD Q

„he lower premol—rs —re uni™uspid with developed disE

Ew @hexEID hexEIID hexEIIWD m—ndi˜ul—r fr—gments with w

I Q

t—l ™ingulumF „he morphology of the lower mol—rs is

Ew @hexEIUSD hexEIPHD eEIIPAD Q m—ndi˜ul—r fr—gment with w I P

simil—r to th—t of the upper onesD ˜ut the linguoEl—˜i—l

Ew @hexEPPUD wrfETD eEIRSAD P m—ndi˜ul—r fr—gments with w P Q

wrfEUAD I m—ndi˜ul—r fr—gment with € Ew @ eERSPAD R € @hexE dimension is proportion—tely sm—llerD the lingu—l —nd Q I R

@hexEIIQD hexEISWD eE ITD hexEIUQD eEISWD eEPVPAD Q w I l—˜i—l ™ingulum is less developedD —nd the third mol—r

PQHAD Rw @hexESRD hexEIWPD hexEPRVD eEIHHAD P r—diusEuln—

Q

A h—s — l—rge t—lonid @„—˜le SAF @w Q fr—gments @ eEVSD eEVTD eEVUAD S t—lus @hexERVD hexEWQD eE

„he metri™—l ™omp—rison of the dent—l spe™imens

RPD eEPQPD eEQSHAD R ™—l™—nei @hexEIRTD eERHD eEISSD eE

from fui— to the good ™olle™tion of rF gorgops from PQWAD —nd m—ny other post™r—ni—l fr—gments @humerusD femursD ti˜i—sD

™—rp—lsD met—podi—ls —nd ph—l—ngesAF uoo˜i por— —nd ‡est „urk—n— @r—rris IWWI—A @pigF TAD

shows th—t in the upper mol—r seriesD the fui— spe™imens

h—ve l—rger mesioEdist—l length for the first —nd the se™E hes™ription

ond mol—r —nd — shorter third mol—rF „he me—surements e l—rge form of rippopot—musD l—rger th—n the

of the lower mol—r series from fui— —re more simil—r to ext—nt rF —mphi˜iusD h—s ˜een do™umented —t fui—

@pigF SAF st is the most —˜und—nt t—xon in the —re— —nd the rF gorgops ™olle™tion from the „urk—n— f—sinD th—n

P1/ P2/ P3/ P4/ M1/ M2/ M3/ L W L W L W L W L WAL WPL L WAL WPL L WAL WPL BUIA UA-223 R. 46,5 66,4 58,6 57,7 53,6 49,5 46,2 UA-223 L. 66,9 53,0 47,8 45,7 UA-54 R. 60,0 60,8 59,6 DAN-78 L. 46,6 58,8 60,0 61,6 57,7 60,0 58,6 DAN-222 L. 63,0 52,9 46,1 DAN-219 R. 55,1 44,4 46,2 DAN-118 L. 59,3 54,5 50,3 DAN-91 L. 56,2 52,0 51,8 DAN-193 L. 52,6 DAN-53 R. 57,7 58,4 53,7 DAN-115 L. 37,9 26,4 TURKANA BASIN WT-19634 L. (1) 16,0 17,0 32,5 25,1 37,4 28,6 39,5 35,4 49,2 44,8 46,0 59,6 57,3 55,6 68,8 62,3 52,3 ER-305 38,6 37,4 38,7 39,8 41,7 ER-1307R. 31,8 32,0 44,2 40,4 41,6 57,6 53,6 53,6 56,1 53,7 48,3 ER-1394 R 39,6 30,3 37,9 35,1 40,1 40,0 46,7 57,6 53,6 53,6 62,2 57,6 49,8 ER-1427 L 34,1 33,9 48,5 37,2 53,1 51,8 49,2 ER-12884 33,4 34,8 ER-637 42,2 45,1 47,4 57,8 59,0 57,4 55,0 54,7 ER-1306 R 48,5 45,7 46,1 59,2 57,3 49,5 ER-5518 71,2 64,2 53,7 ER-5514 58,5 54,3 UBEIDIYA H. behemoth 39,7 28,9 30,2 27,3 56,0 41,0 40,0 60,3 47,3 50,7 65,0 54,0 49,0 Living Hippopotamus

H. amphibius 35,5 27,6 28,8 32,2 42,0 39,9 41,9 49,4 47,8 46,8 48,6 46,8 41,5

„—˜F RE pper dentition of rippopot—mus gorgops from fui—D the „urk—n— f—sin @r—rrisD IWWI—AD rF ˜ehemoth from – ˜eidiy— —nd ext—nt rF

—mphi˜ius @p—ureD IWVTAF

possil v—rge w—mm—ls from fui— TW

P/1 P/2 P/3 P/4 M/1 M/2 M/3 M/1-M/3 L W L W L W L W L WAL WPL L WAL WPL L WAL WPL L BUIA DAN-1 R. 38,3 35,5 56,0 47,9 48,2 78,7 47,6 43,0 174,3 UA-112 R. 40,1 40,9 55,5 41,2 45,8 69,4 165,8 UA-43 L. 55,2 37,8 43,8 DAN-175 L. 60,3 36,0 39,1 66,8 39,2 40,5 DAN-227 L. 49,4 38,3 40,9 39,5 40,2 DAN-54 R. 43,3 42,7 DAN-192 R. 80,4 39,4 35,8 DAN-111-119-120 R. 52,7 39,4 65,6 50,5 48,2 74,5 49,1 50,1 196,0 DAN-159 L. 50,5 UA-230 R. 34,7 35,8 UA-282 L. 29,8 DAN-16 R. 41,6 29,0 DAN-248 L. 78,5 42,9 40,7 DAN-113 R. 63,3 38,9 37,0 UA-100 L. 74,0 UA-145 R. 66,1 41,5 UA-452 L. 36,7 26,9 26,0 53,7 UA-159 R. 40,0 23,2 DAN-173 R. 42,8 27,1 MHB-6 R. 46,4 53,9 43,1 45,5 MHB-7 R. 48,1 45,3 47,5 73,6 47,2 41,7

TURKANA BASIN ER-2405 32,6 22,0 ER-1427 37,6 23,3 36,8 26,9 43,4 31,7 54,1 38,1 41,4 63,0 49,7 42,2 WT-19982 41,1 23,7 42,3 27,2 42,6 33,3 49,2 38,7 40,8 63,8 47,5 45,0 82,6 47,9 44,6 ER-1421 37,5 23,8 37,2 26,9 ER-1425 R 40,3 30,0 ER-1514 41,0 32,9 ER-2405 33,5 26,6 40,1 32,0 34,6 58,2 41,8 44,2 77,6 43,7 41,7 ER-5527 44,3 33,3 ER-785 57,2 37,1 ER-1207 R 45,5 30,4 31,2 ER-1394 R 41,5 31,2 36,8 56,7 39,7 43,7 88,5 42,5 41,9 185,6 ER-1427 R 54,2 38,1 41,4 63,0 49,7 42,2 ER-2044 48,7 35,1 38,9 61,2 45,6 47,7 ER-2405 R 40,1 32,0 34,6 58,2 41,8 44,2 77,6 43,7 41,7 172,5 ER-1307 52,5 34,6 36,3 57,0 ER-5510 57,8 43,9 44,8 ER-1409 89,3 46,0 42,4 ER-5513 80,0 41,5 42,1 ER-5527 L 86,3 46,2 46,1 UBEIDIYA H. behemoth 34,0 19,5 42,5 25,2 41,0 28,5 54,0 33,0 35,0 83,0 47,0 41,0 Living Hippopotamus

H. amphibius 32,4 18,7 36,3 22,4 35,1 27,3 54,6 37,9 39,6 67,8 38,5 37,4

„—˜F S E vower dentition of rippopot—mus gorgops from fui—D the „urk—n— f—sin @r—rrisD IWWI—AD rF ˜ehemoth from – ˜eidiy— —nd ext—nt rF

—mphi˜ius @p—ureD IWVTAF

—re the upper ones from the s—me ˜—sinF „hese metri™—l qenus rex—protodon p—l™oner 8 g—utleyD IVQT

d—t— h—ve —lso ˜een ™omp—red to those of the s—mple

rex—protodon spF from – ˜eidiy—D —s™ri˜ed in p—ure @IWVTA to the spe™ies

rippopot—mus ˜ehemothD whi™h is interpreted —s —n enE

w—teri—lF yne unErestored ™r—nium @hexEPIQAD I unErestored demi™ form from the i—rly €leisto™ene of the vev—ntine

neuro™r—nium @hexEPHSAD I left t—lus @wrfESAD I right m—gnum

‚egionF „he spe™imens from fui— m—t™h the v—ri—˜ility

@hexEPRTAD —nd sever—l other teeth fr—gments —nd post™r—ni—l eleE

of this form from ssr—elF st seems —s if the s—mples from

mentsF

fui— —nd – ˜eidiy— ™orrespond to —n —dv—n™ed form of

rippopot—mus gorgopsF

UH fF w—rtõnezEx—v—rroD vF ‚ook et —lF Â

pigF U E gomp—r—tive view of rippopot—mus gorgops @eEfD iA —nd rex—protodon spF @gEhD pA post™r—ni—F eEfX hexEIHD right m—gnum in

dist—l @eA —nd proxim—l @fA viewsY gEhX hexEPRTD right m—gnum in dist—l @gA —nd proxim—l @hA viewsY iX eEPQPD left t—lus in

posterior viewY pX wrfESD left t—lus in posterior viewF ƒ™—le ˜—r in ™entimetresF

hes™ription „his is the first re™ord of this genus from the

e sm—llEsized ˜r—™hydont hippo is present in the i—rly €leisto™ene of the ef—r regionD where the genus

fui— —ssem˜l—ge together with rippopot—mus gorgopsF is reported to h—ve dis—ppe—red —fter PFS w— @foisserie

st w—s ™ited —s rex—protodon k—rumensis in e˜˜—te PHHPAF

@IWWVA ˜e™—use of its —n—tomi™—l simil—rities with this „he ™omp—rison of the post™r—ni—l elements of

spe™iesF e mostly ™omplete ™r—nium of this sm—ll hippo rippopot—mus gorgops —nd rex—protodon spF from fui—

h—s ˜een found re™ently @hexEPIQAD ˜ut it is still in the shows ™le—r fun™tion—l differen™es ˜etween the two speE

restor—tion pro™ess —nd ™—nnot ˜e in™luded in this studyF ™iesX IA in rF gorgops the —rti™ul—r f—™et for the third

„he —n—tomy of the ™r—niumD the dent—l proportions —nd met—™—rp—l on the dist—l f—™e of the m—gnumD is rel—E

the —n—tomy of the other spe™imens —s™ri˜ed to this tively shorter —nteroEposteriorlyD —nd wider tr—nsverE

spe™ies suggest — different spe™ies th—n rF k—rumensisF selyD th—n in rex—protodon spF „his me—ns th—t the —nteE

‡e h—ve to —w—it the restor—tion of this mostly ™omE roEposterior movement of the m—nus w—s wider in rexE

plete ™r—nium ˜efore we ™—n —ssign — spe™ies n—me to the —protodon th—n in rF gorgops @pigF UAY PA in rex—protoE

fui— rex—protodon spe™iesF don spF the t—lus is rel—tively slender —nd more gr—™ileD

possil v—rge w—mm—ls from fui— UI

ment with w @hexEIWWAD I left m—ndi˜ul—r fr—gment with € @hexE Q R while in rippopot—mus gorgops it is proportion—tely

@hexEPQQAD I w @hexESIAD W w @hexEIQVD hexEVPD IQPAD I €

R I Q shorter —nd more ro˜ustD ™orresponding to — very m—sE

hexEIQSD hexEIQWD hexEIVWD hexEPRWD eEIITD eEPQPD eE

sive spe™iesF „his ™h—r—™teristi™ is rem—rk—˜le in the difE

QPWAF

feren™e of the length —nd tr—nsvers—l di—meter of the

t—lus @pigF UAF

hes™ription

pper teeth ± „he ™—nine @ eEISRA h—s — gre—ter

p—mily ƒuid—e qr—yD IVPI

mesioEdist—l di—meter th—n the dorsoEventr—l oneD —nd

it h—s — longitudin—l groove in the posteroEdors—l f—™eF „hree spe™ies of suids h—ve ˜een identified —t

I

@hexEPPIA is — worn typi™—l tetr—™uspid moE „he w fui—D two pert—ining to the genus uolpo™hoerus @uF oldE

l—rD with l—˜i—l ™uspids @p—r—™one —nd met—™oneA l—rger uv—iensis —nd uF m—jusA —nd one to the genus

—nd higher th—n lingu—l ones @proto™one —nd hypoE wetridio™hoerus @wF —ffF wF modestusAF

™oneAF st —lso shows — developed ™ingulum in the meE

Q

fr—gment is present si—l —nd dist—l f—™esF ynly one w qenus uolpo™hoerus †—n roepen —nd †—n roepenD IWQP

@ eEQTAF

uolpo™hoerus is ˜elived to ˜e the —n™estor of the

vower teeth ± „wo more or less ™omplete m—ndE

ext—nt rylo™hoerusD the forest hogF st is — sexu—lly diE

i˜les @ eEPRP —nd wrfEIA @pigF VA —re present in the

morphi™ suid of moder—te to l—rge sizeF st h—s low

s—mpleF „he m—ndi˜ul—r ˜ody shows — tri—ngul—r se™E

™rowned —nd well rooted mol—rs —nd infl—ted m—ndE

tionF €reserved teeth in™lude the ™—nineD —nd the ™heek

i˜les ˜ene—th the ™heek teeth @r—rris IWVQ˜AF foth speE

Ew F „he premol—rs @€ —nd € A —re dent—l series € Q Q Q R

™ies of uolpo™hoerus re™orded in the efri™—n i—rly

su˜tri—ngul—r uni™uspid pie™esF „he first —nd se™ond

€leisto™eneD uF olduv—iensis —nd uF m—jusD —re well reE

—nd w A —re tetr—™uspid pie™es with lingu—l mol—rs @w I P

presented —t fui—F ™uspids @met—™onid —nd ento™onidA ˜igger —nd higher

nfortun—telyD the ˜est spe™imens of uolpoE th—n the l—˜i—l ones @proto™onid —nd ento™onidAF sn the

™hoerusD the —lmost ™omplete ™r—nium hexEIRP —nd

—nd w l—˜ioElingu—l di—E spe™imen eEPRP the w I P

the muzzle eEPH —re still not restored —nd ™—nnot ˜e meter is gre—ter th—n the mesioEdist—l oneF hexESI

in™luded in this studyF eEPH h—s ˜een ™ited —˜ove ˜eE exhi˜its sm—ll mesi—l —nd dist—l ™ingul—F „he lower

™—use of the hyen— gn—w m—rks found in the n—s—ls —nd

A is the ˜est preserved spe™imenD it third mol—r @w Q

the p—l—te of this spe™imenF h—s tetr—™uspid trigonid @protoED met—ED hypoE —nd enE

to™onidA norm—lly shorter th—n the t—lonidD whi™h is

uolpo™hoerus olduv—iensis @ve—keyD IWRPA very elong—ted —nd ™—n h—ve until four or five p—ir

of ™uspids @pigF W™EdAF st is worth to note the o™™urE

w—teri—l

ren™e of one import—nt —symmetry in the spe™imen

I Q @hexEPPIAD P w @ eEQTD eE yne upper g @ eEISRAD I w

is l—rger th—n the left wrfEID where the right w Q ITUAD I ™omplete m—ndi˜le with ˜oth ˜r—n™hes @wrfEIAD I m—ndi˜le

one @TIFI —nd STFI mmD respe™tivelyAD ˜e™—use it h—s with most of ™omplete left ˜r—n™h @ eEPRPAD I right m—ndi˜ul—r fr—gE

pigF V E uolpo™hoerus olduv—iensisF

eEfX eEPRPD right m—ndE

i˜le in o™™lus—l @eA —nd

˜u™™—l @fA viewsF f—r s™—le in ™entimetresF

UP fF w—rtõnezEx—v—rroD vF ‚ook et —lF Â

l—rger t—lonid with one more p—ir of ™uspidsF „his vower teeth ± xo ™—nines —re present in the ™olE

—symmetry possi˜ly is rel—ted to — high endog—my in Ew D whi™h is shown in le™tionF „he ™heek teeth h—ve € Q Q

the popul—tions of uolpo™hoerus in the —re—F the spe™imens hexEIRI —nd hexEIURF „he premol—rs

uolpo™hoerus olduv—iensis is —n —dv—n™ed form of h—ve one m—in ™usp —nd —re l—rger —nd higher th—n in uF

the type spe™ies uolpo™hoerus limnetesD whi™h in™re—sed

olduv—iensisFw —nd w —re evolution—rily —dv—n™ed I P

the size —nd length of the third mol—rs from the primiE ™omp—red to uF olduv—iensisF „hese mol—rs —re tetr—™usE

tive €ot—mo™hoerus —n—tomyF pidD l—rgerD higher ™rowned —nd proportion—lly more

elong—ted th—n in uF olduv—iensis —nd show developed

uolpo™hoerus m—jus @ropwoodD IWQRA

mesi—l —nd dist—l ™ingul—F „he third mol—r is shorter th—n

in uF olduv—iensisD h—ve developed tetr—™uspid trigonid

w—teri—l

—nd short t—lonid with few p—irs of ™uspidsF sn ™rossE

yne p—l—te fr—gment with the left g @ eEQHQAD I germin—l

P

@hexEIQQD eEPPQD eERQUAD I m—ndi˜ul—r upper g @ eERSUAD Q w se™tion the en—mel is gener—lly more ™ompli™—ted th—n

symphysis @hexEIVUA I in™omplete m—ndi˜le with left —nd right

th—t of uF olduv—iensisF

Ew ˜r—n™hes @hexEIRIAD I right hemim—ndi˜ul—r fr—gment with w

I P

uF m—jus h—s — more ™onserv—tive teeth —n—tomy

@hexEIWRAD I left hemim—ndi˜ul—r fr—gment with w @hexEIURAD I Q

th—n uF olduv—iensisD with less elong—te w D —nd proporE

Q @hexEIRQD eERRD eERTRAF m—ndi˜ul—r symphysis @ eEQUVAD Q w Q

tion—tely more developed w —nd w @„—˜le UAF I P

hes™ription

his™ussion on the genus uolpo™hoerus pper teeth ± „he ™—nines @pigF W—E˜A —re l—rger

P fesides hexEIRP —nd eEPHD the fui— uolpoE @hexEIQQ th—n in uF olduv—iensis @„—˜le TAF „wo w

™hoerus s—mple in™ludes spe™imens th—t m—y ˜e refer—˜le —nd eERQUA h—ve ˜een identifiedD whi™h —re morphoE

to —s m—ny —s two spe™iesD in™ludingX I o™™ipit—l —nd logi™—lly simil—r to uF olduv—iensisD ˜ut —re l—rger —nd

tempor—l ˜one fr—gment @ eERRIAD I left hemim—ndi˜E more hypsodont th—n in uF olduv—iensisF

pigF W E uolpo™hoerus m—jusF eEfX

eEQHQD left m—xill—ry

fr—gment with g —nd in™iE

sors —lveoli in o™™lus—l @eA

—nd ˜u™™—l @fA viewsF uolE

po™hoerus olduv—iensisF gE

in iX hexEPQPD right w Q

o™™lus—l @gAD lingu—l @hA

—nd ˜u™™—l @iA viewsF f—r s™—les in ™entimetresF

C M1/ M2/ M3/ LW LW LW LWL trigL tal K. Olduvaiensis UA-36 24,60 DAN-221 23,70 17,20 UA-154 31,00 15,60 K. Majus UA-437 34,80 15,40 DAN-133 29,50 22,00 UA-303 29,30 27,00

UA-457 21,20 22,00 24,00

„—˜F T E pper teeth of uolpo™hoerus olduv—iensis —nd uF m—jus from fui—F

possil v—rge w—mm—ls from fui— UQ

C P/2 P/3 P/4 M/1 M/2 M/3 LW LW LW LW LW LW LWL trigL tal K. Olduvaiensis UA-242 R. 23,90 17,80 10,60 7,20 14,20 12,70 10,50 15,40 16,40 17,90 67,00 19,70 25,20 41,80 DAN-138 L. 65,40 20,80 28,60 36,80 DAN-82 L. 61,30 20,00 30,10 31,20 UA-116 L. 19,90 42,10 DAN-139 L. 19,20 30,50 DAN-199 R. 57,50 17,60 28,70 28,80 DAN-132 L. 15,00 12,60 DAN-51 L. 23,20 16,10 UA-329 R. 22,50 UA-232 R. 51,10 18,00 23,80 27,30 DAN-249 L. 17,90 37,50 DAN-233 R. 13,90 10,80 MHB-1 L. 17,90 20,60 20,10 56,10 20,70 27,90 28,20 MHB-1 R. 18,60 16,11 11,50 5,30 13,30 10,90 17,40 22,60 20,60 61,10 20,90 28,50 32,60 K. Majus DAN-143 L. 45,20 22,90 28,20 17,00 UA-44 L. 21,80 25,90 DAN-174 L. 14,00 10,60 19,50 11,80 18,70 16,80 26,40 18,60 21,70 27,00 DAN-194 R. 16,70 16,90 27,20 21,10 DAN-141 L. 9,90 16,90 12,60 21,10 14,30 26,10 18,00 40,10 21,00 24,60 15,10 DAN-141 R. 22,50 13,80 26,90 19,30 42,70 20,90 26,10 16,60

UA-464 R. 21,50 12,80

„—˜F U E vower teeth of uolpo™hoerus olduv—iensis —nd uF m—jus from fui—F

ul—r fr—gment with no teeth @ eEQUAD I mol—r fr—gment @ithiopi—AD whi™h is simil—r in its —ge to fui— @esf—w

et —lF PHHPAF „he l—st represent—tives of the uolpo™hoerus @hexESPAD I —tl—s fr—gment @hexEIIRAD I humerus

line—ge in efri™— —re uF olduv—iensis from ylduv—i fed dist—l end @hexEPQTAD I femur dist—l end @ eEIVHAD I

s†D „—nz—ni— @HFUV w—A —nd uF m—jus —t fodoD widdle ti˜i— dist—l end @ eEQIPAD S t—lus @hexEIPD hexESWD

ew—shD ithiopi— @HFT w—A @‡hite IWWSAF uolpo™hoerus hexEIRWD hexEIVID eEPUVAD I proxim—l ph—l—nx

olduv—iensis extended its geogr—phi™ r—nge into the veE @hexETHAD I medi—l ph—l—nx @ eESWAF

v—ntD it is found in – ˜eidiy—D ssr—el @~IFREIFS w—A @qeE uolpo™hoerus m—jus is — more ™onserv—tive spe™ies

r——ds et —lF IWVTAD —nd in ivron u—rryD —lso in ssr—el th—n the line—ge uF limnetesEolduv—iensisD —nd preserves

@~HFVEIFH w—AD ™ited —s uF evronensis @„™hernov et —lF — primitive ™heek teeth —n—tomy ™loser to the —n™ient

IWWRAF sn xorth efri™—D this genus is well known —nd forms rel—ted to €ot—mo™hoerus @r—rris 8 ‡hite IWUWAF

™ited —s uF ph—™o™hoeroides in sever—l sitesD in™luding „hese efri™—n suids —re ˜—si™—lly ™h—r—™terized ˜y

ein r—ne™h —nd ehl —l yughl—m @qer——ds IWWQY ƒh—E the development of the zygom—ti™ —r™h —nd the dist—lE

nouni et —lF PHHPAF w—rd exp—nsion of the lower third mol—rsD with sever—l

pill—rEp—irsF „he en—mel is —lso very thi™kF „he genus

qenus wetridio™hoerus ropwoodD IWPT uolpo™hoerus h—s ˜een interpreted —s —n evolved efri™—n

form whi™h w—s derived from —n esi—n immigr—nt durE wetridio™hoerus is ˜elieved to ˜e —n™estr—l to the

ing the widdle €lio™ene @r—rris 8 ‡hite IWUWY r—rris living w—rthogD €h—™o™hoerus —ethiopi™us @r—rris 8

IWVQ˜Y €i™kford IWWRAF „he oldest re™ognized spe™ies of ‡hite IWUWAF wetridio™hoeres —re sexu—lly dimorphi™

the genus w—s uF —f—rensis from r—d—r @gooke IWUVAD pigs of sm—ll to l—rge sizeF „hey —re ™h—r—™terised ˜y

˜ut — new study of the i—rly —nd widdle €lio™ene suid the redu™tion of the premol—r series —nd the in™re—se

m—teri—l from ithiopi— —nd gh—d h—s determined the in lengthD hypsodonty —nd root fusion of the mol—rsD

o™™urren™e of — primitive spe™ies @uF deheinzeliniA espe™i—lly in the m—xill—ry —nd m—ndi˜ul—r wQsF ix™ept

whi™h g—ve origin to the uolpo™hoerus line—ge in the in the le—st progressive ex—mplesD fusion of the ™rown

efri™—n ™ontinent @frunet 8 ‡hite PHHHAF st would —pE elements of the ™heek teeth ˜egins shortly —fter we—r

pe—r to ˜e derived from the v—te wio™ene esi—n spe™ies ™ommen™es @r—rris 8 ‡hite IWUWAF

€ropot—mo™hoerus hysudri™usF uolpo™hoerus deheinzeliE

ni g—ve rise to uF ™ookeiD — sm—ll —nd hypsodont spe™ies wetridio™hoerus —ffF wF modestus @†—n roepen —nd

†—n roepenD IWQPA whi™h is only found in r—d—rD —nd to uF —f—rensisD whi™h

g—ve origin to uF m—jus —nd to uF limnetesEuF olduv—ienE

w—teri—l

sis @frunet 8 ‡hite PHHHAF uolpo™hoerus olduv—iensis

Ew @hexEPIVAD I m—ndi˜E yne m—ndi˜ul—r fr—gment with € R P

—nd uF m—jus —re ™ommon in most of the i—rly €leistoE

—nd roots of w @ eERRWAD R w @ eEQTUD ul—r fr—gment with w Q P Q

™ene efri™—n —ssem˜l—ges in™luding the site of h—k— hexEIWUD hexEPPHAF

UR fF w—rtõnezEx—v—rroD vF ‚ook et —lF Â

oldestD —nd —rrived in efri™— from esi— in the widdle

€lio™eneF st g—ve origin to two different line—gesD IA the

wF hopwoodiEwF ™omp—™tus line—geD ™h—r—™terised ˜y —

progressive in™re—se of ˜ody sizeD —nd PA to the sm—ll

sized wF modestusD whi™h is ˜elieved to ˜e — pl—usi˜le

—n™estor of €h—™o™hoerusF „he genus wetridio™hoerus is

—lso reported from north efri™—D —t the l—te i—rly €leisE

to™ene site of „ernifine in elgeri—F

ƒu˜order ‚umin—nti— ƒ™opoliD IUUU

p—mily qir—ffid—e qr—yD IVPI

qenus qir—ff— frissonD IUST

qir—ff— ™fF qF jum—e ve—keyD IWTS

w—teri—l

P Q

Ew —nd the roots e left m—xill—ry fr—gment @ eEPSIA with w

P I P of € D one very worn right w or w @hexEIWVAD two right t—lus

@hexEIU —nd hexEUHAD —nd few other fr—gment—ry spe™imensF

hes™ription

e gir—ffid is re™orded in the fui— sitesF ynly few

spe™imens —re —s™ri˜ed to qir—ff— ™fF qF jum—iD whi™h

h—s simil—r size to the ext—nt spe™ies qir—ff— ™—melop—rE

d—lisD —nd it is well known in the i—rly —nd widdle

€leisto™ene from e—stern efri™—F „he me—surements of

the mol—rs from the spe™imens eEPSI —nd hexEIWV

@„—˜le WA f—ll within the r—nge of v—ri—˜ility of qir—ff— pigF IH E wetridio™hoerus —ffF wF modestusF eEgX e RRWD left

in o™™lus—l @eAD lingu—l m—ndi˜ul—r fr—gment with w jum—e from uoo˜i por— @r—rris IWWI˜AD while the two Q

@fA —nd ˜u™™—l @gA viewsF f—r s™—le in ™entimetresF t—lus spe™imens @pigF IIA —re l—rger th—n those of qir—ff—

pygm—e— from uoo˜i por— @r—rris IWWI˜AF

hes™ription

p—mily fovid—e qr—yD IVPI

ynly five dent—l spe™imens from the tot—l fui—

ƒu˜f—mily fovin—e qr—yD IVPI

fossil ™olle™tion ™—n ˜e referred to — sm—ll sized spe™ies

„ri˜e „r—gel—phini terdonD IVUR

of the genus wetridio™hoerusD wetridio™hoerus —ffF wF

wem˜ers of the tri˜e „r—gel—phini —re medium to modestus @„—˜le VAF

l—rge —ntelopes with ™lo™kwise spir—l hornE™ores @on the „he spe™imen hexEPIV is — very old individu—l

right sideAF „hey —re mostly ˜rowsersD h—ving ˜r—™hyE —nd no —n—tomi™—l ™h—r—™teristi™s ™—n ˜e des™ri˜edF „he

dont teeth without go—t fold on the lower mol—rs @r—rE other spe™imensD eERRW @pigF IHAD eEQTUD hexEIWU

ris IWWI™AF „his tri˜e w—s endemi™ to the efri™—n ™onE —nd hexEPPHD —re —ll lower third mol—rs @w AF „hey Q

tinent during the €lioE€leisto™eneD —s it is nowF „he present en—mel in the l—˜i—l —nd lingu—l surf—™esD ˜ut

iur—si—n —ntelopes with spir—l hornE™ores @q—zellospir—D —lso in the mesi—l —nd dist—l onesF sn the trigonidD the

ƒpiro™erusD €onto™erosD —nd othersA —re sometimes inE first p—ir of lo˜ules @proto™onid —nd met—™onidA —re seE

™luded in the „r—gel—phiniD ˜ut —ll of them h—ve well p—r—ted from the se™ond p—ir @hypo™onid —nd ento™oE

m—rked ––g—prini99 fold in the lower mol—rs whi™h is nidA ˜y two p—ir of ™entr—l tu˜er™ulesF „he trigonid is

never present in this efri™—n tri˜eF —lso sep—r—ted from the t—lonid ˜y two ™entr—l tu˜erE

™ulesF „he t—lonid termin—tes in sm—ll imp—ired lingu—l

wF modestus @r—rris 8 ‡hite —nd l—˜i—l pill—rs —s in P/4 M/1 M/2 M/3

LW LW LW LWL trig

IWUWAF elso the en—mel ™orrug—tion is m—rkedF UA-449 45,1 13,5 17,5 ‡e ˜elieve th—t the spe™imens from fui— ™orreE DAN-218 10,9 8,6 12,7 12,2 15,6 14,1

12,9 wetridio™hoerus —ffF wF modestusD spond to — form of DAN-197

DAN-220 R. 42,2 14,4 18,4

wetridio™hoerusF „he fui— s—mE

the sm—ll size spe™ies of UA-367 L 13,7 17,7

ple would represent one of the l—st o™™urren™es of this M. modestus 8,6 6,4 14,5 9,9 17,4 13,2 47,6 15,5 spe™ies in the i—rly €leisto™eneF

‡hite 8 r—rris @IWUWA re™ognize four spe™ies „—˜F V E vower teeth of wetridio™hoerus —ffF wF modestus from

fui—D —nd me—n of ext—nt wF modestusF within the genus wetridio™hoerusF wF —ndrewsi is the

possil v—rge w—mm—ls from fui— US

pigF II E qir—ff— ™fF qF jum—eF eEhX

hexEUHD right t—lus in l—tE

er—l @eAD medi—l @fAD dors—l

@gA —nd ventr—l @hA viewsF

f—r s™—le in ™entimetresF

„r—gel—phus fl—invilleD IVIT M2/ M3/ qenus

L WAL WPL L WAL WPL ™fF „F spekei ƒ™l—terD IVTQ

UA-251 33,50 41,90 27,00 21,00 „r—gel—phus

DAN-198 32,60 w—teri—l

P

@ eEPURAD I left m—ndi˜le with € E€ —nd roots of w E yne w Q R I

„—˜F W E pper teeth of qir—ff— ™fF qF jum—e from fui—

w @ eEPRSAD I ™omplete r—diusEuln— @wrfEPAD I ti˜i— dist—l end P

@wrfEQAF M2/ hes™ription

L WAL WPL ± ynly one spe™imen of — sm—llE

UA-274 L. 18,7 14,8 13,4 pper dentition

Tragelaphus spekei medium size tr—gel—phine h—s ˜een foundD eEPUR@pigF P

F st is — ˜r—™hydont toothD with wide

OM-1895 L. 15,1 17,0 14,7 IPdEgAD — left w

OM-5200 L. 18,0 14,0 13,3 lo˜ules —t the ˜—se th—t de™re—se r—pidly to the —pex of

Tragelaphus scriptus the ™rownF „he lingu—l f—™e h—s no intern—l pill—rF „he

OM-935 L. 14,9 11,3 11,3 l—˜i—l f—™e shows — fl—ttened p—r—™one —nd met—™oneD

OM-926 L. 13,7 11,0 11,1 —nd the three styles @p—r—styleD mesostyle —nd met—styleA

—re presentF „he me—surements of this spe™imen —re siE

„—˜F IH E „r—gel—phus ™fF „F spekeiD wPG me—surements ™omp—red mil—r to those of the m—le of „r—gel—phus spekei @ywE

to ext—nt spe™imens SPHHAD from the x—tion—l wuseum of ueny— @„—˜le IHAF

UT fF w—rtõnezEx—v—rroD vF ‚ook et —lF Â

L PTD PAPD MTD MAPD DTD DAPD MHB-2 201,6 35,3 20,1 24,3 15,9 34,0 23,5 Tragelaphus spekei OM-1895 R. 202,0 33,4 19,0 21,2 14,4 33,5 29,3 OM-5200 R. 227,5 37,2 21,3 25,6 15,7 38,9 24,0 Tragelaphus scriptus

OM-935 L. 201,0 34,6 19,0 24,7 15,1 34,1 19,2

„—˜F IP E „r—gel—phus ™fF „F spekeiD r—dius me—surements ™omp—red

to ext—nt spe™imens

pigF IP E „r—gel—phus ™fF „F spekeiF eEgX e PRSD right m—ndi˜ul—r

E€ in o™™lus—l @eAD ˜u™™—l @fA —nd lingu—l ™orpus with € P Q

P in dist—l @hAD ˜u™™—l @iAD @gA viewsY hEqX e PURD left w

lingu—l @pA —nd o™™lus—l @qA viewsF f—r s™—le in ™entiE

metresF

vower dentition ± ynly one spe™imen h—s ˜een

determinedD the right hemim—ndi˜le eEPRS @pigF IP—E

—nd € D —nd the roots of the w ™AF st preserves the € Q R I

—nd w F „he morphology of the premol—rs is typi™—lly P

„r—gel—phusD with low ™rowns —nd no p—r—™onid —nd

met—™onid jun™tionF nfortun—tely we ™—nnot s—y —nyE

thing —˜out the mol—rs @in „—˜le IID me—surements of

—nd w for eEPRS —re —lveolus me—surementsD the w I P

—nd would ˜e — little ˜it sm—ller th—n the re—l onesAF es

pigF IQ E „r—gel—phus ™fF „F spekeiF eEgX wrfEPD right r—dioEuln—

for the upper spe™imenD the —n—tomy —nd me—surements l—ter—l @eAD dors—l @fA —nd medi—l @gA viewsF f—r s™—le in

of eEPRSD —re more simil—r to the spe™imens of „r—E ™entimetresF

gel—phus spekeiD th—n to „r—gel—phus s™riptus @€—ll—sD

IUTTAD in the ™olle™tions of the x—tion—l wuseum of

ueny— @„—˜le IIAF €ost™r—ni—l skeleton ± ynly one ™omplete right

r—diusEuln— @wrfEPA ™—n ˜e referred to „r—gel—phus

„F spekei @pigF IQAF st is — spe™imen slender —nd P/3 P/4 M/1 M/2 ™fF

L W L W L WAL WPL L WAL WPL

„F spekei th—n

UA-245 L. 11,5 6,6 14,0 7,1 13,1 6,2 6,5 15,9 8,9 9,4 gr—™ileD —nd its me—surements —re ™loser to

Tragelaphus spekei

@„—˜le IPAF OM-1895 L. 11,1 8,0 12,2 8,5 13,2 10,7 10,4 15,9 12,3 12,2 „F s™riptus

OM-5200 L. 12,3 7,0 14,2 7,6 14,8 8,7 8,9 17,5 9,4 9,9 „r—gel—phus

Tragelaphus scriptus ‡e ™on™lude th—t the spe™ies of

„r—E OM-935 L. 10,8 5,2 11,9 5,8 11,6 6,6 7,3 14,3 7,4 8,2 found in the i—rly €leisto™ene of the fui— —re— is

OM-926 L. 11,4 6,0 11,4 6,6 12,3 7,1 7,4 14,0 8,0 8,3 gel—phus ™fF „FspekeiD the sit—tung—D —n —ntelope —d—pted

to w—ter environments with — very long dist—l ph—l—nges „—˜F II E „r—gel—phus ™fF „F spekeiD lower dentition me—surements

™omp—red to ext—nt spe™imens used for swimming —nd w—lking on mudfl—tsF

possil v—rge w—mm—ls from fui— UU

pigF IRE €elorovis oldow—yensisF eE

gX eERSID left hornE™ore

fr—gment in dors—l @eAD

ventr—l @fA —nd medi—l @gA

viewsF f—r s™—le in ™entiE

metresF

„he l—rge ylduv—i ˜uff—lo is well re™orded from the fui— sitesF

ƒu˜f—mily fovin—e qr—yD IVPI e more or less ™omplete ™r—niumD with ˜oth hornE™oresD w—s re™overed

from the site eHWRin xovem˜er PHHQD ˜ut it is still unEprep—red so th—t „ri˜e fovini qr—yD IVPI

its morphology ™—nnot ˜e des™ri˜edF

qenus €elorovis ‚e™kD IWPV

€elorovis oldow—yensis ‚e™kD IWPV

hes™ription

rornE™ores E ep—rt the ™r—nium mentioned —˜oveD

w—teri—l

the l—rgest preserved €elorovis hornE™ore fr—gment in „hree right hornE™ore fr—gments of the s—me individu—l @ eE

the fui— ™olle™tion is the spe™imen eERSI @pigF IRAF IQVD eEIRP —nd eEIRRAD I —pi™—l left hornE™ore fr—gment @ eE

IRQAD I left hornE™ore fr—gment @ eERSIAD I hornE™ore fr—gment st is — stout fr—gmentD dorsoEventr—lly fl—ttenedD with

I P I

Ew @ eERSHAD I w @ eEISHA I left m—xill—ry fr—gment with w

ov—l ™rossEse™tionD th—t h—s intern—l —nd extern—l

Q

@hexETQA I left w @ eEITVAD P upper mol—r fr—gment @ eEVD

groovesD —nd ™orresponds to the middle p—rt of the horn

—nd fr—gmented h€ eEIQRAD I left m—ndi˜ul—r fr—gment with h€ Q R

™oreF sts me—surements @„h —nd h†h of the ˜—s—l preE

@hexEVQAD I left s™—pul—r fr—gment @ eEQTHAD I right s™—pul—r fr—gE

served —re—A IPVFS —nd IHHFH mmD respe™tivelyF „his ment @ eEQSWAD I right r—diusEuln— @ eEQTIAD I dist—l end right r—E

diusEuln— @hexEPQSAD P left t—lus @hexEIPUD hexEPQUAD P right —n—tomy is typi™—l of €elorovisF

I P Q t—lus @ eETSD hexEVSAD I left ™—l™—neum fr—gment @ eEQVHAD I right DIw —nd I w —re pper dentition E „wo w

n—vi™ulo™u˜oideum @ eEQIIAD I dist—l fr—gment of w™ @ eERTAD I

identified in the ™olle™tion @„—˜le IQAF „hese mol—rs

dist—l end left wt @ eERSWAD I dist—l fr—gment of wt @ eEITSAD Q

I

h—ve thi™k en—mel —nd ro˜ust intern—l pill—rsF „he w medi—l ph—l—nges @ eEIHTD hexEIWTAF

M1/ M2/ M3/ L WAL WPL L WAL WPL L WAL WPL

UA-168 36,70 29,80 28,40

€elorovis DAN-63 26,20 28,40 28,70 „—˜F IQ E pper teeth of

UA-450 25,60 31,40 oldow—yensis from fui—F

UV fF w—rtõnezEx—v—rroD vF ‚ook et —lF Â

developedF „he p—r—styleD mesostyle —nd met—style —re

—lso pronoun™edF „he fossettes —re not ™ompli™—tedF

„he v—te €lio™ene spe™ies €elorovis turk—nensis

@r—rris IWWI™A is ™h—r—™terised ˜y the presen™e of —

m—rked en—mel fold in the posteroElingu—l —re— of the

hypo™one lo˜uleF „his fold is present in the fui— spe™iE

mensD ˜ut it is less m—rked th—n in €F turk—nensis from

uoo˜i por—F „his ™h—r—™ter pro˜—˜ly me—ns th—t the

fui— spe™imens were more evolved formsD simil—r to

those of the i—rly €leisto™ene from ylduv—iD €elorovis

oldow—yensisF

vower dentition ± xo lower —dult dentition spe™iE

mens h—ve ˜een re™ordedD —t the momentD in the fui—

sitesF ynly the m—ndi˜ul—r fr—gment @hexEVQA with

—nd h€ is fr—gment—ry —nd poorly preserved h€ Q R

known in the ™olle™tionF

€ost™r—ni—l skeleton ± „he post™r—ni—l spe™imens

@pigF ISA f—ll within the —n—tomi™—l —nd metri™—l v—ri—E

˜ility of €elorovis @„—˜le IRAF

his™ussion —˜out the genus €elorovis

„he genus €elorovis evolved from the i—rly €lioE

™ene efri™—n genus ƒim—therium @†r˜— IWVUAD —nd is the

l—rge v—te €lio™ene —nd €leisto™ene ˜uff—lo of efri™—F st

pigF IS E €elorovis oldow—yensisF eEhX eETSD right t—lus in l—ter—l

is — ™ommon ˜ovine in the efri™—n s—v—nn—hs until the

@eAD medi—l @fAD dors—l @gA —nd ventr—l @hA viewsF f—r

extin™tion of its l—st represent—tive spe™iesD €elorovis s™—le in ™entimetresF

—ntiquusD in the v—te €leisto™eneF €elorovis oldow—yenE

sis h—s ˜een des™ri˜ed from ylduv—iD —nd the holotype

hexETQ represents —n old individu—lF st h—s ™ementum

@field num˜ers ISITD ISIUD —nd ISIVY ‚e™k IWPVA w—s

on the l—˜i—l f—™eD —nd the p—r—styleD p—r—™oneD mesosE pu˜lished from the lo™—lity ‚uD ˜ut hietri™h @IWQQA

tyleD met—™one —nd met—style —re well developedF yn suggested th—t it ™—me from the fed s†F yn the ™ontr—ryD

the lingu—l f—™eD proto™one —nd hypo™one —re —lso develE qentry 8 qentry @IWUVA pu˜lished th—t —ll the rem—ins

opedF „he en—mel of the fossettes is not ™ompli™—tedD of this spe™ies from ylduv—i —re ™onfined to middle —nd

—nd the spe™imen shows —n en—mel isl—nd ˜etween

upper fed ss —nd fed sssF

P Q themF „he w eERSH is poorly preservedF „he w €elorovis oldow—yensis is —n evolved form th—t

eEITV h—s thi™k en—mel —nd thi™k ™ementum over dispersed to the northF st w—s unknown outside of efriE

most of the extern—l surf—™eD espe™i—lly on the l—˜i—l ™— until the first re™ord in – ˜eidiy— @qer——ds IWVTAF

f—™eF „he p—r—™one —nd met—™one —re ro˜ust —nd wellE v—terD other re™ords of €elorovis in the vev—nt h—ve

˜een ™itedF €F ™fF €F oldow—yensisD —t the i—rly €leistoE

RADIUS ™ene deposits of the en p—fud desert in the north of L PTD PAPD MTD MAPD DTD DAPD ƒ—udi er—˜i— @„hom—s et —lF IWWVAF „he s—me spe™ies is

UA-361 365,00 95,50 57,00 38,00 78,00 50,90

reported —t qesher fenot ‰—9—qovD in ssr—el @w—rtõÂnezE TALUS x—v—rro et —lF PHHHAD — site d—ted —t the i—rlyEwiddle

ML EL PTD DTD

DAN-85 75,10 80,80 51,30 53,40 €leisto™ene tr—nsition @qorenEsn˜—r et —lF PHHHAD where DAN-127 73,70 82,40 54,10 52,40 it is represented ˜y — ™r—nium —nd sever—l teeth —nd

UA-65 79,50 84,10 59,50 56,00

DA-237 78,40 82,00 2,09 54,80 post™r—ni—l elementsF e simil—r form w—s —lso re™orded

in the xorth efri™—n site of ein r—ne™h @elgeri—AD deE

fos ˜u˜—loides @er—m˜ourg IWUWAF e™tu—llyD 2nd PHALANX s™ri˜ed —s

L PTD PAPD DTD DAPD

h—s never ˜een found in other —re—s of iur—si—

UA-106 50,00 37,10 35,00 29,20 34,40 €elorovis UA-195 56,70 36,50 39,50 31,20 32,70 outside the vev—ntD ˜ut its —rriv—l into this region of

UA-7 27,70 29,30

DAN-196 55,00 38,20 38,80 30,00 western esi—D together with other open h—˜it—t efri™—n

rumin—ntsD suggests th—t efri™—n s—v—nn—h f—un—s exE

tended into iur—si—n middle l—titudes during i—rly „—˜F IRE €ost™r—ni—l spe™imens of €elorovis oldow—yensis from

fui—F €leisto™ene timesF

possil v—rge w—mm—ls from fui— UW

pigF IT E rippotr—gus gig—sF eEfX hexEIVPD ™—lv—rium fr—gment in front—l @eA —nd l—ter—l @fA viewsY gEpX eESID right met—™—rpus in ventr—l

@gAD l—ter—l @iAD dors—l @hA —nd medi—l @pA viewsY qErX eERQQD left horn ™ore in —nterior @qA —nd medi—l @rA viewsF f—r s™—le in ™entimetresF

VH fF w—rtõnezEx—v—rroD vF ‚ook et —lF Â

pigF IU E uo˜us ™fF uF ellipsiprymnusF eEgX eEIRHEIRID left horn ™ore fr—gment in medi—l @eAD —nterior @fA —nd l—ter—l @gA viewsF f—r s™—le in

™entimetresF

hes™ription ƒu˜f—mily rippotr—gin—e ‚etzius 8 voveÂnD IVRS

„he ™r—ni—l fr—gment hexEIVP @„—˜le ISA preE „ri˜e rippotr—gini ‚etzius 8 voveÂnD IVRS

serves ˜oth front—lsD joined ˜y the interEfront—l suture qenus rippotr—gus ƒundev—llD IVRT

@pigF ITAF „he left or˜it—l ™—vity is mostly ™ompleteD —nd

rippotr—gus gig—s ve—keyD IWTS

is situ—ted just ˜elow the hornE™ore pedi™elD whi™h is

tot—lly pneum—ti™ in the ˜—seF „he front—l sinus is develE w—teri—l

oped ˜ut not —s in the el™el—phiniF „he frontoEp—riet—l yne ™r—nium fr—gment @hexEIVPAD P hornE™ore fr—gments

@ eERQQ —nd hexEPISAD I medi—l ph—l—nx @ eEQUUAD I dist—l end of suture is situ—ted QH mm ˜ehind the posterior f—™e of the

— met—™—rp—l @ eESIAD —nd I dist—l ph—l—nx @ eEQUTAF hornE™oresD —nd is „Esh—ped to the interEfront—l oneF „he

˜r—in™—se roof is fl—ttenedD ˜ut its uppermost point is situ—ted on the interfront—l sutureF „he l—ter—l edge of

Biparietal Transv. Diam. 86,50 the roof is m—rked ˜y — ™rest whi™h st—rts —t the ™entr—l

Supraorbital foramens Transv. Diam. 63,00 posterior f—™e of the hornE™oreF „he ˜—se of the hornE External Transv. Diam. at the base of the Horn-cores 117,80 ™ore is ov—l with no keelsF

Tansv. Diam. Left horn-core 47,40 „he hornE™ore fr—gments eERQQ —nd hexEIPSD

Antero-Posterior Diam. Left horn-core 53,20

—re stout —nd h—ve — su˜ ov—l se™tion with — posterior

keelD m—rked in the hexEIPS spe™imenF „hey h—ve —

„—˜F IS E we—surements of the rippotr—gus gig—s ™r—ni—l fr—gment

longitudin—l ™urv—ture with no torsionF eERQQ h—s — hexEIVP

preserved length of PSH mmD —nd ˜—s—l tr—nsverse —nd

—nteroEposterior di—meters of QWFUH mm —nd RIFHH mm

Preserved Transverse Antero-posterior respe™tivelyY hexEIPS h—s — preserved length of IHQ

Length Diameter (base) Diameter (base) mmD —nd ˜—s—l tr—nsverse —nd —nteroEposterior di—E UA-60 227,00 46,30 55,70 meters of QUFPH mm RUFQH mmD respe™tivelyF

UA-30 95,50 37,00 40,00

is — l—rge rippotr—gini deE

UA-141+UA-140 50,70 45,70 57,00 rippotr—gus gig—s UA-136 297,00 43,60 54,50 s™ri˜ed from ylduv—iD widely distri˜uted in efri™—D

UA-137 255,00 41,00 50,70

UA-276 90,00 34,70 37,90 —nd ™ommon in i—rly €leisto™ene efri™—n sites @qentry DAN-99 490,00 51,00 64,30 8 qentry IWUVAF UA-436 105,00 42,70 50,10

DAN-148 466,00 41,90 56,60

DAN-148 115,00 43,90 51,30 ƒu˜f—mily ‚edun™in—e vydekker 8 fl—ineD IWIR

DAN-207 145,00 45,70 55,20 „ri˜e ‚edun™ini vydekker 8 fl—ineD IWIR

qenus uo˜us ƒmithD IVRH

„—˜F IT E rornE™ore fr—gments me—surements of uo˜us ™fF uF elE

lipsiprymnus from fui—F uo˜us ™fF uF ellipsiprymnus @ygil˜yD IVQQA

possil v—rge w—mm—ls from fui— VI

hes™ription „he most —˜und—nt —ntelope from fui— is — w—terE

R ˜u™k simil—r to the ext—nt uo˜us ellipsiprymnusF „he reE „he upper ™heek teeth —re hypsodontD the € h—s

present—tives of the genus uo˜us —re l—rge size hypsodont the styles well m—rked on the l—˜i—l f—™e —nd the posE

‚edun™ini th—t live in gr—ssl—nds ne—r perm—nent w—terF teroElingu—l region h—s — sm—ll ™ingul—r pill—r @„—˜le IVAF

Q

shows — sm—ll intern—l pill—r —nd h—s no m—rked „he w

w—teri—l

styles in the l—˜i—l f—™e @pigF IWeEhAF „he me—surements

„wo nonEme—sur—˜le ™r—ni—l fr—gments @ eEIRHC eEIRID

of the post™r—ni—l elements @pigF IW—EdD iEmA —re proE hexEPHUAD hornE™ore fr—gments @ eEQHD eETHD eEIQTD eEIQUD

vided in „—˜le IWF eEPUTD eEIQS—D eEIRQ˜D eERQTD hexEWWD hexEIRVAD I left

I Q P Q

Ew @hexEISPAD I w Cw @hexEPHWAD I m—xill—ry fr—gment with w „hese d—t— do not —llow — spe™ifi™ referr—l for the

@hexEVUAD I w @ eEPSSAD I upper mol—r fr—gment @hexEPITAD I w

P Q fui— q—zell—F „he genus q—zell— is re™orded in efri™—

w™ dist—l end @hexEVVAD Rwt dist—l ends @ eEQPD eESVD eEPTID

eEPUSAD I wet—podi—l dist—l end @hexEIQHAD P t—lus @ eESPD hexE

UIAD I n—vi™ulo™u˜oideum @ eEPSTAD Q proxim—l ph—l—nges @ eEIUD

eESHD hexEIPTAD P medi—l ph—l—nges @ eEPTSD hexEIWHAF „here

—re —lso other post™r—ni—l spe™imens pro˜—˜ly ˜elonging to this t—xonF

hes™ription

gr—nium ± „he ™r—ni—l fr—gment eEIRHCIRI

@pigF IUA is he—vily ˜uilt —nd l—™ks — pneum—tised front—l

˜oneF st h—s — simil—r —n—tomy to the ext—nt m—le indiE

vidu—ls of uo˜us ellipsiprymnus housed —t the x—tion—l

wuseum of iritre—F rornE™ores —re longD with the ˜—se

™urving ˜—™kw—rds —nd the —pex ™urving upw—rdD forE

w—rd —nd inw—rdD mediol—ter—lly ™ompressed with su˜E

tri—ngul—r se™tions —t the ˜—se —nd fl—ttened posteroEinE

tern—l f—™es @„—˜le ITAF „he spe™imens exhi˜it wellE

m—rked tr—nsverse ridges @pigF IV™EdAF

hentition ± „he ™heek teeth —re hypsodontF

pper mol—rs show the three stylesD p—r—™oneD —nd meE

t—™one m—rked in the l—˜i—l f—™eD —nd pronoun™ed loE

˜ules @proto™one —nd hypo™oneAD with presen™e of —n

intern—l pill—r on the lingu—l f—™eF „he fossettes —re simE

ple @pigF IV—E˜AF vower mol—rs h—ve m—rked go—t foldsD

— prominent extern—l pill—rD —nd three m—rked intern—l

stylids with the met—™onid —nd ento™onid ˜eing proE

noun™edF „he me—surements @„—˜le IUA f—ll within the

v—ri—˜ility r—nge of this l—rge redun™ine —ntelope @r—rris

IWWI™AF

€ost™r—ni—l skeleton ± „he post™r—ni—l elements

f—ll within the v—ri—˜ility of uo˜us ellipsiprymnusF

„he ext—nt w—ter˜u™k spe™ies uo˜us ellipsiprymE

nusD pro˜—˜ly evolved from uF sigmoid—lisD is — ™ommon

l—rge ‚edun™ini in l—te i—rly €leisto™ene i—stern efriE

™—n fossil —ssem˜l—ges @qentry 8 qentry IWUVY qentry

IWVSY r—rris et —lF IWVVY r—rris IWWI™AF

ƒu˜f—mily entilopin—eD qr—y IVPI

„ri˜e entilopini qr—yD IVPI

qenus q—zell— fl—invilleD IVIT

q—zell— spF

w—teri—l

R Q

@ eEPTWAD I w @hexEPIIAD I left m—ndi˜ul—r fr—gment yne € pigF IV E uo˜us ™fF uF ellipsiprymnusF eEfX hexEISPD left m—xilE

I P

E€ @hexEPPWAD I „—lus @ eEPVIAD I r—dius proxim—l with —lveolus € Ew in ˜u™™—l @eA —nd o™™lus—l @fA l—ry fr—gment with w P Q

viewsY gEhX hexEIRVD left horn ™ore in medi—l @gA —nd end @hexETRAD I wet—™—rp—l dist—l end @hexEIWSAD —nd I proxim—l

l—ter—l @hA viewsF f—r s™—les in ™entimetresF ph—l—nx @ eEQTVAF

VP fF w—rtõnezEx—v—rroD vF ‚ook et —lF  M1/ M2/ M3/ „—˜F IU E pper —nd lower dentition

L WAL WPL L WAL WPL L WAL WPL

uo˜us ™fF

DAN-152 18,00 19,10 19,00 21,00 22,90 24,00 me—surements of

from fui—F DAN-209 22,70 22,70 21,30 28,00 21,70 17,70 uF ellipsiprymnus

M/2 M/3 L WAL WPL L WAL WPL DAN-87 23,00 13,90 13,50

UA-255 32,70 13,50 12,70

q—zell— spF from fui—F P4 M3 „—˜F IV E pper teeth spe™imens of L W L WAL WPL UA-269 L. 8,40 10,40

DAN-211 L. 14,20 11,00 9,90

EL IL PTD PAPD MTD MAPD DTD DAPD „—˜F IW E we—surements of post™r—ni—l

q—zell— spF Talus spe™imens of UA-281 28,00 25,00 16,20 16,40

Mc from fui—F DAN-195 24,50 19,10 1st phalanx

UA-368 10,90 15,50 9,60 10,50

Q pigF IW E q—zell— spF eEhX eEPVID right t—lus in dors—l @eAD ventr—l @fAD medi—l @gA —nd l—ter—l @hA viewsY iErD hexEPIID left w in lingu—l

@iAD ˜u™™—l @pAD o™™lus—l @qA —nd mesi—l viewsY sEwX hexEIWSD left fr—gment—ry met—™—rp—l in dors—l @sAD ventr—l @tAD l—ter—l @uAD

medi—l @vA —nd dist—l viewsF f—r s™—le in ™entimetresF

w—teri—l from the v—te wio™ene @„hom—s IWUWA —nd persists until

yne right m—ndi˜ul—r fr—gment with w Ew @hexEPIAD I w

P Q I

presentF

P

@ eEPUQAD I w @hexEIPWAD —nd I w @hexETPAF „here —re —lso Q dist—l P

humeri —nd I proxim—l ph—l—nx fr—gments th—t pro˜—˜ly ™orrespond to

this t—xonF

ƒu˜f—mily g—prin—e qr—yD IVPI

„ri˜e g—prini ƒimpsonD IWRS

hes™ription qenus indetF

„he tri˜e g—prini is represented —t fui— ˜y

g—prini genF et spF indetF some dent—l spe™imensF „hese dent—l spe™imens ™orreE

possil v—rge w—mm—ls from fui— VQ

pigF PH E g—prini indetFY eEhX

P in disE hexETPD left w

t—l @eAD ˜u™™—l @fAD linE

gu—l @™A —nd o™™lus—l

@hA viewsF iEqX hexE

PID right m—ndi˜ul—r ™orE

pus with w Ew in ˜u™E P Q

™—l @iAD lingu—l @pA —nd

o™™lus—l @qA viewsF f—r

s™—les in ™entimetresF

oogeogr—phi™ ™omp—rison of the fui— l—rge m—mm—l spond to — medium size ™—prine @pigF PHY „—˜le PHAF

—ssem˜l—ge using ƒimpson9s —nd hi™e9s ––qenus p—un—l „hey —re very hypsodontF „he upper mol—r @hexE

P ‚esem˜l—n™e sndex99 th—t shows prism—ti™ stru™ture with no TPAD is —n w

pronoun™ed p—r—E —nd met—stylesD more pronoun™ed

„o ™omp—re the fui— l—rge m—mm—l f—un— with

mesostyle —nd rel—tively pronoun™ed pill—rs of p—r—E

other €lioE€leisto™ene efri™—n —nd xe—r i—st f—un—l —sE

™one —nd met—™one on the l—˜i—l f—™eF yn the lingu—l

sem˜l—gesD we h—ve performed —n —n—lysis of genusEr—nk

f—™eD the —nterior lo˜ule is more mesioEdist—lly ™omE

f—un—l resem˜l—n™e indi™es @qp‚sAF ‡e sele™ted — numE

pressed th—n the posterior oneD —nd this l—st one h—s

˜er of efri™—n —nd xe—r i—st f—un—s whose —ges r—nge

— ridge on the posteroElingu—l —re—F „he lower mol—rs

˜etween P —nd HFS w—F p—un—l lists from whi™h this

h—ve no go—t fold —nd no mesostylidD the pill—rs of the

™omput—tion is ˜—sed —re derived from the —v—il—˜le litE

met—™onid —nd ento™onid —re well m—rked in the linE er—ture @qer——ds IWVIY „urner et —lF IWWWY esf—w et —lF

gu—l f—™eD —nd the se™tion of the l—˜i—l lo˜ules is very PHHPA in some ™—ses upd—ted ˜y the —uthors @fwx —nd

—ngul—rF v‚A person—l o˜serv—tions @„—˜le PIAF ‡e follow porE

qo—ts h—ve —n iur—si—n originD ˜ut ˜e™ome rel—E telius et —lF @IWWTA in ™—l™ul—ting ˜oth hi™e —nd ƒimpson

tively ™ommon in the e—stern i—rly €leisto™ene efri™—n qp‚s9sF hi™e is the one most highly re™ommended ˜y

—ssem˜l—gesD ™ited in the „urk—n— f—sin —t the x—rikoE er™her 8 w—ples @IWVUA —nd w—ples 8 er™her @IWVVA

—nd is ™—l™ul—ted —sX Pe G @Pe Cf CgA D where e is the tome —nd u—itio wem˜ers of the x—™hukui porm—tion

num˜er of t—x— present in ˜oth f—un—sD f the num˜er of @r—rris et —lF IWVVAF „he spe™ies g—pr— prim—ev— is —lso

t—x— present in f—un— I ˜ut —˜sent in f—un— PD —nd g the re™orded in the v—te €lio™ene of xorth efri™— —t the site

num˜er of t—x— present in f—un— P ˜ut —˜sent in f—un— IF of ein frim˜—D „unisi— @er—m˜ourg IWUWAF

M2/ L WAL WPL DAN-62 18,4 16,1 14,3 M1 M2 M3

L WAL WPL L WAL WPL L WAL WPL DAN-21 17,5 11,2 11,2 25,7 12 11,5 „—˜F PH E pper —nd lower teeth me—E

DAN-129 18,4 10,8 10,5

UA-273 16,2 9,8 9,8 surements of g—prini indetF from fui—F

VR fF w—rtõnezEx—v—rroD vF ‚ook et —lF Â

ogyY esf—w et —lF PHHPAD ylduv—i fed PD x—riokotomeD ‡e —lso use ƒimpson9s @IWRUD IWTHA f—un—l resem˜l—n™e

—nd ykote show the highest resem˜l—n™e v—lues to fui—F index ˜e™—use it h—s — long tr—dition of use in p—leontolE

– ˜eidiy— —nd qesher fenot ‰—9—qov @ssr—elY w—rtõÂnezE ogy —nd it ™orre™ts for differen™es in diversity of f—un—s

x—v—rro et —lF in prepY qorenEsn˜—r et —lF PHHHA —s well —s ˜eing ™omp—red @plynn IWVTY fernor 8 €—vl—kis IWVUY

„ernifine @qer——ds IWVIA —nd ein r—ne™h @ƒ—hnouni et fernor et —lF PHHIAF ƒimpson9s f—un—l resem˜l—n™e index

—lF PHHPA in ˜oth plots shows the lowest v—luesF „his is is ™—l™ul—ted —sX e G @e CiA where i is the sm—ller of f

either due to — differen™e in —ge in some of themD —nd to or gF „he l—tter —djusts for differen™es in s—mple sizes

the strong influen™e of ––iur—si—n99 t—x— o™™urring —t ˜etween p—irEwise f—un—s ˜eing ™onsidered @—s the ™—se

those l—tter sites @eFgF deerD ––iur—si—n99 pro˜os™ide—ns of the s—mple used hereA —nd h—s — demonstr—˜le ro˜ustE

—nd ™—rnivoresAF st is worth to noteD howeverD th—t the ness in p—leontology @fernor IWUVAF

qp‚s v—luesD —lthough low @with respe™t to those shown pigure PI in™ludes two plots of qp‚sF „he first

˜y the i—st efri™—n sitesAD do™ument — rel—tively high one provides p—irEwise ™omp—risons ˜etween fui— —nd

degree of simil—rity ˜etween the xe—r i—st f—un—s —nd the set of lo™—lities under ™onsider—tionD the se™ond one

fui— @note the ƒimpson9s v—lue of HFSHH for the fui—E provides the p—irEwise ™omp—risons ˜etween x—riokoE

– ˜eidiy— p—irEwise ™omp—risonAF „his is m—inly due to tome —nd the other lo™—litiesF sn the first one it is to note

the ™ommon o™™urren™e of t—x— su™h —s „heropithe™usD how the ithiopi—n site h—k— @this f—un— is the ™losest to

uolpo™hoerusD rippopot—musD q—zell— —nd €elorovisF fui— ˜oth in terms of geogr—phi™ dist—n™e —nd ™hronolE

„he s—me is evident —lso for the

™omp—rison fui—Eqesher fenot GFRI to Buia ‰—9—qovF „he l—tter site m—int—ins

1 — rel—tively signifi™—nt degree of

0,9 simil—rity with fui— @with — ƒimpE

0,8 son9s v—lue of HFQVRA —nd gener—lly

0,7 spe—king with — €leisto™ene 0,6 ––efri™—n ˜ioprovin™e99 @™fF the

0,5

0,4 qp‚s to x—riokotomeD low ˜ut

0,3 —˜ove HFPA th—nks to the o™™urE

rippopot—E

0,2 ren™e of t—x— su™h —s

D q—zell— —nd €elorovisF 0,1 mus

0 „he p—irEwise ™omp—rison to x—riokotome is p—rti™ul—rly

Okote Daka Olduvai 1 Olduvai 2 Olduvai 3 Ternifine 'Ubeidiya inform—tiveF „he highest simil—rE

Ain Hanech Nariokotome ity indi™es —re found with ykote

Gesher Benot Y. —nd ylduv—i fed P @™loser in sp—™e —nd time to x—riokotomeAF

GFRI to Nariokotome e highD —lthough slightly lower

v—lueD is —lso shown ˜y h—k—

1 —nd fui— @˜oth very ™lose in time

0,9 —nd sp—™eAF ylduv—i feds I —nd QD

0,8 on the other h—ndD show low siE

0,7 mil—rity v—luesD ˜ut this ™ould ˜e 0,6 due to the rel—tive sm—ll s—mple 0,5

0,4 from the two lo™—litiesD while

0,3 even lower v—lues ™h—r—™terize

0,2 ein r—ne™hD „ernifineD – ˜eidiy—

0,1 —nd qesher fenot ‰—9—qovF „he

0 l—tter is —n expression of their m—E jor dist—n™e in sp—™e —ndGor time

Daka Buia Okote from x—riokotomeF Olduvai 1 Olduvai 2 Olduvai 3 Ternifine 'Ubeidiya Ain Hanech

Gesher Benot Y.

gon™luding rem—rks

pigF PI E qenus p—un—l ‚esem˜l—n™e sndex @qp‚sA plots ™omp—ring the set of lo™—lities to fui—

„he l—rge m—mm—l —ssemE @topA —nd to x—riokotome @fottomAF qr—y ˜—rs hi™e9s qp‚sD fl—™k ˜—rs ƒimpson9s

qp‚s @see text for expl—n—tion —nd dis™ussionAF ˜l—ge from fui— represents — l—te

possil v—rge w—mm—ls from fui— VS

1234567891011 1234567891011 XXX Paranthropus Ceratotherium XX X XXXX XXXX XXXXXX Homo Diceros XX X XX XX XX Theropithecus Stephanorhinus X X XX Parapapio Kolpochoerus XXXXXXXX X XX Papio Metridiochoerus XXXXXXX X X Cercocebus Sus X X X Macaca Hexaprotodon XX XX Cercopitecoides Hippopotamus XX X XXXXXX XXXX Colobus Sivatherium XXX X Atillax Giraffa X X XXXX X Crossarchus Camelus XXX Herpestes X X Beatragus XX Ichneumia X Alcelaphus XX Mungos XX Sigmoceros X Pseudocivetta X X Connochaetes XXXX X Lutra XX X Damaliscus XX Vormela X Megalotragus XXXXXX Pannonictis X Parmalarius X XXXX X Homotherium X Hippotragus XXXXX Dinofelis X Brabovus X Megantereon X X Oryx X X X Panthera XXXX Kobus XXXXXXX X Acinonyx X Menelikia X Felis X Redunca XXX Lynx X Gazella X XX XXXXXX Pachycrocuta X Antidorcas X XXX Crocuta XXXX XXXX Tragelaphus XXXXXXX X Hyaena XXX Taurotragus XX Canis X X Syncerus XXX Lycaon XXX XX Pelorovis X XXXXXXXX Vulpes X X Bos X Ursus XX Aephiceros XXXX X Loxodonta X Rabaticeras X Elephas XXXXX Bouria X Mammuthus X X Nitidarcus X Stegodon X Numidocapra X Palaeoloxodon X Praemegaceros X Hipparion X XXX Megaloceros X Equus XX X XXXXXX Dama X

Cervus X

IA ykote mem˜erY PA x—riokotomeY QA ylduv—i fed IY RA ylduv—i ˜ed PY SA ylduv—i ˜ed QY TA h—k—Y UA fui—Y VA r—in ene™hY WA „ernifineY IHA

– ˜eidiy—Y IIA qesher fenot ‰—9—qov

„—˜F PI E qenus o™™urren™es in the lo™—lities used fro the qp‚s —n—lysisF

predomin—ntly w—terEdependent t—x— sin™e the ™ro™oE i—rly €leisto™ene —sso™i—tion of the i—stern efri™—n

dile —nd the pelomedusid ™heloni—n rem—ins —re found ‚iftD ™h—r—™terised ˜y evolved l—st represent—tive forms

stri™tly —sso™i—ted to —ll kind of —qu—ti™ environments of „heropithe™us ™fF „F osw—ldiD ileph—s re™kiD rippopoE

—nd the efri™—n ro™k pythons —re most —˜und—nt ne—r t—mus gorgopsD uolpo™hoerus olduv—iensisD uolpo™hoerus

low —ltitude riversD l—kes —nd sw—mps @helfino et —lF m—jusD wetridio™hoerus —ffF wF modestus —nd €elorovis

PHHRAF „he p—leoe™ologi™—l ™h—r—™teristi™s of the f—un— oldow—yensisD —nd some others th—t persist until present

fully —gree with the kind of environment do™umented like ger—totherium simumD iquus ™fF iF grevyi „r—gel—E

˜y the sediment—ry depositsD whi™h ™orrespond to — phus ™fF „F spekei or uo˜us ™fF uF ellipsiprymnusF „he

fluvioEdelt—i™ —nd l—™ustrine environments @e˜˜—te et geologi™—l —nd sedimentologi™—l surveyD p—leom—gnetE

—lF PHHRAF ism —nd fission tr—™k d—tings —re in good —greement with

„he s™—r™e presen™e of ™—rnivores in the f—un—l d—t— of m—mm—l —ssem˜l—ge ˜io™hronologyD —ttesting to

—ssem˜l—ge ™—n ˜e expl—ined —s due to poor s—mpling the romoE˜e—ring level —n —ge of —pproxim—tely I w—

@e˜˜—te et —lF PHHRY el˜i—nelli 8 x—poleone PHHRY fiE in the fui— ˜edsD ˜ut m—y —lso ˜e rel—ted to hum—n

g—zzi et —lF PHHRAF —™tivity in the sites @piore et —lF PHHRAD where most of

„he m—mm—l —ssem˜l—ge is predomin—ntly ™omE the fossil l—rge verte˜r—te ˜ones —re —sso™i—ted with

posed of t—x— with — strong w—ter dependen™e @hipposD —˜und—nt e™heuli—n lithi™ —rtef—™tsF

w—terE˜u™kD sit—tung—D uolpo™hoerusAD —sso™i—ted with „he —n—lysis of ––p—un—l ‚esem˜l—n™e sndex99

mu™h less represented m—mm—li—n t—x— typi™—l of more @qp‚sA with some well known f—un—l —ssem˜l—ges of

efri™— —nd xe—r i—st reve—led some interesting ™luesF open l—nds™—pesF „he herpetof—un— —lso is ™omposed of

VT fF w—rtõnezEx—v—rroD vF ‚ook et —lF Â

‚ese—r™h @w ‚ƒ„Gws ‚AY st—li—n x—tion—l ‚ese—r™h goun™il @gx‚E es expe™tedD the fui— —ssem˜l—ge over—ll shows the

g„HSD —nd gx‚ €rogetto pin—lizz—to feni gultur—liAY the ve—key pounE highest resem˜l—n™e with i—st efri™— —ssem˜l—ges like

d—tionY —nd the niversity of ploren™eF ‡e —re deeply inde˜ted to pFgF

ykoteD ylduv—i ˜ed PD —nd h—k—F roweverD in respe™t

rowell —nd ‚FvF fernor for supportD suggestionsD —nd ™riti™—l re—ding of

to other i—st efri™—n —ssem˜l—gesD fui— h—s — rel—tively

—n e—rly version of the m—nus™riptF fFwFxF th—nks the ve—key —nd

high simil—rity with some xe—r i—st f—un—s like – ˜eiE ge‚i pound—tions for fin—n™ing the ™omp—r—tive studies —t x—iro˜i

@ueny—A —nd terus—lem @ssr—elAF xumerous friends in iritre— —nd —round diy—D —nd qhesher fenot ‰—9—qovF

the world in different w—ys h—ve ˜een of help —nd support to the proje™tF

emong theseD we mention the people from the fui— vill—ge for their e™knowledgmentsF „he fui— €roje™t —™tivities h—ve ˜een possiE

w—rm hospit—lityF por prelimin—ry prep—r—tion of spe™imensD in the field ˜le th—nks to fin—n™i—l support ˜y sever—l funding institutionsX st—li—n

poreign winistry @weiAY st—li—n winistry for idu™—tion niversity —nd —nd in esm—r—D we th—nks pF gozzini —nd pF v—ndu™™i @ploren™eAF

‚ipi‚ixgiƒ

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m—ri—m fFD fruni €FD gipri—ni xFD gl—rke ‚F tFD pi™™—rE woden—F

elli qF w—™™hi—relli ‚FD x—poleone qFD €—pini wFD fernor ‚FvF 8 8 €—vl—kis €F @IWVUA E oogeogr—phi™ rel—E

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the h—n—kil @ef—rA hepression of iritre—F x—tureD QWQX ziry eF‡FD de reinzelin tF 8 fo—z hFhF @edsA ± xeoE

RSVERTHD vondonF gene €—leontology —nd qeology of ƒ—h—˜iX QQUEQRVF

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gionD iritre—n h—n—kil hepressionAF ‚ivF stF €—leontF in „he el—t porm—tion of the h—ndiero qroup @h—E

ƒtr—tFD IIH @supplementAX SEQRD wil—noF n—kil hepressionD iritre—AF ‚ivF stF €—leontF ƒtr—tFD IIH

el˜—nelli eF 8 x—poleone qF @PHHRA E w—gnetostr—tigr—phy @supplementAX RSERWD wil—noF

of the romoE˜e—ring €leisto™ene h—ndiero f—sin foisserie tFE‚F @PHHPA E xouve—ux rippopot—mid—e du wioE

@h—n—kil hepressionD iritre—AD ‚ivF stF €—leontF ƒtr—tFD €lio™eÁne du „™h—d et d9ithiopieX impli™—tions phyloE

IIH @supplementAX QSERRD wil—noF genetiques et p—leoenvironnement—lesF €hh thesisD   Â

er—m˜ourg gF @IWUWA E †erte˜res †ill—fr—n™hiens d9efrique niverity of €oitiersD ppF IERRHF

du xord @ertiod—™tylesD g—rnivoresD €rim—tesD ‚epE frunet wF 8 ‡hite „FhF @PHHHA E heux nouvelles espeÁ™es de

tilesD yise—uxAF ƒingerE€olign—™D €—risF ƒuini @w—mm—li—D ƒuid—eA du ™ontinent efri™—in

 er™her eF‡F 8 w—ples gFqF @IWVUA E wonte g—rlo simul—E @ithiopieY „™h—dAF gF ‚F e™—dF ƒ™iFD QQPX SIESUD €—risF

tion of sele™ted ˜inomi—l simil—rity ™oeffi™ient @sAX efE gh—v—illon tFD ferthelet eF fois—u˜ert tF vFD p—ure wFD queÂE

fe™t of num˜er of v—ri—˜lesF €—l—iosD PX THWETIUD flooE ren gF 8 ‡—rs—m—F ƒF eF @IWWHA E n ileph—s re™kiD

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ithiopi—F x—tureD RITX QIUEQPHD vondonF r—d—rD ithiopi—F uirtl—ndi—D PWX IETQD glevel—ndF

feden wF @IWVUA E ves p—unes €lioE€leisto™enes de l— †—lle helfino wFD ƒegid eFD ‰osief hFD ƒhosh—ni tFD ‚ook vF 8

de l9ymo @ithiopieAD „ome PX ves ileÂph—ntideÂs vi˜sek—l ‰F @PHHRA E possil ‚eptiles from the €leistoE

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@PAX ITSD terus—lemF sndi— —nd the t—xonomy of the genusF snX t—˜lonski

fernor ‚FvF @IWUVA E „he w—mm—li—n ƒystem—ti™sD fiostr—E xFqF @edFA E „heropithe™usX „he rise —nd f—ll of — priE

tigr—phy —nd fio™hronology of w—r—gheh —nd its imE m—te genusX ISUEIVWF g—m˜ridge niversity €ressD

port—n™e for nderst—nding v—te wio™ene rominoid g—m˜ridgeF

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fernor ‚FvFD portelius wF 8 ‚ook vF @PHHIA E ivolution—ry PWWEQHRD ƒtuttg—rtF

fiogeogr—phy —nd €—leoe™ology of the ––yreopithe™us iisenm—nn †F @IWVQA E p—mily iquideF snX r—rris tFwF @edFA E

˜—m˜olii p—un—l one99 @l—te wio™eneD „us™oEƒ—rdiE uoo˜i por— ‚ese—r™h €roje™tF †ol PX „he possil nE

possil v—rge w—mm—ls from fui— VU

gul—tesX €ro˜os™ide—D €erissod—™tyl—D —nd ƒuid—eX ISTE qer——ds hFD queÂrin gF 8 p—ure wF @IWVTA E ves suideÂs du

€leÂisto™eÂne —n™ien d9yu˜eidiyeh @ssr—elAF snX „™herE PIRD gl—rendon €ressD yxfordF È

nov iF @edFA E ves w—mmifeÁres du €leÂisto™eÁne infeÂrE iisenm—nn †F @IWVTA E ves iquideÂs du €leÂisto™eÁne d9yu˜eiE

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