‚ivist— st—li—n— di €—leontologi— e ƒtr—tigr—fi— volume IIQ noF I ppF TUEUU w—r™h PHHU

ew€rswegres‚yh ƒ @pivsheiD wewwevseA p‚yw ƒerefs

@ve„iƒ„ wsygixiEie‚vsiƒ„ €vsygixiD vsf‰eAD ‡s„r e ‚i†si‡

yp ep‚sgex wsygixi wegres‚yhyx„sxei

I P

‚eppeivi ƒe‚hivve 8 ve‚ƒ ‡i‚hivsx

‚e™eivedX w—y PRD PHHTY —™™eptedX t—nu—ry IHD PHHU

uey wordsX efri™—D vi˜y—D g—rnivor—D pelid—e w—™h—irodontiE ture of iur—si—n —nd efri™—n verte˜r—tesX eleph—ntidsD

n—eD wio™eneD €lio™eneF

—nthr—™otheres —nd hippos @q—ziry IWVU—Y IWVU˜Y

IWVU™AD suids @gooke IWVUAD ˜ovids @vehm—nn 8 „hoE

e˜str—™tF ‡e des™ri˜e —nd illustr—te — p—rti—l skull —nd m—ndi˜le

m—s IWVUAD the shortEne™ked gir—ffid ƒ—motherium @r—rE of — l—rge s—˜ertooth ™—t from ƒ—h—˜iD vi˜y—D —nd refer it to emphim—E

™h—irodus —ffF eF k—˜irF e review shows the wio™ene w—™h—irodontin—e ris IWVUAD the rhino™eros hi™eros @ager—totheriumA neuE

from efri™— to ˜e — heterogeneous —ssem˜l—geD with ˜oth sm—ll —nd m—yri @reissig IWWTAD hipp—rions @fernor 8 ƒ™ott PHHQAD

l—rge forms sp—nning the entire v—te wio™eneF „he ƒ—h—˜i form ˜elongs

sever—l ™—rnivores @rowell IWVPY IWVUY ‚ook 8 w—rtiE

to the group of l—rger sized t—x—D —long with eF k—˜ir from gh—d —nd

nez x—v—rro PHHRAD the rodent ƒ—yimys @wunthe IWVUY some previously undes™ri˜ed spe™imens from the ‡em˜ereEw—nong—

egustõÁ et —lF PHHHAD —nd — l—rge —n—tid @f—llm—nn IWVUAF porm—tionD „—nz—ni—F foth the ƒ—h—˜i —nd gh—d spe™imens h—ve rel—E

tively slender lower ™—rn—ssi—lsD simil—rly to romotheriumD though deE es su™hD ƒ—h—˜i is import—nt for ˜iogeogr—phi™ re™onE

rived fe—tures of the skull —nd m—ndi˜le suggest th—t they —re not in the

stru™tion —nd p—leoe™ologi™ ™omp—risons @fernor 8

dire™t —n™estry of th—t genusF

€—vl—kis IWVUAF emong the ™—rnivore rem—ins —re — skull

—nd m—ndi˜le of — l—rgeEsized s—˜ertoothed ™—t stored —t

‚i—ssuntoF n ™r—nio e un— m—ndi˜ol— di un— gr—nde tigre d—i

the €—leontologi™—l wuseum —t ––v— ƒ—pienz—99 niverE denti — s™i—˜ol— provenienti d— ƒ—h—˜i @vi˜i—A sono qui des™rittiD illuE

str—ti e —ttri˜uiti —d emphim—™h—irodus —ffF eF k—˜irF v— revisione dei sity in ‚omeF „hese fossils —re illustr—ted herein for the

m—™—irodontini —fri™—ni del wio™ene mostr— ™ome questi ™ostituissero

first time @pigF IA —nd des™ri˜ed in rel—tionship to other

un insieme eterogeneoD ™omprendente forme si— di pi™™ole si— di gr—ndi

efri™—n v—te wio™ene w—™h—irodontin—eF dimensioni distri˜uite per tutto il wio™ene superioreF v— form— di ƒ—E

„he e—rliest studies on the fossiliferous —re— of h—˜i —pp—rtiene —l gruppo dei t—x— piuÁ gr—ndiD insieme — eF k—˜ir del

—sr —sEƒ—h—˜i @vi˜y—A were ™—rried out ˜y the st—li—n gh—d e —d —l™uni esempl—ri m—i des™ritti in pre™edenz— provenienti d—ll—

porm—zione di ‡em˜ereEw—nong—D in „—nz—ni—F v9esempl—re di ƒ—h—˜i geologist erdito hesioD who led field —™tivities in IWQIE

e quello del gh—d h—nno in ™omune l— presenz— di ferini inferiori seE

QPF prom IWQREQWD further geologi™—l —nd p—l—eontoloE

™odontiD ™ome si osserv— in romotheriumD m— i ™—r—tteri deriv—ti di

gi™—l work w—s ™—rried out ˜y g—rlo €etro™™hiF et th—t

™r—nio e m—ndi˜ol— suggeris™ono ™he questi non possono essere ™onsiE

timeD p—rt of the m—teri—l w—s t—ken from vi˜y— to st—lyD der—ti ™ome i diretti progenitori di questo genereF

where it w—s stored in different museums @oologi™—l

givi™ wuseum —nd €—leontologi™—l wuseum —t ––v— ƒ—E

pienz—99 niversityD ˜oth in ‚omeD —nd the €—leontoloE sntrodu™tion

gi™—l wuseum of ––pederi™o ss99D niversity of x—plesAF

€etro™™hi @IWRIY IWRQY IWSIA identified TP fossiliferous „he l—test wio™ene verte˜r—te f—un— from ƒ—h—˜i

lo™—litiesF re mentioned the presen™e of — m—stodonD — @giren—i™—D vi˜y—A is — ri™h —nd diversified oneF es reE

˜ovidD —n —nthr—™othereD —nd one skull —nd two m—ndE ™ently emph—sized ˜y sever—l —uthors @qer——ds IWWVY

fernor 8 ƒ™ott PHHQAD the ƒ—h—˜i f—un— in™ludes — mixE i˜les of ™—rnivoresF re provided — des™ription of the

Á I hip—rtimento di ƒ™ienze dell— „err—D niversit— di ‚om— v— ƒ—pienz—D €i—zz—le eldo woro SD sEHHIVS ‚om—D st—lyF

iEm—ilX r—ff—eleFs—rdell—dunirom—IFit P hep—rtment of €—l—eozoologyD ƒwedish wuseum of x—tur—l ristoryD fox SHHHUD ƒEIHR HS ƒto™kholmD ƒwedenF iEm—ilX werdelindnrmFse

TV ƒ—rdell— ‚F 8 ‡erdelin vF

pigF I E emphim—™h—irodus —ffF eF k—˜ir from ƒ—h—˜iX skull in —A s—gitt—lD ˜A p—l—t—lD ™A o™™ipit—l —nd dA l—ter—l viewY left m—ndi˜le in eA ˜u™™—lD fA lingu—l —nd gA o™™lus—l viewF

emphim—™h—irodus from ƒ—h—˜iD vi˜y— TW

pigF P E vower right mID ‡w IWQUGWP—D ‡em˜ereEw—nong— pmFD „—nz—ni—D in —A ˜u™™—lD ˜A lingu—lD ™A o™™lus—l viewY dEfX lower left mID ‡w

IWQUGWP˜D ‡em˜ereEw—nong— pmFD „—nz—ni—D in dA ˜u™™—lD eA lingu—lD fA o™™lus—l viewY gEiX left m—ndi˜ul—r fr—gmentD ƒewE€

vPHSHSD v—nge˜——nwegD ƒouth efri™—D in gA lingu—lD hA ˜u™™—lD iA o™™lus—l viewF

—nd ueny— @wor—les et —lF PHHSAF huring the IWUH —nd eleph—ntid ƒtegotetr—˜elodon syrti™us @see perretti et —lF

IWVHs further field ™—mp—igns org—nized ˜y fo—z —nd PHHQ for — dis™ussionAD ˜ut no t—xonomi™ ™onsider—tion

ilEern—uti @sntern—tion—l ƒ—h—˜i ‚ese—r™h €roje™tA were of the ™—rnivore rem—ins

™—rried out ˜y —n intern—tion—l te—m of geos™ientists —nd sn IWVHD isu 8 uots—kis des™ri˜ed one of the two

IRI new fossiliferous lo™—lities were do™umented in the ™—rnivore m—ndi˜les —nd referred it to s™titherium —rkeE

—re— @he reinzelin 8 ilEern—uti IWVUAF sil—iD — new spe™ies of hy—enidF ‡erdelin 8 ƒolouni—s

sn his IWVU —n—lysis of the ƒ—h—˜i ™—rnivoresD roE @IWWIA revised hy—enid system—ti™s —nd re—ssigned the

well presented — syntheti™ des™ription without illustr—E spe™ies to ––ry—eni™titherium99 n—m—quensisD otherwise

tions of the skull —nd m—ndi˜le of the €etro™™hi ™olle™E known from ƒouth efri™— @ƒtromer IWQIY rendey IWURA

UH ƒ—rdell— ‚F 8 ‡erdelin vF

tionF re referred them to w—™h—irodus spFD together the x—w—t— pmFD voth—g—mD ueny— @™—F UFSETFS w—AF

„his spe™ies is known from — p—rti—l skeleton —nd —sE with — pQD — d€Q —nd — few post™r—ni—l ˜ones re™overed

sorted other skelet—l elementsF st is quite distin™t from in the IWVHsF sn his €hh dissert—tion ƒ—rdell— @IWWRA

other w—™h—irodontin—e known from efri™— in its deE underlined the pe™uli—r morphologi™—l fe—tures of the

rived dentition —nd slender m—ndi˜leF ƒ—h—˜i m—™h—irodontine ™—tD suggesting th—t it might

r—ileEƒel—ssie @PHHIY r—ileEƒel—ssie et —lF PHHRA ˜elong to — new spe™iesF

des™ri˜ed —s w—™h—idorus spF —n extensive m—teri—l from „he t—xonomy of xeogene m—™h—irodontine ™—ts

the v—te wio™ene eduEes— pmFD widdle ew—shD ithioE h—s ˜een dis™ussed in — series of re™ent p—pers @entoÂn et

pi— @d—ted SFVESFP w—AF „his m—teri—l in™ludes ˜oth ™r—E —lF PHHRY qer——ds et —lF PHHRY €eigne et —lF PHHSY ƒotniE

niodent—l —nd post™r—ni—l elementsD ˜ut is quite fr—gE kov— 8 xoskov— unpu˜lishedAF sn p—rti™ul—rD —n—lysis of

ment—ryD espe™i—lly the ™r—niodent—l m—teri—lF — ri™h s—mple of ™r—niom—ndi˜ul—r elements of w—™h—iE

‚e™entlyD €eigne et —lF @PHHSA des™ri˜ed m—™h—iE rodus —ph—nistus from f—t—llonesEI @ƒp—inA @ento n et —lF

rodont spe™imens from „orosEwen—ll—D gh—d @v—te PHHRA shows th—t this spe™iesD whi™h is the type spe™ies

wio™eneD —˜out the s—me —ge —s voth—g—mAD —s the of the genusD differs m—rkedly in morphology —nd —d—pE

new spe™ies w—™h—irodus k—˜irF „his spe™ies is l—rge t—tions from ––w—™h—irodus99 gig—nteusF „hese —uthors

—nd h—s — ro˜ust m—ndi˜leD quite different from th—t suggest restri™ting the use of the genus w—™h—irodus

of vokotunj—ilurusD ˜ut simil—r to the m—teri—l from ƒ—E to the type spe™ies —nd thus 99wF99 gig—nteus ™—n ˜e

h—˜i des™ri˜ed hereinF pl—™ed the genus emphim—™h—irodus uretzoiD IWPWD —s

pin—llyD rendey @IWURA des™ri˜ed —s w—™h—irodus —lso suggested ˜y fe—umont @IWUVAF

spF — ™onsider—˜le m—teri—l from v—nge˜——nwegD ƒouth

efri™— @e—rliest €lio™eneA @pigF PgEiAF „his m—teri—l reE

sem˜les vokotunj—ilurus in some respe™tsD yet differs efri™—n l—rger w—™h—irodontin—e

signifi™—ntly from it in othersF sts —ffinities rem—in to

˜e est—˜lishedF sn the s—me pu˜li™—tionD rendey deE ‡ith the re™ent remov—l of efrosmilus from the

s™ri˜ed —n upper ™—nine —s ˜elonging to romotherium pelid—e @wor—les et —lF PHHIY worlo et —lF PHHRAD the fossil

spF „his spe™imen —nd some post™r—ni—l ˜ones were reE re™ord of preE€lio™ene l—rger w—™h—irodontin—e @exE

™ently re—ssigned ˜y ‡erdelin 8 ƒ—rdell— @PHHTA to emE ™luding hinofelis —nd wet—ilurusA in efri™— h—s ˜e™ome

phim—™h—irodusF st is ™onsider—˜ly different from the very limitedF „he oldest re™ord is — left m—ndi˜ul—r r—E

m—teri—l referred to w—™h—irodus spFD m—king v—ngeE mus —nd upper ™—nine fr—gment from fled hou—r—h in

˜——nweg the only efri™—n site with rem—ins of more „unisi— @ci—rly †—llesi—nAF „hese spe™imens were reE

th—n one spe™ies of m—™h—irodont ™—t l—rger th—n hinoE ferred to w—™h—irodus ro˜insoni ˜y uurteÂn @IWUTAF „his

felisF spe™ies is of un™ert—in —ffinitiesD though it is likely to ˜e

efri™—n m—™h—irodont m—teri—l younger th—n th—t ™lose to the ™oev—l iur—si—n form wF —ph—nistusF

from v—nge˜——nweg ™—n ˜e referred to the €lioE€leistoE e m—™h—irodont spe™imenD most re™ently referred

—nd weg—ntereonF „he forE ™ene gener— romotherium to wF —ph—nistus @qer——ds et —lF PHHPA is known from

mer h—s its e—rliest —ppe—r—n™e in the vonyumun w˜F of the ghoror— pmFD ithiopi— @™—F IHFS w—AF „he spe™imen

the uoo˜i por— porm—tionD ueny—D while the e—rliest ™onsists of the symphyse—l region of — left m—ndi˜ul—r

o™™urren™e of the l—tter is from ƒouth „urkwelD ueny— r—musF

@‡erdelin 8 vewis PHHHAF „he re™ord of the former m—y

€etter @IWWRA referred —n upper ™—nine —nd right

˜e —nd the l—tter is the glo˜—lly e—rliest re™ord of the w„ sss from the xkondo pmF of the el˜ertine ‚iftD

respe™tive t—x—F

g—nd— to romotherium spF „he —ttri˜ution is m—inly

˜—sed on the simil—rity ˜etween the w„ sss —nd th—t of

romotherium from ƒeneÁze @f—llesio IWTQA —nd on the

w—teri—l —nd wethods reported presen™e of romotherium spF —t v—nge˜——nE

weg @rendey IWURAF ep—rt from the ƒ—h—˜i fossils des™ri˜ed hereinD — l—rge —mount

of ™omp—r—tive m—teri—l h—s ˜een studiedF efri™—n fossil m—™h—irodonts „hree lower ™—rn—ssi—ls of m—™h—irodonts —re

from voth—g—mD v—nge˜——nweg —nd the ‡em˜ereEw—nong— pmFD —s known from the s˜ole w˜F of the ‡em˜ereEw—nong—

well —s —ll efri™—n romotherium spF used in pigF QD h—ve ˜een studied

porm—tionD „—nz—ni— @r—rrison IWWUY v‡ person—l o˜E

˜y one of us @v‡A from origin—l spe™imensF snform—tion on spe™imens

serv—tionsAF „wo of theseD possi˜ly ˜elonging to the from fled hou—r—hD the el˜ertine ‚iftD „orosEwen—ll— —nd ghoror— h—s

s—me individu—lD from ƒhosh—m—g—i PD —re l—rge @pigF ˜een o˜t—ined from the relev—nt pu˜li™—tions @uurteÂn IWUTY €etter IWWRY

qer——ds et —lF PHHPY €eigne et —lF PHHSAF gomp—r—tive m—teri—l of emE P—EfAD —nd the thirdD from snolelo ID is ™onsider—˜ly

phim—™h—irodus gig—nteus from iur—si— h—s ˜een studied ˜y ˜oth of us sm—llerF ynly the former two —re ™onsidered hereinD —s

from origin—l spe™imens in ™olle™tions in pps—l— @€—l—eontologi™—l

the l—tter likely ˜elongs to hinofelis or — rel—ted t—xonF

hivisionD wuseum of ivolutionAD feijing @snstitute of †erte˜r—te €—E

‡erdelin @PHHQA des™ri˜ed — new genus —nd speE leontology —nd €—leo—nthropologyAD ‚ome @wuseum of €—leontologyD

v— ƒ—pienz— niversityAD ploren™e @wuseum of €—leontologyAD vondon ™ies of m—™h—irodontD vokotunj—ilurus em—geritusD from

emphim—™h—irodus from ƒ—h—˜iD vi˜y— UI

@„he x—tur—l ristory wuseumA —nd €—ris @x—tur—l ristory x—tion—l the lower m—rgin of the extern—l —uditory me—tusF

wuseumAF snform—tion on w—™h—irodus —ph—nistus from f—t—llonesEI

w—ndi˜le @pigF IeEgAF „he —lmost ™omplete left

—nd w—™h—irodus kurteni h—s ˜een o˜t—ined from the liter—ture @ƒotniE

m—ndi˜le shows eviden™e of we—theringD espe™i—lly on

kov— IWWIY ento n et —lF PHHRAF ell me—surements used in pigF Q —re

the lingu—l sideF „he in™isors —nd the lower ™—nine —re origin—l ex™ept for those on wF —ph—nistusD wF kurteniD wF ro˜insoniD

—nd wF k—˜irF l—™kingY the ™heek teethD pR —nd mI in p—rti™ul—rD —re

e˜˜revi—tions usedX uxwX ueny— x—tion—l wuseumsD x—iro˜iY he—vily wornF woreoverD the —nterior h—lf of pQ w—s

xw„X x—tion—l wuseum of „—nz—ni—D h—r is ƒ—l——mY ƒewX sziko

—™™ident—lly ˜roken in the l—te IWVHsF „he ˜ro—d we—r

ƒouth efri™—n wuseumD g—pe „ownY „wwX „ex—s wemori—l wuseumD

surf—™es on pR —nd mI indi™—te th—t the spe™imen is —n

niversity of „ex—sD eustinY fwX „he x—tur—l ristory wuseumD vonE

old individu—lF donY s†€€X snstitute of †erte˜r—te €—leontology —nd €—leo—nthropolE

ogyD feijingY €w X €—l—eontologi™—l hivisionD wuseum of ivolutionD „he horizont—l r—mus is slenderD in p—rti™ul—r —t

pps—l—Y €sxX €—leontologi™—l snstituteD wos™owY f€sX fern—rd €ri™e

the ™EpQ di—stem—F e m—rked ment—l fl—ngeD slightly d—E

snstituteD toh—nnes˜urgF

m—ged in its —nterior p—rtD is presentF tudging ˜y its

—lveolusD the lower ™—nine w—s quite l—rgeD —nd elev—ted

rel—tive to the level of the ™heek teethF iven if p—rti—lly

hes™ription of the ƒ—h—˜i m—™h—irodont m—teri—l

d—m—gedD the redu™ed size of the dou˜leErooted pQ is

evidentF rowell @IWVUA des™ri˜ed it —s followsX ––pQ is

gr—nium @pigF I—EdAF „he well preserved neuroE

present —nd it is quite wornD ˜ut the ™usps —re preservedY

™r—nium l—™ks the left zygom—ti™ —nd glenoid pro™essesD

the m—in ™usp is pointed —nd ™—nted r—ther ˜—™kw—rdD the posterior p—rt of the s—gitt—l ™rest —nd the right p—rt

—nd there —re su˜st—nti—l —nterior —nd posterior —™™esE of the o™™ipit—l ™restF „he o™™ipit—l pl—ne is verti™—lF

sory ™usps —nd — seemingly low dist—l ™ingulumF „he pQ enteriorlyD the skull is ˜roken just —˜ove the or˜itF

is quite ™losely —pproxim—ted to pRF99 „he l—m˜doid ™rest is strongly developedD while the

foth pR —nd mI —re l—rge —nd wornD espe™i—lly on tempor—l foss— is wideD verti™—lly highD —nd delimited

the lingu—l sideF xeverthelessD there is overl—p ˜etween ˜y — m—ssive zygom—ti™Eglenoid pro™essF „he glenoid

the —nterior edge of mI —nd the posterior edge of pRF sn foss— is very wide —nd deepF „he p—ro™™ipit—l pro™esses

mID the met—™onidEt—lonid ™omplex is missingF —re l—rge —nd posterol—ter—lly oriented rel—tive to the

yther dent—l rem—ins referred ˜y rowell @IWVUA m—stoid pro™essesD whi™h —re ˜ro—d —nd m—ssiveF „he

to w—™h—irodus spF —re — right pQ @IVS€PVeA from lo™—lE l—tter extend verti™—lly ˜elow —nd p—rti—lly ˜ehind the

ity €PVe —nd — d€Q from lo™—lity €PWeF ‡e h—ve not respe™tive extern—l —uditory me—tusD whi™h is lo™—ted

seen these spe™imens —nd they —re not figured ˜y rowell medi—lly ˜elow —nd ˜ehind the zygom—ti™ root —nd

@IWVUAF re doesD howeverD figure sever—l post™r—ni—l eleE —nterodors—l to the root of the m—stoid pro™essF

ments @rowell IWVUD pigF VA refer—˜le to w—™h—irodonE purthermoreD the glenoid pro™ess protrudes ventr—lly

tin—eF „hese will ˜e ™omp—red to m—teri—l from voth—E ˜elow the lower end of the m—stoid pro™essD —pproxiE

m—tely —s mu™h —s the m—stoid pro™esses extend ˜elow g—m —nd v—nge˜——nweg in — sep—r—te ™ontri˜utionF

pigF Q Efiv—ri—te di—gr—m of length

versus width of the lower

™—rn—ssi—l of some wioE€lioE

™ene m—™h—irodonts —s l—E ˜eledF

UP ƒ—rdell— ‚F 8 ‡erdelin vF

gomp—rison of the ƒ—h—˜i m—™h—irodont m—teri—l to ™imens of romotherium listed —˜ove differ in size —nd

other v—te wio™ene m—™h—irodonts in some ™r—niodent—l fe—turesD ˜ut the gener—l setting of

the m—stoid —re— is quite homogeneousF

„he emphim—™h—irodus from ƒ—h—˜i shows —dE

gomp—red to romotheriumD the ƒ—h—˜i spe™imen

v—n™ed ™h—r—™ters in its ˜—si™r—nium —nd lower ™heek

h—s — l—rger p—ro™™ipit—l pro™essD rel—ted to — powerful

teethF emong „uroli—n s—˜ertoothed ™—tsD only — few

mF dig—stri™usD — ˜ro—der m—stoid pro™essD with — wide

skulls with well preserved ˜—si™r—ni— —re knownD —nd

insertion —re— for the mF o˜liquus ™—pitis ™r—ni—lisF v—tE

some of these —re juvenile spe™imensF emong iurope—n

er—l to this —re— —re the insertions for mmF spleniusD

eF gig—nteus spe™imensD the morphology of the m—stoid

longissimus ™—pitisD —nd sternoE™eph—li™usF „he ˜r—™hioE

—nd o™™ipit—l —re— ™—n ˜e o˜served in the —lmost ™omE

™eph—li™us —nd p—rt of the sterno™eph—li™us insert on the

plete —dult skull from r—lmyropot—mos des™ri˜ed —nd

tip of the pro™essF tudging ˜y the ˜—si™r—ni—l morpholE

figured in welentis @IWUHA @—t th—t time referred to wF

ogyD these mus™les —re very strong —nd more —nteroposE

—ph—nistusA —nd in — juvenile spe™imen from ƒ—mos pu˜E

teriorly oriented in the ƒ—h—˜i spe™imen th—n in romoE

lished —nd figured ˜y fe—umont @IWUVAF purther inforE

theriumF „he o™™iput is more verti™—l th—n in romoE

m—tion on the ™r—ni—l morphology of these felids ™—n ˜e

therium —nd the other spe™ies of emphim—™h—irodusF

o˜t—ined from skulls from different lo™—lities of the forE

sn this fe—ture —nd in the set of the m—stoidEglenoid —re—D

mer ƒoviet nionD —mong them the re™ently prep—red

the ƒ—h—˜i spe™imen seems to p—r—llel the smilodontine

skull with m—ndi˜le of emphim—™h—irodus t—r—™liensis

™—tsF e gre—t development of the upper ™—nine m—y ˜e

@holotypeAD — previously unpu˜lished skull of emphiE

expe™ted in the ƒ—h—˜i spe™iesF „he m—ndi˜le from ƒ—E

m—™h—irodus ™opei @yƒ PTQVA from qre˜eniki —nd

h—˜i —lso h—s some fe—tures of interestD in™luding — rel—E

the skull with m—ndi˜le of emphim—™h—irodus kurteni

tively slender horizont—l r—musD in p—rti™ul—r —t the level

@ƒotnikov— IWWIY ƒotnikov— 8 xoskov— person—l ™omE of the ™EpQ di—stem—D — strong ment—l fl—ngeD redu™ed pQ

muni™—tionAF —nd wide we—r surf—™es on the lingu—l side of ˜oth pR

„he ghinese fossil re™ord —lso in™ludes some very @posterior h—lfA —nd mIF

well preserved ™omplete skullsD su™h —s th—t referred to „he o™™urren™e of su™h — distin™t ment—l fl—nge is

eF p—l—nderi @a eF gig—nteusA ˜y gh—ng @IWSUA —nd the v—ri—˜le in „uroli—n m—™h—irodontsF tudging ˜y the ilE

su˜—dult spe™imen from lo™F IIQ of f—odeD ƒh—nxiD ghiE lustr—tionsD it is present in the spe™imens refer—˜le to eF

n—A @d—nsky IWPRAF „he ƒ—h—˜i emphim—™h—irodus h—s gig—nteus from €ikermi —nd ƒ—mos @fe—umont IWUVAD in

—lso ˜een ™omp—red to — ™—st of — quite ™ompleteD eF gig—nteus figured ˜y gh—ng @IWSUA —nd in eF kurteni

though slightly ™rushedD skull with —rti™ul—ted m—ndi˜le @ƒotnikov— IWWIAD ˜ut not in spe™imen wQVSI from

of eF gig—nteus from —n unknown lo™—lity in ghin— f—ode lo™F QH @d—nsky IWPRAF e dire™t ™omp—rison with

@stored —t the ƒwedish wuseum of x—tur—l ristoryD the —lmost ™omplete lower j—w from lo™F IIQ referred to

ƒto™kholmAF eF p—l—nderi @a eF gig—nteusA @d—nsky IWPRY uurteÂn

sn ™omp—rison to —ll the spe™imens listed —˜oveD IWSPAD whi™h ˜elongs to — su˜—dult individu—lD —nd with

the ƒ—h—˜i m—™h—irodont shows the most derived o™™iE eF gig—nteus from —n unknown lo™—lity in ghin—D deE

pit—l —nd m—stoid —re—D with l—rger p—ro™™ipit—l pro™esses monstr—tes m—rked differen™es in the gener—l morpholE

—nd — ˜ro—der m—stoid pro™ess th—t is more ventr—lly ogy of the ™oronoid pro™ess @more redu™ed in the ƒ—h—˜i

oriented —nd is lo™—ted —pproxim—tely midw—y —long — spe™imenA —nd in the sh—pe of the m—sseteri™ foss—D

line sep—r—ting the p—ro™™ipit—l pro™ess from the glenoid whi™h is quite rounded in emphim—™h—irodus from ƒ—E

pro™essF ƒu™h fe—tures in the ƒ—h—˜i spe™imen suggest —n h—˜i —nd more tr—pezoid in the other spe™imensF „he

—d—pt—tive set ™onverging on smilodontine ™—tsF vi˜y—n s—˜ertoothed ™—t —lso h—s — more redu™ed pQF

„he —v—il—˜le d—t— indi™—te ™onsider—˜le simil—riE effinities with ––w—™h—irodus99 k—˜ir from „oros

ties ˜etween eF gig—nteus —nd romotheriumD in p—rtiE wen—ll— @€eigne et —lF PHHSA —re evidentF „he gh—d speE

™ul—r in the sh—pe —nd development of the p—ro™™ipit—l ™imen is somewh—t sm—ller th—n the vi˜y—n oneD

—nd m—stoid pro™essesF ƒu™h fe—tures ™—n ˜e seen on — though they —re ™losely simil—r in dent—l me—surementsF

well preserved —nd ™omplete skull of rF nesti—nus from „he m—ndi˜le „w PTTEHPEIHP shows — developed menE

the v—te €lio™ene of ‚o™™— xeyr— @pr—n™eA @fonis IWUTA t—l fl—nge —nd — redu™ed pQD ˜ut differs in h—ving —

—nd on — ™—st of — skull of rF ™ren—tidens from ƒeneze l—rger ™oronoid pro™ess @judging ˜y the photoA —nd

@pr—n™eA @f—llesio IWTQAF e neuro™r—nium from €irro eviden™e of — sm—ll met—™onidGt—lonid ™omplexF sn

xord @i—rly €leisto™eneD st—lyA h—s —lso ˜een used for these fe—turesD ––wF99 k—˜ir seems to ˜e slightly more

™omp—rison @ƒ—rdell— IWWRAF e det—iled study of the ˜—E primitive th—n the ƒ—h—˜i spe™imen —s would ˜efit it9s

si™r—ni—l morphology of well preserved spe™imens of somewh—t gre—ter —geF

romotherium from sn™—r™—l @i—rly €leisto™eneD ƒp—inA e very wide we—r surf—™e ™—n ˜e seen on the linE

w—s re™ently pu˜lished @ento n 8 q—lo˜—rt IWWWA —nd gu—l side of the lower ™—rn—ssi—lD —s well —s on the posE

terior ™usp of pRF ƒu™h — ™ondition is seen in the eF provides — good ™omp—r—tive s—mpleF „he different speE

emphim—™h—irodus from ƒ—h—˜iD vi˜y— UQ Catalog number Species Locality Lp3 Wp3 Lp4 Wp4 Lm1 Wm1 „—˜F I Ewe—surement d—t— for the

La Sapienza - Amphimachairodus Sahabi 18.9 6.5 21.2 11.2 36.4 14.0 lower dentition of efri™—n —nd iur—si—n m—™h—iroE cf. A. kabir donts —s used in pigF QF KNM-LT 26178 Lokotunjailurus Lothagam 8.1 5.2 17.9 8.5 27.8 10.9 emageritus T-491 Machairodus Bled Douarah 25.0 11.5 robinsoni NMT WM Amphimachairodus Shoshagamai 2 29.0 13.3 1937/92a sp. NMT WM Amphimachairodus Shoshagamai 2 30.8 13.4 1937/92b sp. SAM-PQ L20505 "Machairodus" sp. Langebaanweg 28.8 10.7 TMM 266-02- Amphimachairodus Toros-Menalla 34.7 13.7 102 kabir BM M 48437 Amphimachairodus Siwaliks 24.5 29.0 13.8 giganteus Composite, from Machairodus Batallones 1 26.3 10.2 30.5 13.6 Antón et al. 2004 aphanistus IVPP V.906 Amphimachairodus Yushe 21.0 10.0 29.0 12.0 33.0 14.0 giganteus PMU – (Ex2) Amphimachairodus Loc 113, Baode 15.6 7.4 26.1 10.3 28.1 12.6 giganteus PIN 2433/287 Amphimachairodus Kalmakpai 26.8 11.5 31.2 13.0 kurteni PIN 2433/524 Amphimachairodus Kalmakpai 17.3 7.9 25.3 11.1 31.4 14.0 kurteni BPI M8280 Homotherium Makapansgat 34.0 12.0 problematicum KNM-ER 931 Homotherium sp. Koobi Fora, KBS 36.1 13.7 Mb.

KNM-KP 30420 Homotherium sp. Kanapoi 32.0 12.5

—˜le is the mIF sn pigF QD the length —nd width of the gig—nteus ™omplexD in™luding the derived eF kurteniD

—v—il—˜le lower ™—rn—ssi—ls —re ™omp—red to lower ™—rE —nd is typi™—l of the genus romotheriumF

n—ssi—ls of some iur—si—n wio™ene l—rge m—™h—irodontsD xeverthelessD the —dv—n™ed fe—tures of the ˜—siE

—s well —s some efri™—n romotherium spF @„—˜F IAF ™r—ni—l morphology in the ƒ—h—˜i spe™imen ™omp—red

prom this it is evidentD first of —ll th—t the efri™—n wioE to the less derived ™h—r—™ters o™™urring in romotheE

™ene m—™h—irodont m—teri—l ™—n ˜e sep—r—ted into two rium suggest th—t the vi˜y—n t—xon should ˜e ™onsidE

size ™l—ssesF e group of sm—ll forms in™ludes wF ro˜inE ered — very —dv—n™ed form in the evolution—ry trend of

soniD vF em—geritus —nd ––wF99 spF from v—nge˜——nweg @it the eF gig—nteus groupD ˜ut not the dire™t —n™estor of

should ˜e noted th—t the mI from snolelo ID ‡em˜ereE romotheriumF

w—nong— pm —lluded to —˜ove is ™onsider—˜ly sm—ller ƒimil—r ™onsider—tions h—ve ˜een pointed out —lso

th—n this groupAD while — group of l—rger forms in™ludes for emphim—™h—irodus kurteni @ƒotnikov— IWWIAD whi™h

––wF99 k—˜irD emphim—™h—irodus spF from ƒhosh—m—g—i PD sh—res with the ƒ—h—˜i spe™imen — distin™t ment—l —poE

—nd the ƒ—h—˜i spe™imen des™ri˜ed —˜oveF physis —nd —˜sen™e of the met—™onidGt—lonid ™omplexF

‡ithin e—™h of these groups there —re differen™es

˜etween spe™imensD however —nd these differen™es ™—n

˜e rel—ted to simil—r differen™es in the ™omp—r—tive m—E

‚e—ssessment of efri™—n v—te wio™ene m—™h—irodonts

teri—lF sn the group of sm—ll spe™imensD wF ro˜insoni h—s

„he most ™ommonly preserved element in the — ˜ro—d mID while the mu™h younger voth—g—m —nd

efri™—n m—™h—irodont m—teri—l th—t is ™urrently —v—ilE v—nge˜——nweg spe™imens h—ve slender lower ™—rn—sE

UR ƒ—rdell— ‚F 8 ‡erdelin vF

spe™imens dis™ussed —˜oveD ˜ut in the —˜sen™e of dent—l si—lsF st is tempting to suggest th—t the former is ˜ro—d

˜e™—use it is older —nd more primitiveD sin™e there is — m—teri—l other th—n the upper ™—nine it ™—nnot ˜e deterE

trend —t le—st in l—ter m—™h—irodonts @see ˜elowA for the mined if it ˜elongs with the ˜ro—d lower ™—rn—ssi—l form

mI to ˜e™ome progressively more slenderF „his tends to represented ˜y the ƒhosh—m—g—i P spe™imens or the

˜e ™onfirmed ˜y the ˜ro—d mI in the simil—rly —ged wF slender lower ™—rn—ssi—l form represented ˜y the m—teE

—ph—nistus @in the plot this spe™ies is represented ˜y the ri—l from vi˜y— —nd gh—dF

me—n of the f—t—llonesEI s—mple from ento n et —lF pin—llyD r—ileEƒel—ssie et —lF @PHHRA note th—t the

PHHRAF „he simil—rity ˜etween the voth—g—m —nd v—nE ––w—™h—irodus99 spF m—teri—l from the eduEes— pmF ––is

ge˜——nweg forms lends ™reden™e to suggestions th—t l—rger in size ‘th—n vF em—geritus from voth—g—mD iFeFD

they —re ™losely rel—tedD though differen™es in other the size of wF gig—nteus“D with — shorter —nd wider mI99F

dent—l —nd ™r—ni—l proportionsD —s well —s in skelet—l qiven the length of mI st—ted in r—ileEƒel—ssie @PHHIAD

fe—tures pre™lude spe™ifi™ identityF guriouslyD the older we —ssume the l—tter to me—n rel—tively shorter —nd

voth—g—m form —ppe—rs more derived th—n the younger widerF „hese st—tements suggest th—t the eduEes— pmF

v—nge˜——nweg formD ˜ut —g—inD this requires further mI is most simil—r to the two lower ™—rn—ssi—ls from

—n—lysisF ƒhosh—m—g—i PF gle—rlyD the eduEes— pmF spe™imens

sn the group of l—rger forms there —re —lso spe™iE should —t le—st provision—lly ˜e referred to —n emphiE

mens with ˜ro—d lower ™—rn—ssi—ls —nd spe™imens with m—™h—irodus spF simil—r to th—t from ƒhosh—m—g—i P ˜ut

more slender onesF sn the first group we h—ve the two different from the vi˜y—Ggh—d spe™iesF

spe™imens from ƒhosh—m—g—i P —nd in the l—tter group

we h—ve ––wF99 k—˜ir —nd the ƒ—h—˜i spe™imenF st should

˜e emph—sized th—t the differen™e ˜etween these spe™iE gon™lusions

mens in mI width h—s no o˜vious str—tigr—phi™ ™ompoE

„he verte˜r—te f—un— of ƒ—h—˜i h—s ˜een v—rE nentD —s —ll spe™imens —re ˜etween S —nd U million ye—rs

iously interpreted —s ˜eing of l—test wio™ene @on the oldF snste—dD the differen™es m—y refle™t — t—xonomi™

˜io™hronology of its f—un—l elementsA or ˜—s—l €lio™ene distin™tion —nd differenti—l history of the t—x— involvedF

—ge @˜—sed on region—l geology ™onsider—tionsAF et „he ƒhosh—m—g—i P spe™imens m—t™h iur—si—n eF gig—nE

presentD most —uthors seem to ™onsider ƒ—h—˜i to lie teus ™losely in mI proportionsD while the other two

very ne—r the wio™eneE€lio™ene tr—nsition @wx IQAF spe™imens —ppro—™h romotherium spF in proportionsF

€—leoe™ologi™—l studies suggest th—t —t ƒ—h—˜i wooded st thus seems pl—usi˜le to suggest th—tD given the simil—r

h—˜it—ts o™™urred —long —dj—™ent ˜—nks of — l—rge river —ges of the spe™imensD the more slender vi˜y—n —nd

™ontr—sting with semi—rid ™onditions —w—y from the gh—di—n spe™imens either represent — se™ond—ry immiE

riverF ƒu™h ™onditions pro˜—˜ly ˜e™—me intensified gr—tion of emphim—™h—irodus into efri™— or —re des™enE

during — well m—rked dry se—son @fo—z IWVUAF emphiE d—nts of —n efri™—n form simil—r to th—t represented in

m—™h—irodus w—s the l—rgest spe™i—list pred—tor in the the m—teri—l ˜y the spe™imens from ƒhosh—m—g—i PF f—E

ƒ—h—˜i f—un—D with —n —d—pt—tion for killing l—rge prey si™r—ni—l morphology seems to pre™lude the presen™e of

@young pro˜os™ide—ns or rhinosD hipposD —ntr—™otheresD the ƒ—h—˜i form in the dire™t —n™estry of romotherium

et™FAF whose first @glo˜—lA o™™urren™e is in the vonyumun

„he ƒ—h—˜i emphim—™h—irodusD with its strongly wem˜er of the uoo˜i por— porm—tionD d—ted RFQSERFI

developed m—stoid —nd glenoid pro™essesD —nd redu™ed w— @‡erdelin 8 vewis PHHSAD ˜ut the group th—t did

pQ —nd ™oronoid pro™ess ™—n ˜e ™onsidered —s — derived give rise to romotherium must surely ˜e very ™lose in

form within the p—r—Eor polyphyleti™ emphim—™h—iroE time —nd morphology to the ƒ—h—˜i formF por t—xoE

dus spe™ies ™omplexF sn some ™r—ni—l fe—tures it ™onE nomi™ purposes it is —lso quite ™le—r th—t the gh—d form

verges on smilodontine ™—tsF „he vi˜y—n spe™ies seems must ˜e referred to emphim—™h—irodusF

too derived to ˜e ™onsidered —n™estr—l to the genus yf m—teri—l th—t is not in™luded in the dis™ussion

romotheriumF gomp—red to other xeogene efri™—n —˜oveD the spe™imen from the ghoror— pmF is spe™ifiE

spe™imens the ƒ—h—˜i form differs gre—tly in ˜oth size ™—lly indetermin—teD ˜ut ˜ro—dly simil—r to the wF —ph—E

—nd morphology from vokotunj—ilurus em—geritus of nistusGwF ro˜insoni groupF „he m—teri—l from the xkonE

voth—g—m @ueny—AD while it shows ™le—r —ffinities with do pmF h—s not ˜een —v—il—˜le for studyF roweverD the

the l—rge emphim—™h—irodus k—˜ir from „oros wen—ll— dimensions —nd morphology of the spe™imensD —s well —s

@gh—dAF „he l—tter seems more primitive in h—ving — the illustr—tions in €etter @IWWRA provide no indi™—tion

more developed ™oronoid pro™ess —nd ret—ining — met—E th—t this m—teri—l ™ould not ˜e referred to emphim—E

™onidGt—lonid ™omplex on mIF st seems pro˜—˜le th—t ™h—irodus spFD whi™h would ˜e more in line with its v—te

the two forms —re rel—tedD —nd should ˜e pl—™ed in — wio™ene —geF es we h—ve shown elsewhere @‡erdelin 8

ƒ—rdell— PHHTAD the l—rger m—™h—irodont from v—ngeE different t—xonomi™ group th—n the iur—si—n l—rge eF

˜——nweg would fit ™omfort—˜ly in the group of l—rger gig—nteusF ‡hether the „orosEwen—ll— —nd ƒ—h—˜i spe™iE

emphim—™h—irodus from ƒ—h—˜iD vi˜y— US

e™knowledgementsF ‡e would like to th—nk —ll those museum mens represent the s—me spe™ies is not possi˜le to deE

™ur—tors who h—ve provided —™™ess to spe™imensD espe™i—lly ƒolweig termine with ™ert—inty on the ˜—sis of the m—teri—l —v—ilE

ƒtuenesD pps—l—D ilis—˜ett— gioppiD pirenzeD w—rg—ret every —nd

—˜leD ˜ut indi™—tions —re th—t the differen™es present ™—n

herek yl—ndD g—pe „ownD the government of the ‚epu˜li™ of ueny—D

˜e —s™ri˜ed to ontogenyD interspe™ifi™ v—ri—tion —nd str—E

we—ve ve—keyD imm— w˜u—D predri™k uy—lo w—nthiD w—ry wuungu

tigr—phi™ —geF „orosEwen—ll— —nd ƒ—h—˜i —re not very —nd m—ny others for permissions —nd —ssist—n™e in the study of m—E

different in —ge in view of the o™™urren™e of — simil—r teri—l housed in x—iro˜iD —nd gyƒ„igrD „erry r—rrison —nd em—nE

dus uwek—son for permissions —nd —ssist—n™e with the study of m—E f—un—l ™omplex in ˜oth lo™—lities @eFgF the —ntr—™othere

teri—l housed in h—r is ƒ—l——mF ‡e th—nk the referees vorenzo ‚ook

vy˜i™os—urus petro™™hiiA @†ign—ud et —lF PHHPAD ˜ut overE

—nd w—nuel ƒ—les— for ™riti™—l re—ding the m—nus™ript —nd the useful

—ll the „orosEwen—ll— f—un— must ˜e ™onsidered someE

suggestionsF ‡e —lso th—nk w—rin— †F ƒotnikov— for —™™ess to —n unE

wh—t older th—n th—t of ƒ—h—˜iF ‡e provision—lly refer

pu˜lished m—nus™riptD ƒteph—ne €eigne for dis™ussions —˜out eF k—˜irD

the ƒ—h—˜i spe™imens to emphim—™h—irodus —ffF eF k—E vu™— fellu™™i —nd elessi— gipullo for te™hni™—l —ssist—n™eF ‚ƒ —™E

˜irF knowledges funding through the ––righ v—t99 progr—m whi™h w—s

m—de —v—il—˜le ˜y the iurope—n gommunity Ee™™ess to ‚ese—r™h „he presen™e of — derived emphim—™h—irodus —t

snfr—stru™ture —™tion of the smproving rum—n €otenti—l €rogr—mD v—nge˜——nweg @e—rliest €lio™eneD ƒouth efri™—A @‡erdeE

—s well —s through ƒ‰x„riƒ‰ƒD proje™t ƒi„epEIHRPD whi™h w—s

lin 8 ƒ—rdell— PHHTA —nd the ‡em˜ereEw—nong— pmF

m—de —v—il—˜le ˜y the iurope—n gommunity E‚ese—r™h snfr—stru™E

@l—test wio™eneD „—nz—ni—A suggests th—t the evolution

ture e™tion under the p€T ––ƒtru™turing the iurope—n ‚ese—r™h ere—99

of s—˜ertoothed ™—ts in efri™— —t the wio™eneE€lio™ene €rogr—mF v‡ gr—tefully —™knowledges funding from the ƒwedish ‚eE

tr—nsition is more ™omplex th—n previously envis—gedF se—r™h goun™ilD †‚F

‚ipi‚ixgiƒ

egustõÁ tFD ilEern—uti eFD woy—Á ƒol—Á ƒFD uohler wFD q—lo˜—rt the ƒ—h—˜i hipp—rons @l—test wio™eneA @vi˜y—AF qeodiE È

eFD q—ete ‚F 8 vlen—s wF @PHHHA E‚esults of — fieldE versit—sD PSX PWUEQIWD €—risF

™—mp—ign in the v—te wio™ene of the ƒ—h—˜i porm—E fo—z hF hF @IWVUA E„—phonomy —nd p—leoe™ology —t the

tion @vy˜i—AF e˜str—™ts ˆs ‚gwxƒ gongressD pez €lio™ene site of ƒ—h—˜iD vi˜y—F snX fo—z xF „FD ilE

@woro™™oAD IX SWF ern—uti eFD q—ziry eF ‡FD he reinzelin tFD 8 fo—z

ento n wF 8 q—lo˜—rtF eF @IWWWA Exe™k fun™tion —nd preE hF hF @idsA E xeogene €—leontology —nd qeology of

d—tory ˜eh—vior in the s™imit—r toothed ™—t romoE ƒ—h—˜iX QQUEQRVD xew ‰orkF

therium l—tidens @ywenAF tF †ertF €—leontFD IWX UUIE fonis de vF @IWUTA E n feÂlide — longues ™—nines de l— ™olline

UVRD xorth˜rookD v—wren™eF de €errier @€uyEdeEhoà meAX ses r—pports —ve™ les feÂE

ento n wFD ƒ—les— wF tFD wor—les tFD 8 „urner eF @PHHRA E lineÂs m—™h—irodontesF ennF €—leontFD TPXISWEIWVD Â

pirst known ™omplete skulls of the s™imit—rEtoothed €—risF

™—t w—™h—irodus —ph—nistus @pelid—eD g—rnivor—A from gh—ng rF @IWSUA Eyn new m—teri—l of some m—™h—irodonts

the ƒp—nish l—te wio™ene site of f—t—llonesEIF tF †ertF of €onti—n —ge from ƒh—nsiF †ertF €—lesi—tFD IX IWQE

€—leontFD PRX WSUEWTWD v—wren™eF PHHD feijingF

f—llesio ‚F @IWTQA Ewonogr—phie d9un m—™h—irodus du giE gooke rF fF ƒF @IWVUA Epossil ƒuid—e from ƒ—h—˜iD vi˜y—F snX

sement †ill—fr—n™hien de ƒeneÁzeX romotherium ™reE fo—z xF „FD ilEern—uti eFD q—ziry eF ‡FD he reinE

n—tidens p—˜riniF „r—v—ux du v—˜or—toire de qeologie zelin tFD 8 fo—z hF hF @idsA E xeogene €—leontology Â

X QUERPD xew ‰orkF de l— p—™ulte des ƒ™ien™es de vyonD WX IEIPWD vyonF —nd qeology of ƒ—h—˜i Â

he reinzelin tF 8 ilEern—uti eF @IWVUA E „he ƒ—h—˜i porE f—llm—nn €F @IWVUA Ee fossil ˜ird f—un— from the €lio™ene

m—tion —nd rel—ted depositsF snX fo—z xF „FD ilEerE ƒ—h—˜i porm—tion of vi˜y—F snX fo—z xF „FD ilEern—uE

n—uti eFD q—ziry eF ‡FD he reinzelin tFD 8 fo—z hF ti eFD q—ziry eF ‡FD he reinzelin tFD 8 fo—z hF hF

hF @idsA E xeogene €—leontology —nd qeology of ƒ—E @idsA E xeogene €—leontology —nd qeology of ƒ—h—˜iX

h—˜iX IEPPD xew ‰orkF IIQEIIVD xew ‰orkF

isu hFD 8 uots—kis „F @IWVHA E s™titherium —rkesil—i spF nF fe—umont de qF @IWUVA Exotes ™ompleÂment—ires sur quelE

@ry—enid—eD g—rnivor—A del „erzi—rio superiore di ƒ—E ques feÂlideÂs @g—rnivoresAF er™hives des ƒ™ien™esD QIX

h—˜i @giren—i™—D vi˜i—AF ‚ivF stF €—leontF ƒtr—tFD VTX PIWEPPUD qeneÁveF

PRIEPSTD wil—noF fernor ‚F vF 8 €—vl—kis €F €F @IWVUA Eoogeogr—phi™ rel—E

perretti wF€FD ‚ook vF 8 „orre hF @PHHQA E ƒtegotetr—˜eloE tionships of the ƒ—h—˜i l—rge m—mm—l f—un—F snX fo—z

don ™fF syrti™us @€ro˜os™ide—D ileph—ntid—eA from the xF „FD ilEern—uti eFD q—ziry eF ‡FD he reinzelin tFD

pper wio™ene of gess—niti @g—l—˜ri—D southern st—lyA 8 fo—z hF hF @idsA E xeogene €—leontology —nd

—nd its ˜e—ring on v—te wio™ene p—leogeogr—phy of qeology of ƒ—h—˜iX QRWEQVRD xew ‰orkF

™entr—l wediterr—ne—nF tF †ertF €—leontFD PQ@QAX TSWE fernor ‚F vF 8 ƒ™ott ‚F ƒF @PHHQA Exew interpret—tions of

TTTD v—wren™eF the system—ti™sD ˜iogeogr—phy —nd p—leoe™ology of

UT ƒ—rdell— ‚F 8 ‡erdelin vF

„FD ilEern—uti eFD q—ziry eF ‡FD he reinzelin tFD 8 q—ziry eF ‡F @IWVU—A E‚em—ins of €ro˜os™ide— from the

fo—z hF hF @idsA E xeogene €—leontology —nd qeolE e—rly €lio™ene of ƒ—h—˜iD vi˜y—F snX fo—z xF „FD ilE

ogy of ƒ—h—˜iX ISQEIVID sn™FD xew ‰orkF ern—uti eFD q—ziry eF ‡FD he reinzelin tFD 8 fo—z hF

uretzoiD wF @IWPWA EpelidenEƒtudienF e w—gy—r uir— hF @idsA E xeogene €—leontology —nd qeology of ƒ—E lyi Â

h—˜iX IVQEPHRD xew ‰orkF poldt—ni sntezet r—zinyomd—j—D PRXIEPPD fud—pestF  È

q—ziry eF ‡F @IWVU˜A E wery™opot—mus petro™™hii @ertiod—™E uurteÂn fF @IWSPA E„he ghinese ripp—rion f—un—F gommenE

tyl—D w—mm—li—A from ƒ—h—˜iD vi˜y—F snX fo—z xF „FD t—tF ˜iolF IQXIEVPD relsinkiF

ilEern—uti eFD q—ziry eF ‡FD he reinzelin tFD 8 fo—z uurteÂnD fF @IWUTA Epossil g—rnivor— from the v—te „erti—ry

hF hF @idsA E xeogene €—leontology —nd qeology of of fled hou—r—h —nd gheri™hir—D „unisi—F xotes du

ƒ—h—˜iX PVUEQHPD xew ‰orkF ƒervi™e qeologique de „unisieD RPX IUUEPIRD „unisF Â

q—ziry eF ‡F @IWVU™A E rex—protodon s—h—˜iensis @ertiod—™E vehm—nnD F 8 „hom—s rF @IWVUA Epossil fovid—e @w—mE

tyl—D w—mm—li—AX e new hippopot—mus from vi˜y—F m—li—A from the wioE€lio™ene of ƒ—h—˜iD vi˜y—F snX

snX fo—z xF „FD ilEern—uti eFD q—ziry eF ‡FD he fo—z xF „FD ilEern—uti eFD q—ziry eF ‡FD he reinE

reinzelin tFD 8 fo—z hF hF @idsA E xeogene €—leonE zelin tFD 8 fo—z hF hF @idsA E xeogene €—leontology

tology —nd qeology of ƒ—h—˜iX QHQEQITD xew ‰orkF —nd qeology of ƒ—h—˜iX QPQEQQTD xew ‰orkF

qer——ds hF @IWWVA Efiogeogr—phy of ™ir™umEmediterr—ne—n welentis tF uF @IWUHA Eƒtudien u ˜er possile †erte˜r—ten È

wio™eneE€lio™ene rodentsX — revision using f—™tor qrie™henl—ndsF IWF hie €ikermif—un— von r—lmyroE

—n—lysis —nd p—rsimony —n—lysis of endemi™ityF €—E pot—mos @iu˜o— Eqrie™henl—ndAF I „eilX ydontologie È

l—eogeogrFD €—l—eo™lim—tolFD €—l—eoe™olFD IQUX PUQEPVVD und ur—niologieF ennF qeolF €—ys rellenF IWX PVSE  Â

emsterd—mF RIID ƒ—loniqueD ethensF

qer——ds hFD elemseged F 8 fellon rF @PHHPA E„he v—te wor—les tFD €i™kford wF 8 ƒori— hF @PHHSA Eg—rnivores from

wio™ene m—mm—li—n f—un— of ghoror—D ew—sh f—sinD the v—te wio™ene —nd ˜—s—l €lio™ene of the „ugen

ithiopi—X system—ti™sD ˜io™hronology —nd the RHuE rillsD ueny—F ‚evF ƒo™F qeolF isp—n—D IVX QWETID w—E Ä

RHer —ges of the —sso™i—ted vol™—ni™sF „erti—ry ‚eE dridF

se—r™hD PIXIIQEIPPD vondonF wor—les tFD ƒ—les— wFD €i™kford wF 8 ƒori— hF @PHHIA Ee new

qer——ds hFD u—y— „F 8 „un— †F @PHHRA Ee skull of w—™h—iE tri˜eD new genus —nd two new spe™ies of f—r˜ourofeE

rodus gig—nteus @pelid—eD w—mm—li—A from the v—te lin—e @pelid—eD g—rnivor—D w—mm—li—A from the i—rly

wio™ene of „urkeyF xeues t—hr˜F qeolF €—l—ontFD wFD wio™ene of i—st efri™— —nd ƒp—inF „r—nsF ‚oyF ƒo™F È

PHHRX WSEIIHD ƒtuttg—rtF idin˜urghD i—rth ƒ™ien™esD WPX WUEIHPD idin˜urghF

r—ileEƒel—ssie ‰F @PHHIA E v—te wio™ene w—mm—li—n p—un— worlo wFD €eigne ƒF 8 x—gel hF @PHHRA Ee new spe™ies of

from the widdle ew—sh †—lleyD ithiopi—F †F of RPS €ros—ns—nosmilusX impli™—tions for the system—ti™ reE

ppFD €hF hF thesisD hep—rtment of sntegr—tive fiologyF l—tionships of the f—mily f—r˜ourofelid—e new r—nk

niversity of g—liforni—D ferkeleyF @g—rnivor—D w—mm—li—AF oolF tF vinnF ƒo™FD IRHXRQE

r—ileEƒel—ssie ‰FD ‡oldeq—˜riel qFD ‡hite „F hFD fernor ‚F TID vondonF

vFD hegust— hFD ‚enne €F ‚FD r—rt ‡F uFD †r˜— iFD wunthe tF @IWVUA Eƒm—llEm—mm—lfossils from the €lio™ene

em˜rose ƒF 8 rowell pF gF @PHHRA EwioE€lio™ene ƒ—h—˜i porm—tion of vi˜y—F snX fo—z xF „FD ilEern—uE

m—mm—ls from the widdle ew—shD ithiopi—F qeo˜iosD ti eFD q—ziry eF ‡FD he reinzelin tFD 8 fo—z hF hF

QUX SQTESSPD €—risF @idsA E xeogene €—leontology —nd qeology of ƒ—h—˜iX

r—rris tF wF @IWVUA Epossil qir—ffid—e from ƒ—h—˜iD vi˜y—F PVQEPWUD xew ‰orkF

snX fo—z xF „FD ilEern—uti eFD q—ziry eF ‡FD he €eigne ƒFD de fonis vFD vikius eFD w—™k—ye rF „FD †ign—ud €FD

reinzelin tF 8 fo—z hF hF @idsA E xeogene €—leonE 8 frunet wF @PHHSA Ee new m—™h—irodontine @g—rE

tology —nd qeology of ƒ—h—˜iX QIUEQPPD xew ‰orkF nivor—D pelid—eA from the v—te wio™ene hominid loE

r—rrison „F @IWWUA Exeogene €—leontology of the w—nong— ™—lity of „wPTTD „orosEwen—ll—D gh—dF gF ‚F €—leE

†—lleyD „—nz—ni—X e ‡indow into the ivolution—ry volFD RX PRQEPSQD €—risF

ristory of i—st efri™—F snX v—ndm—n xF rF 8 tones €etro™™hi gF @IWRIA Esl gi—™imento fossilifero di ƒ—h—˜iF follF

hF ƒF @idsA E „opi™s in qeo˜iologyX IERIVD xew ‰orkF ƒo™F qeolF stFD THX IHUEIIRD ‚om—F

reissig uF @IWWTA E„he str—tigr—phi™—l r—nge of fossil rhinoE €etro™™hi gF @IWRQA Esl gi—™imento fossilifero di ƒ—h—˜iF

™eroses in the v—te xeogene of iurope —nd the e—stE gollF ƒ™iF ho™umentF winF efrF st—lFD IPX IEITPD ‚om—F

ern wediterr—ne—nF snX fernor ‚F vFD p—hl˜us™h †FD 8 €etro™™hi gF @IWSIA Exote sull— f—un— terzi—ri— di ƒ—h—˜iF

wittm—nn rFE‡F @idsA E „he ivolution of ‡estern ‚el—zioni dell— ƒo™iet—Á st—li—n— per il €rogresso delle

iur—si—n xeogene w—mm—l p—un—sX QQWEQRUD xew ƒ™ienzeD RPX RUWERVID ‚om—F

‰orkF €etter qF @IWWRA Eg—rnivores des reÂgions de xkondo et de

rendey F fF @IWURA E„he l—te genozoi™ g—rnivor— of the uisegiExy—˜usosi @‚ift o™™ident—lD yug—nd—AF snX fF

southEwestern g—pe €rovin™eF ennF ƒouth efri™—n ƒenut 8 €i™kford wF @idsA E qeology —nd €—l—eo˜iolE

wusFD TQXIEQTWD g—pe „ownF ogy of the el˜ertine ‚ift †—lleyD g—nd—E—ireF†FssX

rowell pF gF @IWVPA E€relimin—ry o˜serv—tions on g—rniE €—l—eo˜iologyX PHUEPITD gspiqD yrle—nsF

vor— from the ƒ—h—˜i porm—tion @vi˜y—AF q—ryounis ‚ook vF 8 w—rtinez x—v—rro fF @PHHRA E †iverr— howelli nF

ƒ™iF fullF ƒpe™F sssueX RWETID fengh—ziF spFD — new viverrid @g—rnivor—D w—mm—li—A from the

rowell pF gF @IWVUA E€relimin—ry o˜serv—tions on g—rniE f—™™inelloEginigi—no ˜—sin @l—test wio™eneD st—lyAF

vor— from the ƒ—h—˜i porm—tion @vi˜y—AF snX fo—z xF ‚ivF stF €—leontF ƒtr—tFD IIH@QAX UIWEUPQD wil—noF

emphim—™h—irodus from ƒ—h—˜iD vi˜y— UU

wio™ene „orosEwen—ll— hominid lo™—lityD gh—dF x—tE ƒ—rdell— ‚F @IWWRA Eƒistem—ti™— e histri˜uzione ƒtr—tigr—fi™—

dei w—™—irodontini d—l wio™ene ƒuperiore —l €leistoE ureD RIVX ISPEISSD vondonF

™eneF †F of IQU ppF €hh dissert—tion in €—l—eontoloE ‡erdelin vF @PHHQA EwioE€lio™eneg—rnivor— from voth—E

gi™—l s™ien™esD niversities of woden—D fologn—D g—mD ueny—F snX ve—key wF qF 8 r—rris tF hF @idsA E

pirenze —nd ‚om— ––v— ƒ—pienz—99F voth—g—mX h—wn of hum—nity in e—stern efri™—D PTIE

ƒotnikov— wF †F @IWWIA Ee new spe™ies of w—™h—irodus from QPVD xew ‰orkF

the l—te wio™ene u—lm—kp—i lo™—lity in e—stern u—E

‡erdelin vF 8 vewis wF iF @PHHHA Eg—rnivor— from the

z—khst—n @ ƒƒ‚AF snX porsteÂn eFD portelius wF 8 ‡erE

ƒouth „urkwel hominid siteD northern ueny—F tF €—E

delin vF @idsA E fjorn uurten E— memori—l volume X  È

leontFD URX IIUQEIIVHD sow— gityF

QTIEQTWD relsinkiF

‡erdelin vF 8 vewis wF iF @PHHSA E€lioE€leisto™eneg—rniE

ƒtromer iF @IWQIA E‚este ƒu ssw—sser und v—nd ˜ewohnenE È

vor— of e—stern efri™—X spe™ies ri™hness —nd turnover

der ‡ir˜eltiere —us den hi—m—ntfeldern uleinEx—m—E

p—tternsF oolF tF vinnF ƒo™FD IRRX IPIEIRRD vondonF

qu—l—ndes @ƒudwest—frik—AF ƒitzungs˜eri™hte der

È

‡erdelin vF 8 ƒ—rdell— ‚F @PHHTA E„he –– romotherium99

m—them—tis™hEn—turwissens™h—ftli™hen e˜teilung der

from v—nge˜——nwegD ƒouth efri™— —nd the origin of

f—yeris™hen ek—demie der ‡issens™h—ften zu wun™hE È

romotheriumF €—l—eontogr—phi™—D PUU@IETAX IPQEIQHD

enD IWQHX IUERUD wun™henF È

ƒtuttg—rtF

†ign—ud €FD huringer €FD w—™k—ye rF „FD vikius eFD flondel

‡erdelin vF 8 ƒolouni—s xF @IWWIA E„he ry—enid—eX t—xE

gFD foisserie tFE‚FD de fonis vFD iisenm—nn †FD itienne

onomyD system—ti™s —nd evolutionF possils —nd ƒtr—t—D wFEiFD qer——ds hFD quy pFD vehm—nn „FD vihore—u pFD

QHX IEIHRD ysloF vopezEw—rtinez xFD wourerEgh—uvire gFD ytero yFD

d—nsky yF @IWPRA Etungterti— re g—rnivoren ghin—sF €—E È ‚—ge tFEgFD ƒ™huster wFD †iriot vFD —zzo eF 8 frunet

leontF sinFD serF g P@IAX IEIRWD fejingF wF @PHHPA Eqeology —nd p—leontology of the pper