‚ivist— st—li—n— di €—leontologi— e ƒtr—tigr—fi— volume IIU noF P ppF QHWEQPI tuly PHII

„ri ps‚ƒ„ ‚igy‚h yp e xy„yƒ grsex g‚ygyh‰vspy‚w p‚yw s„ev‰

I P

pefsy wF hevve †iggrse 8 exh‚ie ge

‚e™eivedX he™em˜er PWD PHIHY —™™eptedX epril TD PHII

uey wordsX iphodont toothD gro™odyliformesD xotosu™hi—D s™ri˜edF „he tooth is stored in the ™olle™tions of the

hor—todonD er—ripesu™hus wegeneriD pper gret—™eousD €ol—zzo fossil

€—l—eontologi™—l ƒe™tion of the wuseo dell— ‚o™™— di siteD u—rst €l—te—uF

wonf—l™one with the spe™imen num˜er w€gw IIUPH

@the origin—lfieldnum˜er w—s ƒWUGIIVAF e˜str—™tF e serr—ted tooth from the goni—™i—nEƒ—ntoni—n

@ pper gret—™eousA €ol—zzo fossil site @u—rstD xi st—lyA is the first „he —im of this work is to des™ri˜e the spe™imen

re™ord of — notosu™hi—n ™ro™odyliform from st—lyF elthough it sh—res

in det—il—nd dis™uss its —ffinityF ist—˜lishingthe t—xoE

syn—pomorphies with teeth referred to the iurope—n genus hor—todon

nomi™ st—tus of w€gw IIUPH is ne™ess—ryD —s there h—ve

—nd with the qondw—n—n genus er—ripesu™husD it is distin™t in the

˜een suggestions th—t the tooth ™ould in f—™t ˜elong to — unusu—l™om˜in—tion of fe—turesD suggesting the presen™e of — yet unE

reported notosu™hi—n t—xon in the edri—ti™Ehin—ri™ g—r˜on—te €l—tE theropod dinos—ur @€F gurrieD persF ™ommFAF es dinos—ur

form lo™—ted in the „ethys ˜etween the efro—r—˜i—n ™ontinent —nd

eviden™e is r—re in st—lyD it would ˜e worth of noteF sn the xorth iurope—n l—ndm—ss during v—te gret—™eous timesF xotosuE

the me—ntimeD the tooth h—s ˜een re™ently mentioned in

™hi—ns were typi™—lly terrestri—l ™ro™odyliformsD supporting the preE

liter—ture —s eviden™e of the genus hor—todon in the sen™e of emergent —re—s on the ™—r˜on—te pl—tformF

pper gret—™eous of st—ly @helfino PHHIY ‚—˜i PHHVAD

‚i—ssuntoF n dente munito di ™—rene denti™ol—te rinvenuto nel

with potenti—l p—leogeogr—phi™ impli™—tionsD despite the sito fossilifero di €ol—zzo @g—rsoD st—li— nordorient—leA e d—t—˜ile —l

f—™t th—t its identifi™—tion in previous p—pers w—s ™le—rly goni—™i—noEƒ—ntoni—no @gret—™i™o ƒuperioreA eÁl— prim— testimoni—nz—

tent—tive —nd prelimin—ryF di un ™o™™odrilliforme notosu™o in st—li—F ƒe˜˜ene ™ondivid— sin—poE

morfie ™on denti —ttri˜uiti —lgenere europeo hor—todon e ™on ilgondE

w—ni—no er—ripesu™husD esso si distingue per l9inusu—le ™om˜in—zione di

™—r—tteriD ™he suggeris™e l— presenz— di un— spe™ie di notosu™o non

„erminology —nd methods

—n™or— identifi™—t— nell— €i—tt—form— g—r˜on—ti™— edri—ti™oEhin—ri™—D

lo™—lizz—t— tr— il ™ontinente efro—r—˜i™o e quello xord iuropeoF s

„he odontologi™—l terminology —nd v—ri—˜les used in this p—per

notosu™hi er—no ™o™™odrilliformi prett—mente terrestri e questo eÁindiE

—re those of ƒmith 8 hodson @PHHQA —nd ƒmith et —lF @PHHSAF „he terms

™—tivo dell— presenz— di —ree emerse sull— pi—tt—form— ™—r˜on—ti™—F

–™ell—e9 —nd –™—ud—e9 —re used in the sense of e˜ler @IWWPAF €—r—meters of

hor—todon teeth from eustri—D ‚om—ni— —nd rung—ry —re ˜—sed on

me—surements t—ken from photogr—phs pu˜lished in fuffet—ut @IWUWAD

w—rtin et —lF @PHHTAD —nd ™ourtesy of w—rton ‚—˜iD respe™tivelyF sntrodu™tion

„he —verostr—n theropods —re ™onsidered —™™ording to iz™urr—

huring the IWWU field work in pper gret—™eous @PHHTA —s those ˜elonging to the le—st in™lusive ™l—de in™luding neo™erE

—tos—urs —nd tet—nur—ns @pigF PeAF „he ™ro™odyliform phylogeneti™ €ol—zzo fossil site @pogli—noE‚edipugli— wuni™ip—lityD

fr—mework —nd t—xonomy followed herein is th—t of ƒereno 8 v—rsson qorizi— €rovin™eD xi st—lyY pigF IAD —n unusu—l serr—ted

@PHHWA —nd ƒereno et —lF @PHHIAF e™™ordinglyD the gro™odyliformes —re

tooth w—s found @‚igo IWWUAF „he spe™imen w—s preliE

the le—st in™lusive ™l—de in™luding protosu™hids —nd ™rownEgroup ™roE

min—rily —nd tent—tively referred to the ziphodont ™roE

™odiles @gro™odyli—AD the wesoeu™ro™odyli— is the most in™lusive ™roE

™odyliform hor—todon in some overviews of the €ol—zE ™odyliform ™l—de ex™luding protosu™hidsD —nd the xotosu™hi— is the

zo fossil —ssem˜l—ge @eFgFD h—ll— †e™™hi— et —lF PHHI˜Y most in™lusive ™l—de in™luding xotosu™husD ˜ut ex™luding ™rownEgroup

h—ll— †e™™hi— 8 „entor PHHRAD ˜ut rem—ined undeE ™ro™odiles @pigF PfAF

I snstitut g—t—l—Áde €—leontologi— @sg€AD ere— de re™er™— del wesozoi™D is™ol— sndustri—l PQD iEHVPHI ƒ—˜—dellD ƒp—inF P wuseo qeologi™o qiov—nni g—pelliniD vi— —m˜oni TQD sERHIPT fologn—D st—lyF

QIH h—ll— †e™™hi— pFwF 8 g—u eF

seÂum x—tion—ldu xigerD xi—meyD ‚eÂpu˜lique de xigerY w€gwD wuE

seo dell— ‚o™™— di wonf—l™one ± ƒezione qeop—leontologi™— @former

wuseo €—leontologi™o gitt—dino di wonf—l™oneAD qorizi—D st—lyY ‡€D

€—leontologi™—l snstituteD niversity of †ienn—D eustri—F

qeologi™—l —nd p—leontologi™—l notes

„he €ol—zzo fossil site o™™urs in the western porE

tion of the st—li—n se™tor of the u—rst €l—te—u n—med

–g—rso ssontino9 in st—li—nF st —™tu—lly in™ludes two sitesD

n—med site e —nd site f —nd pro˜—˜ly representing two

distin™tD ˜ut ™loseD str—tigr—phi™ se™tions @see h—ll— †e™E

™hi— et —lF PHHI˜ —nd h—ll— †e™™hi— 8 „entor PHHR for

more inform—tionAF

pieldwork —t site f where w€gw IIUPH w—s

found h—s ˜een undert—ken system—ti™—lly sin™e IWWT

˜y the wuseo €—leontologi™o gitt—dino of wonf—l™one

@now €—l—eontologi™—l ƒe™tion of the wuseo dell— ‚o™™—

di wonf—l™oneAF

foth site e —nd f h—ve yielded — pe™uli—r fossil

—ssem˜l—ge ™omposed m—inly of perfe™tly —rti™ul—ted pigF I E vo™—tion of the €ol—zzo site @pogli—noE‚edipugli—D st—lyA

—nd wellEpreserved ostei™hthy—n fishesF „he i™hthyof—uE where w€gw IIUPH w—s foundF

n— in™ludes the py™nodontiform €ol—zzodus ™oron—tus

€oy—toEeriz—D PHIHD the —lopiform ‚hyn™hoder™etisD

sm—ll ™lupeomorphsD r—re gh—nid—eD in™hodus spFD—n

—nguilliformD —nd some sm—llEsized ˜ery™iform

—™—nthopterygi—ns @quidotti IWVQY h—ll— †e™™hi— 8

„entor PHHRY €oy—toEeriz— PHIHAF „errestri—lpl—ntreE

m—ins @prenelopsis spFD fr—™hiphyllum spFD €—giophyllum

spFD ™fF ƒphenolepisD gunningh—mites ™fF eleg—nsD some

pro˜—˜le ginko—les —nd indetermin—te —ngiospermsA —re

rel—tively ™ommon —t site eD —nd —re mu™h r—rer —t site f

@h—ll— †e™™hi— 8 „entor PHHRAF snverte˜r—te eviden™e is

represented only ˜y — few de™—pod ™rust—™e—nsD some

rudist ˜iv—lve shells —nd — level with tr—™e fossils @h—ll—

†e™™hi— 8 „entor PHHRAF „etr—pod rem—ins were found

ne—rly ex™lusively —t site e —nd ™onsist of dis—rti™ul—ted

˜ones of nonEm—rine ™heloni—ns —nd — worn tooth of —

™ro™odyliform @h—ll— †e™™hi— 8 „entor PHHRAF

„he fossiliferous se™tion of site f is only IPPFS ™m

thi™k —nd is ™omposed of thinly l—min—tedD light gr—yE

pigF P E €hylogeneti™ rel—tionships —mong the m—in theropod @eA greenish limestone @h—ll— †e™™hi— et —lF PHHI˜Y h—ll—

—nd ™ro™odyliform @fA ™l—des mentioned in the textF ƒin™e

†e™™hi— 8 „entor PHHRAF wi™rof—™ies —n—lysis suggests

the position of the „h—l—ttosu™hi— —mong the gro™odyliE

th—t deposition w—s in — restri™ted l—goon—l environment

formes is ™ontroversi—lD —n unresolved polytomy is preE

with intertid—lepisodes @ƒ—ndro †enturiniD persF ™ommF sented hereF

to pF wF hF †FAF w€gw IIUPH w—s found —t the ˜oundE

—ry ˜etween l—yers f —nd gD PHH —nd IWH mm thi™k

e˜˜revi—tions for v—ri—˜lesF evD —pi™—llengthYgeD ™rown

respe™tively @see h—ll— †e™™hi— 8 „entor PHHRD figF PUAF —ngleY gfvD ™rown ˜—se lengthY gf‡D ™rown ˜—se widthY gf‚D ™rown

foth l—yers —re ™omposed of sets of strom—tolite l—miE ˜—se r—tio @gf‡GgfvAY grD ™rown heightY gr‚D ™rown height r—tio

@grGgfvAY heD dist—l—pi™—ldenti™lesdensity @per mmAY hfD dist—l n—e —nd thinD ™o—rser storm levelsF „he tooth w—s deE

˜—s—l denti™les density @per mmAY hgD dist—l midE™rown denti™les denE posited —t the ˜—se of the l—min— fT th—t is the lowerE

sity @per mmAY wfD mesi—l˜—s—ldenti™lesdensity @per mmAY wgD mesi—l

most storm level of l—yer fY other fossils found in fT

midE™rown denti™les density @per mmAF

in™lude the fish ‚hyn™hoder™etisF „his se™tion is p—rt of

— thi™ker str—tigr—phi™ se™tion m—de of pl—tyD wellE snstitution—l —˜˜revi—tionsF pqq fD p—™ult—te— de qeologie

sÎi qeofizi™—D niversit—te— fu™uresÎtiD fu™h—restD ‚om—ni—Y wxxD wuE ˜edded limestone ™ropping out e—st of the vill—ge of

„he first re™ord of — notosu™hi—n ™ro™odyliform from st—ly QII

€ol—zzoD ˜elonging to the thi™k pper gret—™eous ™—rE

˜on—te pl—tform sequen™e of the u—rstF „his sequen™e is

domin—ted ˜y limestone with rudist ˜iv—lves —nd is

known in liter—ture under the n—me of –g—l™—ri di

wFte ƒ—n wi™hele9 @w—rtinis IWTPAF „he lithostr—tiE

gr—phi™ unit ™ont—ining the €ol—zzo se™tion is ™—lled

lo™—lly –g—l™—ri di eurisin—9 @known inform—lly —lso —s

–™—l™—ri — ‚udiste9A —nd its —ge is ™onsidered „uroni—nE

e—rly ƒenoni—n @„entor et —lF IWWRAF „intori et —lF @IWWQA

referred site e to the lower ƒenoni—n ˜e™—use of the

presen™e of the for—minifer e™™ordiell— ™oni™— —nd the

—lg—e ƒgrossoell— p—rtenopei— —˜ove the fish levels —nd

the for—minifer wont™h—rmonti— —pennini™— ˜elowF

wont™h—rmonti— spF w—s found —lso —t site fY „he forE

—minifers wF —pennini™—D €seudo™y™l—mmin— sph—eroiE

de—D e™™ordiell— spFD —nd ƒ™—ndone— s—mniti™—D —nd the

—lg—e ƒgrossoell— spF were found in — se™tion immediE

—tely ˜elow the fossiliferous se™tion of site fF „his —sE

sem˜l—ge suggests —n e—rly ƒenoni—n —ge @iFeFD goni—E

™i—nEƒ—ntoni—nA —lso for site fD in —greement with the

str—tigr—phi™ fr—mework of the u—rst €l—te—u @ƒF †entuE

riniD persF ™ommF to pF wF hF †FAF

„he u—rst €l—te—u is formed m—inly ˜y ™—r˜on—tes

gener—lly supposed to h—ve ˜een deposited in — sh—llow

pigF Q E „he western „ethys during ƒ—ntoni—n @v—te gret—™eousA m—rine environment —t tropi™—ll—titudes—nd domin—ted

timesD ˜—sed on i˜erli et —lF @IWWQAD with lo™—tion of the

˜y rudist –reefs9 during v—te gret—™eous timesF pollowE

u—rst €l—te—u @—steriskA on the edri—ti™Ehin—ri™ g—r˜oE

ing re™ent p—l—eogeogr—phi™ re™onstru™tion of western

n—te €l—tformF h—rk gr—y ™olor represents deep se— on

„ethys @seeD for ex—mpleD i˜erli et —lF IWWQY g—moin et o™e—ni™ ™rustD light gr—y epi™ontinent—l se— —nd m—rine

—lF IWWQY €hilip et —lF PHHHAD the u—rst €l—te—u w—s situE ˜—sins on ™ontinent—l™rustD white emergent l—ndD—nd

˜ri™kEp—ttern identifies the ™—r˜on—te pl—tformsF —ted —t the northern point of — wideD isol—ted intr—o™e—E

ni™ ™—r˜on—te pl—tform of f—h—mi—n type @edri—ti™EhiE

n—ri™ ™—r˜on—te pl—tformA during ƒenoni—n times @pigF

hes™riptionF w€gw IIUPH is — shed ™rownY the QAF imersion events —ffe™ted lo™—lly or extensively this

root w—s ne—rly ™ompletely reE—dsor˜ed —nd the ˜—s—l pl—tform sever—l times during gret—™eous timesY the

side of the ™rown —ppe—rs ™on™—ve in l—˜iolingu—l view most import—nt o™™urred during the l—te g—mp—ni—nE

@pigF ReEfAF foth mesi—l—nd dist—l™utting m—rgins ˜e—r w——stri™hti—n when — rel—tively l—rge isl—nd formed

@edri—ti™ ssl—ndX ytoni™Ï—r PHHUY h—ll— †e™™hi— PHHVD serr—ted ™—rin—eF sts ev is IPFPS mm —nd gr is IIFQTY

PHHWAF „he emergent p—rts of the ™—r˜on—te pl—tform the gfv is SFVP mmF „he gf‡ is not signifi™—tive ˜eE

˜elonged to —n —r™hipel—go of isl—nds pl—™ed ˜etween ™—use the l—˜i—l side is d—m—ged —nd the lingu—l side is

the xorthern iurope—n l—ndm—ss —nd the efro—r—˜i—n slightly ™oll—psedY width —t midE™rown is QFHU mmF „he

™ontinent @the iurope—n er™hipel—goX see h—ll— †e™™hi— gf‚ ™—nnot ˜e ™—l™ul—tedY howeverD the ™rown is not

PHHTY €ered—Eƒu˜er˜iol— et —lF PHHWAF strongly ™ompressed l—˜iolingu—llyX the width —t the

midE™rown is more th—n h—lf the gfv @SV7AF „he

™rown is rel—tively elong—ted ˜—so—pi™—llyD —s the gr‚

ƒystem—ti™ €—l—eontology equ—ls IFWSF

„he l—˜iolingu—l pol—rity is likely indi™—ted ˜y the

position of the mesi—l™—rin— th—t twists on one sideD

er™hos—uri— gopeD IVTW presum—˜ly the lingu—l oneF

„he outline of the ™rown in side view is tri—ngul—rD gro™odylomorph— ‡—lkerD IWUH

moder—tely ™urved dist—lly @ge is TV degreesAD with —

gro™odyliformes r—yD IWQH

™onvex mesi—lm—rgin th—t ™urves smoothlyfrom the

wesoeu™ro™odyli— ‡hetstone —nd ‡hy˜rowD IWVQ

˜—s—l p—rt to the —pexD —nd — ne—rly str—ight —nd slightly

xotosu™hi— q—sp—riniD IWUI in™lined dist—l m—rgin @pigF ReEfAF „he —pi™—l h—lf of the

™rown is slightly ™urved lingu—llyF „he ˜—se of the xotosu™hi— genF —nd spF indetF

pigs REUe ™rown is ™onstri™tedY the ™onstri™tion is wellEm—rked

QIP h—ll— †e™™hi— pFwF 8 g—u eF

pigF R E w€gw IIUPHD xotosu™hi— spF —nd genF indetF eA lingu—l viewY fA l—˜i—l viewY gA dist—l viewY hA mesi—l viewF ƒ™—le ˜—r equ—ls I mmF

—long the dist—l m—rginD where — not™h is evident @pigF the outline of the m—rgin just —˜ove the indent—tion is

SAD ˜ut it is imper™epti˜le —long the mesi—l m—rginF „he slightly ˜ut distin™tly ™onvex @–˜ump9A @pigF SAY interdenE

dist—lm—rgin presents — –˜ump9 just —˜ove the not™hF ti™le slits in the ˜—s—l p—rt —re shorter —nd ™ell—e less

„he l—˜i—l side is symmetri™—lly ™onvex @iFeFD the —pex developed th—n in su™™essive denti™lesY hf is VF widE

of the ™onvexity is ™entr—lon the sideY pigF RfAY the ™rown denti™les —re more distin™tD ™hiselEsh—pedD —nd

lingu—l side h—s — ™entr—lD longitudin—l relief with — fl—t ˜—so—pi™—lly longer th—n high @pigF TeEfAY denti™les —re

surf—™e —nd step —nd ™on™—ve mesi—l—nd dist—lslopes slightly in™lined tow—rd the —pex of the ™rownD ˜ut they

th—t —re ™onfluent with the serr—ted ™—rin—e —nd emph—E —re symmetri™—lF hg is SESFSF ƒh—llow ™—ud—e extend on

size them @pigF ReD UeAF the ™—rin— —nd —re —ngled ˜—somesi—llyD where—s the inE

„he midE™rown ™rossEse™tion is su˜re™t—ngul—r to terdenti™le slits —nd ™ell—e o™™ur only in the most exterE

dropElikeD with — rounded —nd wider mesi—l sideD — n—rE n—l p—rt of the ™—rin—Y very thin en—mel ridges p—r—llel to

rower dist—lside with — sh—rp ™—rin—D — ™onvex l—˜i—l the ™—ud—e extend onto the lingu—l surf—™e of the ™rown

sideD —nd — more fl—ttened lingu—l sideF @pigF TeAF „here is —n en—melridge on the midlineof

„he —pex of the ™rown presents — sm—ll we—r f—™et e—™h midEdist—ldenti™le@pigF TfAF „ow—rd the —pexD denE

extending slightly on the lingu—l surf—™eF „he ™rown ti™les ˜e™ome sm—ll —g—in @he is UFSEVA —nd they —re not

surf—™e is not flutedY it shows thin —nd irregul—r longE ™le—rly distin™t in the uppermost PFQ mm of the ™—rin—F

itudin—l wrinkles of the en—melF ƒm—ll ™ir™ul—r pits ™—n „he mesi—l ™—rin— is ™lose to the midline ne—r the

˜e o˜served in the midElower p—rt of the lingu—l side —pex of the ™rownD ˜ut it shifts on the lingu—l surf—™e

@pigF ReD UeAF „he l—rgest h—s — di—meter of HFRQ mmD just I mm ˜elow the —pex —nd twists lingu—lly @soD denE

where—s the sm—llest h—s — di—meter of HFIR mmF „he ti™les point lingu—lly —nd only slightly mesi—llyAF „husD

l—rgest pit presents ™on™entri™ lines of ˜re—k—ge —nd only — short —pi™—l segment ™oin™ides with the mesi—l

™oll—pse of the tooth surf—™eD whi™h suggest it is — stru™E m—rgin of the ™rown @pigF RhAF „he mesi—l™—rin—

ture ™—used ˜y ™ompression ˜y — sm—llD ™oni™—l pointD re—™hes the ˜—s—lp—rt of the ™rownD ˜ut the ˜—s—lsegE

possi˜ly the tooth of — ™run™hing fish @pun™ture m—rkAF ment @~PFW mmA is d—m—ged —nd denti™ul—tion is only

„he dist—l™—rin— extends from the —pex to the ˜—s—l p—rtly preservedF e tot—l of PV denti™les ™—n ˜e ™ounted

not™h —nd is ™ontiguously serr—ted @pigF RgAF f—s—llyD it in the rem—ining portion of the ™—rin—F wg is TD thus the

shifts on the lingu—l side in ™orresponden™e of the mesi—l denti™les —re slightly sm—ller th—n the ™orreE

–˜ump9F e tot—lof RS denti™les™—n ˜e ™ountedF „hey sponding dist—l denti™lesF wid mesi—l denti™les differ

—re slightly sm—ller in the ˜—s—l p—rt of the ™—rin— where from mid dist—lones in ˜eing lessdistin™t ˜e™—use interE

„he first re™ord of — notosu™hi—n ™ro™odyliform from st—ly QIQ

denti™le slits —re very short —nd ™ut only the m—rgin of

the ™—rin—D ™ell—e —re redu™edD —nd ™—ud—e —re not develE

oped @pigF TgEhAF „he few unworn denti™les —re —symE

metri™—l @pointing —pi™—llyA —nd longer th—n highF „he

denti™les diminish in size tow—rd the —pex @~VGmmAY

individu—ldenti™les—re not distin™t in the uppermost

QFS mm of the ™—rin—D ˜e™—use the interdenti™le slits

—re very short —nd ™ut only the outer p—rt of the ™—rin—

@they look like p—pill—eY pigF TgAF

his™ussion

„he tooth o˜viously ˜elongs to — ™—rnivorousD

gn—thostom—te verte˜r—te —ndD —ssuming th—t the shift

of the mesi—l ™—rin— is lingu—lD it ™omes from the left

upper j—w or from the right lower j—wF ƒerr—ted teeth

o™™ur in different gn—thostom—te groupsD from ™honE

dri™hthy—n fishes to m—mm—li—ns @eFgFD p—rlow et —lF

IWWIY e˜ler IWWPAF €resen™e of teeth with l—˜iolingu—lly

™ompressed ™rowns ˜e—ring finely denti™ul—ted ™—rin—e

@ziphodont teethA is ™onsidered —n —r™hos—uriform syE

n—pomorphy @eFgFD fenton IWWWA th—t is se™ond—rily lost

in sever—l derived line—gesX m—ny ™ro™odyliformsD most

pteros—ursD —nd ornithis™hi—n —nd s—uropodomorph diE

nos—ursF emong gret—™eous —r™hos—ursD ziphodont

teeth —re reported in most @˜ut not —llA theropod ™l—des

—nd sever—l™ro™odyliformsF „hereforeD the presen™e of

ziphodont teeth is — symplesiomorphi™ ™ondition th—t

—lone ™—nnot ˜e used for referring isol—te teeth to — less

in™lusive —r™hos—ur ™l—deF elthough ziphodonty is

pro˜—˜ly plesiomorphi™ for theropods dinos—ursD isoE pigF S E w€gw IIUPHD xotosu™hi— spF —nd genF indetF €—rti™ul—r of

l—ted ziphodont teeth th—t —re found in gret—™eous the ˜—s—l™onstri™tionF „he —rrow points to the more

m—rked dist—l ™onstri™tionF ƒ™—le ˜—r equ—ls I mmF ro™ks origin—ted from ™ontinent—lsediments ± —nd

pigF T E w€gw IIUPHD xotosu™hi— spF —nd genF indetF eEfA dist—lm—rgin in lingu—l@eA—nd dist—l@fA viewY gA the midE—pi™—lp—rt of the mesi—l m—rgin in lingu—l viewY hA midEdenti™les of the mesi—l m—rginF ƒ™—le ˜—r equ—ls I mmF

QIR h—ll— †e™™hi— pFwF 8 g—u eF

sometimes —lso from sediments deposited in tr—nsiE —nd the presen™e of — str—ightD serr—ted dist—lm—rgin

tion—lGsh—llow m—rine settings ± —re tr—dition—lly reE @ƒmith et —lF PHHSAF st differs from the prem—xill—ry teeth

ferred to th—t ™l—de @eFgFD gurrie et —lF IWWHY piorillo 8 of known —˜elis—uroids ˜e™—use the ™rown shows —

gurrie IWWRY vee IWWUY f—szio IWWUY „ori™es PHHQY more —symmetri™—l ™rossEse™tion profile in the l—tterD

„ori™es et —lF PHHRY ƒ—nkey et —lF PHHPD PHHSY ƒ—nkey more m—rkedly ™onvex l—˜i—lly —nd fl—tter lingu—lly

PHHVY vongri™h PHHVAF @ƒmith PHHUAF yne pe™uli—r fe—ture of w€gw IIUPH is

w€gw IIUPH ™omes from — se™tion whose sediE the presen™e of — slight ™onstri™tion —t the ™rownEroot

ments pro˜—˜ly origin—ted in —n intertid—l to l—goon jun™tion —long the dist—l m—rginD m—rked ˜y — low

environment ™ont—ining r—re m—rine inverte˜r—tes —nd –˜ump9 of the serr—ted m—rgin just —˜ove the indent—E

for—minifersD fishes with — m—rine to fresh w—ter distriE tionF elthough widespre—d —mong ˜—s—l dinos—uriformsD

˜ution @iFeFD €y™nodontiformesA —nd fishes with — m—rine ornithis™hi—nsD ˜—s—ls—uropodomorphsD —nd the primiE

distri˜ution only @‚hyn™hoder™etis —nd fery™iformesAF tive „ri—ssi™ s—uris™hi—n ior—ptor @v—nger 8 fenton

roweverD the r—re rem—ins of terrestri—lveget—tion PHHTAD the ™om˜in—tion of ™h—r—™teristi™s o˜served in

found in the toothE˜e—ring se™tionD —nd the nonEm—rine

w€gw IIUPH is —˜sent in —ll known nonE™oeluros—ur

turtles —nd —˜und—nt pl—nts ™olle™ted —t site eD testify to

theropods @‚—uhut PHHQAF emong goeluros—uri—D — ˜—E

the presen™e of ™lose emergent —re—sF pootprints of

s—l™onstri™tion of the ™rown is reported in the ˜—s—l

sm—ll to mediumEsized ˜iped—l dinos—ursD mostly referE

ornithomimos—ur €ele™—nimimus @€erezEworeno et —lF

—˜le to theropods with — size simil—r to th—t of the posE

IWWRAD therizinos—uroids @—nno PHIHAD —lv—rezs—urids

si˜le owner of the des™ri˜ed toothD —re rel—tively ™omE

@€erle et —lF IWWRAD troodontids @gurrie IWVUAD the droE

mon in the upper el˜i—n —nd upper genom—ni—n ™—r˜oE

m—eos—urid wi™ror—ptor @rw—ng et —lF PHHPAD toothed

n—te pl—tform limestones of the ne—r˜y sstri—n €eninsuE

˜irds @€—di—n 8 ghi—ppe IWWVAD —nd in the prem—xill—ry

l—D gro—ti— @eFgFD h—ll— †e™™hi— IWWVY h—ll— †e™™hi— 8

teeth @˜ut not in m—xill—ry or dent—ry teethA of the ˜—s—l

„—rl—o PHHHY h—ll— †e™™hi— et —lF PHHI—D PHHPAF ƒm—ll

tyr—nnos—uroid €ro™er—tos—urus @‚—uhut et —lF PHIHAF

theropod teeth —re reported from the w——stri™hti—n site

„he €ol—zzo spe™imen differs from —ll these t—x— in

of uozin— @ƒ‡ ƒloveni—Y he˜elj—k et —lF IWWWD PHHPA th—t

the presen™e of — serr—ted ˜—s—lh—lfof the mesi—l™—rin—F

is —˜out RH km ƒi of the €ol—zzo siteF „husD the identiE

w€gw IIUPH is ™le—rly different from the teeth of the fi™—tion of w€gw IIUPH —s — theropod tooth would not

therizinos—uroidsD those of the few dent—te ornithomiE

™ontr—st with the lo™—l—ndregion—lgeologi™—lhistory

mos—ursD —nd those of the troodontids @in™luding those

—nd f—un—l—ssem˜l—gesF

˜—s—l t—x— th—t ˜e—rs finely denti™ul—ted ™—rin—eAD ˜eE

™—use the ™onstri™tion of the ™rown ˜—se is not m—rkedD

gomp—risons with theropod dinos—urs

expe™i—lly mesi—lly @f—rrett PHHWAF „he €ol—zzo tooth

„heropod teeth —re ™onsidered di—gnosti™ —t low

differs from —ll known theropod teeth in the in™omplete t—xonomi™ level @eFgFD gurrie et —lF IWWHY piorillo 8 gurE

serr—tion of the mesi—l™—rin—F „his ™ondition h—s never rie IWWRY f—szio IWWUY ƒ—nkey et —lF PHHPD PHHSY ƒmith et

reported —mong theropodsX —ll des™ri˜ed spe™imens —re

—lF PHHSY ƒ—nkey PHHVY vongri™h PHHVAF f—sed on the

either –truly9 serr—ted or unserr—ted @eFgFD gurrie et —lF su˜tri—ngul—r sh—pe of the ™rown —nd its limited ™urv—E

IWWHY f—szio IWWUY „ori™es PHHQY „ori™es et —lF PHHQY ture in l—˜iolingu—l viewD the rel—tively high l—˜iolingu—l

ƒ—nkey et —lF PHHPD PHHSY ƒ—nkey PHHVY vongri™h PHHVAF widthD —nd the —˜sen™e of —pi™—l™urv—ture of the dist—l

sn ™on™lusionD the ™om˜in—tion of ™h—r—™ter ™onditions m—rginD w€gw IIUPH differs from the l—ter—l @m—xill—ry

seen in w€gw IIUPH is unreported in theropod teeth —nd dist—ldent—ryA teeth of most theropodsD the l—tter

@„—˜F IAF sn p—rti™ul—rD the ™om˜in—tion of in™omplete t—x— showing l—˜iolingu—lly fl—ttenedD ˜l—deElike —nd

mesi—lserr—tion —nd — ˜—s—l–˜ump9 h—s never ˜een reE —pi™—lly re™urved ™rowns in l—˜iolingu—l view @eFgFD gurE

ported in —ny theropod t—xonF elthough some single rie et —lF IWWHY f—szio IWWUY ƒ—nkey et —lF PHHPD PHHSY

fe—tures @over—ll tooth morphologyD denti™le sh—pe —nd fro™hu PHHQY ƒ—nkey PHHVY vongri™h PHHVAF w€gw

density in the dist—l™—rin—A —re sh—red with —˜elis—urE IIUPH sh—res the presen™e of — lingu—lly shifted mesi—l

oidsD tyr—nnos—uroids —nd drom—eos—uridsD —ll —ltern—E ™—rin— with prem—xill—ry —nd mesi—lEmost @symphysi—lA

tive pl—™ements of w€gw IIUPH —mong the „heropod— dent—ry teeth of most —verostr—n theropods @gurrie et

require — ™omp—r—˜le —mount of homopl—syF €ending —lF IWWHY ‚—uhut PHHQY ‚—uhut et —lF PHIHAF st sh—res

—ddition—l eviden™eD it is unlikely th—t w€gw IIUPH some derived fe—tures with mesi—lmost m—xill—ry —nd

˜elongs to — yetEunreported line—ge of theropods with dent—ry teeth of —˜elis—uroid neo™er—tos—ursX lingu—lly

shifted mesi—l™—rin— @ƒ—mpson et —lFPHHIY ƒmith PHHUAD —n unusu—ldent—lmorphologyF

low ˜ut distin™t ˜—so—pi™—lly elong—te ™onvex —re— runE

gomp—risons with gro™odyliformes ning —long the ™enter of the lingu—l surf—™e with fl—t to

slightly ™on™—ve —re—s —dj—™ent to the serr—ted ™—rin—e gro™odyliform teeth —re usu—lly ™oni™—l with unE

@‚—uhut PHHRY p—nti 8 „herrien PHHUY ƒmith PHHUAD serr—ted ™—rin—e @idmund IWTWAD ˜ut the xotosu™hi—

„he first re™ord of — notosu™hi—n ™ro™odyliform from st—ly QIS

„—˜F I E histri˜ution of the odontoE Lingual shift of mesial carina logi™—l fe—tures dis™ussed in Carinae Mid-crown Basal crown (in non-premaxillary Taxon/grade

ornamentation cross-section constriction and non-symphysial

the text —mong ™ro™odyliE dentary teeth) form —nd theropod t—x—F Crocodyliformes

“Protosuchids” denticles elliptical absent absent

“Sphenosuchids” denticles elliptical absent absent Non-Geosaurini absent rounded absent absent thalattosuchians denticles/ Geosaurini elliptical absent absent pseudo-serrations Most neosuchians absent rounded absent absent

Pristichampsus denticles elliptical absent absent denticles/ Trematochampids elliptical absent absent pseudo-serrations “Sebecosuchians” denticles/absent elliptical absent absent

“Notosuchids” cusps teardrop-shaped marked absent

Araripesuchus denticles elliptical moderate in A. wegeneri

Doratodon denticles elliptical moderate absent

Theropod dinosaurs elliptical or Abelisauridae denticles absent in Majungasaurus D-shaped Non-maniraptoran elliptical or denticles absent absent tetanurans D-shaped Troodontidae denticles/absent rounded/elliptical present absent ellitptical or in Dromaeosauridae denticles/absent in Dromaeosaurus D-shaped Microraptor Denticles, not completely subrectangular to MPCM 11720 moderate present individualized in the drop-like

apical part of carinae

™le—rly individu—lized ˜y interdenti™le slits —nd ™ell—e @–iphosu™hi—9 sensu gomp—ny et —lF PHHSAD — few meE

extending on the whole ™—rin—F triorhyn™hids @endr—de et —lF PHIHAD €risti™h—mpsus

e™™ording to €r—s—d 8 de v—pp—rent de froin —nd the „rem—to™h—mpsid—e @fuffet—ut IWURA —re ™h—rE

@PHHPAD ziphodonty is highly homopl—sti™ within groE —™terized ˜y l—˜iolingu—lly ™ompressed teeth with serE

™odyliformes @„—˜F IAF por ex—mpleD the m—rine metriorE r—ted ™utting m—rgins m—de of distin™t denti™les @€r—E

hyn™hids qeos—urus gig—nteus —nd h—kos—urus m—xiE s—d 8 de v—pp—rent de froin PHHPY gomp—ny et —lF

mus from pper tur—ssi™ —nd lowermost gret—™eous PHHSY ƒereno 8 v—rsson PHHWY „urner 8 ƒerti™h PHIHAF

h—ve denti™ul—ted ™utting m—rginsD where—s their ™lose e™™ording to €r—s—d 8 de v—pp—rent de froin @PHHPX

rel—tive „orvoneustes shows — –f—lse9 ziphodonty @see QTA ––true ziphodonts —re those with teeth h—ving l—tE

endr—de et —lF PHIHAF er—lly ™ompressed ™rowns with posterior re™urved

xotosu™hi— is — ™l—de th—t w—s ™ommon in ƒouth —pexD —nterior —nd posterior ™—rin—e ˜e—ring — num˜er

emeri™— —nd efri™— during the gret—™eous @q—sp—rini et of isol—ted festoonElike denti™les99 where—s –f—lse ziE

—lF IWWIY ƒereno et —lF PHHQY ƒereno 8 v—rsson PHHWAD phodont teeth9 —re those where ––the —nterior —nd posE

where—s in iuropeD ziphodont t—x— ˜e™—me more freE terior ™—rin—e —re rel—tively ™o—rse —nd ˜e—r ™renul—E

quent during the io™ene @s˜erosu™husD fergisu™husD the tions gener—lly formed ˜y —n—stomisingD irregul—r

–i˜ero™™it—ni—n ™ro™odile9D —nd the eusu™hi—n €ristiE ridges issued from the m—in ˜ody of the ™rown99F e™E

™h—mpsusY veg—s— et —lF IWWRY €r—s—d 8 de v—pp—rent ™ording to veg—s— et —lF @IWWRX IQPA true ziphodont

de froin PHHPAF m—rgins —re those h—ving ™—rin—e with isol—ted dentiE

„he referr—lof w€gw IIUPH to €risti™h—mpsusD ™lesD —nd ––—n –isol—ted denti™le9 is — dis™rete morphoE

the ƒe˜e™osu™hi— @in™luded in the xotosu™hi— —™™ordE logi™—l unitD ™le—rly individu—lized ˜y —n interdenti™le

ing to ƒereno et —lF PHHIAD —nd the trem—to™h—mpsids is groove or not™h99F pollowing this definition of ziphoE

dismissed ˜e™—use of the —˜sen™e of ˜oth the ˜—s—l dontyD w€gw IIUPH is not — –f—lse ziphodont9 tooth

–˜ump9 —nd the lingu—lly shifted ™—rin—D —nd the preE —nd is ™loser to — true ziphodont ™onditionD —lthough

sen™e of — ™omp—r—tively ˜lunter —nd rounded —pex of the denti™les of the —pi™—l h—lf of the mesi—l ™—rin— —nd

the ™rown in those ™l—des @ghi—ppe IWVVY q—sp—rini et those of the the —pi™—lp—rt of the dist—l™—rin— —re not

QIT h—ll— †e™™hi— pFwF 8 g—u eF

—lF IWWIY fu™kley 8 fro™hu IWWWY „urner 8 g—lvo somewh—t sm—ller th—n w€gw IIUPHD the ˜—so—pi™—l

PHHSAF height r—nging IIEU mmF sn f—™tD fuffet—ut @IWUWX WIA

reported th—t the l—rgest tooth is II mm –long9 —nd —

gomp—risons with hor—todon sm—ller one is ––only U mm long ‘FFF“ the —nteroposterior

e™™ording to gomp—ny et —lF @PHHSA —nd w—rtin et di—meter is R mmD the l—˜iolingu—l one PFS mm99D ˜ut he

—lF @PHIHAD the notosu™hi—n hor—todon is the only true did not report the museum num˜er of the spe™imensD

ziphodont ™ro™odyliform known in the pper gret—™E thus we don9t know to whi™h of the figured spe™imens

eous of iuropeF f—sed on its sm—ll sizeD its s™—r™e l—˜ioE he refersF

lingu—l ™ompression @it is not ˜l—deElikeAD —nd modest hor—todon i˜eri™us gomp—nyD €ered— ƒu˜er˜iol—D

™urv—tureD h—ll— †e™™hi— et —lF @PHHI˜A —nd h—ll— †e™E ‚uizEymen—™— 8 fus™—lioniD PHHS is ˜—sed on —n inE Ä

™hi— 8 „entor @PHHRA tent—tively referred w€gw IIUPH ™omplete lower j—w with teeth found in the ƒierr— €eE

to the only ™ro™odyliform with denti™ul—ted teeth ren™hiz— porm—tion —t the gher— site @†—len™i—D ƒp—inAD

known in the pper gret—™eous of iuropeD hor—todonD l—te g—mp—ni—n or e—rly w——stri™hti—n in —geF „eeth —re

inste—d of — theropod t—xonF hor—todon is reported heterodont in size —nd sh—peY the fourth tooth is su˜E

from the uppermost gret—™eous of eustri— @funzel ™oni™—l@gf‚ is —˜out HFTPAD where—s ™rowns in — more

IVUIY ƒeeley IVVIY fuffet—ut IWUWAD ƒp—in @gomp—ny et dist—l position —re l—ter—lly ™ompressed ˜ut str—ightF „he

—lF PHHSAD ‚om—ni— @w—rtin et —lF PHHTA —nd rung—ry fourth dent—ry tooth resem˜les w€gw IIUPH in over—ll

@‚—˜i PHHVAF ƒp—nishD ‚om—ni—nD —nd rung—ri—n spe™iE sh—pe —nd size @gomp—ny et —lF PHHSD figF QAF roweverD

mens were found in ro™ks origin—ted from fluvi—l —ndG its mesi—l ™—rin— does not shift —nd twist lingu—lly —s

or l—™ustrine sedimentsY the eustri—n spe™imens ™ome well —s th—t of the other teethD in™luding the mesi—lEmost

from — sequen™e of ™o—lse—ms —nd freshw—terGne—rshore ones @gomp—ny et —lFD PHHSY tulio gomp—nyD persF

m—rine ™l—sti™ ro™ks @uv—™Ïek 8 rerm—n PHHRAF ™ommF to pF wF hF †FAF purthermoreD serr—tions of the

hor—todon ™—r™h—ridens @funzelD IVUIA is the type ™—rin— —re —lw—ys formed ˜y ™le—rly individu—lized denE

spe™ies of the genus hor—todon ƒeeley @IVVIA —nd is ti™les in this t—xon @gomp—ny et —lF PHHSAF „here —re

˜—sed on —lmost ™omplete m—ndi˜le with poorly preE three —nd five denti™les per mm —t midE™rown in l—rge

served teethD — fr—gment—ry m—xill— —nd eight isol—ted —nd sm—ll teethD respe™tively @gomp—ny et —lF PHHSAF

teeth @fuffet—ut IWUWAF ell spe™imens ex™luding one henti™les —re ™omp—r—tively l—rgerD —nd p—ired denti™les

tooth —re from the lower g—mp—ni—n qrun˜—™h porm—E joined —t the ˜—se —re irregul—rly presentD —t le—st in the È

tion of the wuthm—nnsdorf lo™—lityD eustri— @ƒummesE figured se™ond m—ndi˜ul—r tooth @gomp—ny et —lF PHHSD

˜erger et —lF PHHUAF „eeth —re homodont in size @gomE figF RAF e m—rked ˜—s—l™onstri™tion —longthe dist—l

p—ny et —lF PHHSAD ˜ut heterodont in sh—pe @fuffet—ut m—rgin —nd the low –˜ump9 just —pi™—l to itD —nd the

IWUWAF fuffet—ut @IWUWD plF IA figured four isol—ted ™entr—lD longitudin—l relief with — fl—t surf—™e in the linE

™rowns th—t —ppe—r more ˜ul˜ous th—n w€gw IIUPHD gu—lside —re not present @gomp—ny et —lFPHHSY tF gomE

—lthough fuffet—ut @IWUWX WIA des™ri˜ed them —s p—nyD persF ™ommF to pF wF hF †FAF

––strongly ™ompressed l—ter—lly99F purthermoreD their ƒever—lteeth from the lowerw——stri™hti—n or

™rossEse™tion is reported —s ov—l—nd the mesi—l™—rin— possi˜ly ˜—s—l upper w——stri™hti—n p—Ãnt—Ãnele —nd €ui

does not shift on the lingu—l side @fuffet—ut IWUWY pigF lo™—lities of the r—teg f—sin of „r—nsylv—ni— @‚om—ni—A

UfAF „he dist—l m—rgin is sigmoid in l—˜iolingu—l view in were referred ˜y w—rtin et —lF @PHHTA to hor—todon spF

‡€ PQRWGSV —nd TH @fuffet—ut IWUWD plF ID figs T —nd U „he —ttri˜ution to the genus is ˜—sed on the ™—rin—e

respe™tivelyAD where—s it is ne—rly str—ight in w€gw displ—ying — ––™le—r pli™—tion of the en—mel —nd the denE

IIUPHF ‡€ PQRWGSW @fuffet—ut IWUWD plF ID figF SA h—s tineD whi™h is ™h—r—™teristi™ of the genus99 @w—rtin et —lF

— less sigmoid dist—l m—rgin respe™t to the other hF PHHTX QPAD me—ning with th—t the presen™e of true denE

™—r™h—ridens teethD —nd — ˜—s—ldist—l™onstri™tion @pigF ti™les @teremy w—rtinD persF ™ommF to eF gFAF roweverD

UfAF yne side of ‡€ PQRWGSW @in the ™—ption of plF ID we note th—t su™h — fe—ture is not —ut—pomorphi™ for

fuffet—ut IWUW does not spe™ify whether the lingu—l or hor—todonD this ˜eing — notosu™hi—n symplesiomorphy

l—˜i—l is figuredA seems to h—ve — ™entr—lD longitudin—l sh—red ˜y sever—lother t—x— @v—g—s— et —lFIWWRY yrteg—

relief with — fl—t surf—™e @see pigF UfAD possi˜ly present et —lF PHHHY ƒereno 8 v—rsson PHHWAF „he only referen™e

fuffet—ut —lso in ‡€ PQRWGSVF roweverD this would ˜e in ™onE to teeth in the di—gnosis of hor—todon ˜y

@IWUWX WHEWIA is ––teeth strongly ™ompressed l—ter—llyD tr—st with the des™ription of the ™ross se™tion of the

with finely serr—ted edges —nd smooth en—mel99D where—s hor—todon ™—r™h—ridens teeth —s ov—lgiven in fuffet—ut

hor—todon is not di—gnosed ˜y —ny dent—lfe—tures in @IWUWAF ‡€ PQRWGSW is only slightly less ™urved th—n

the revision ˜y gomp—ny et —lF @PHHSAF yne of the two w€gw IIUPH @ge is UH degreesA —nd less elong—ted

„r—nsylv—ni—n morphotypes @pqq f ‚FIWQWY w—rtin ˜—so—pi™—lly @gr‚ equ—ls IFUVAF e™™ording to the deE

et —lF PHHTD plF ID figF IEPY pigF UhA resem˜les w€gw s™ription ˜y fuffet—ut @IWUWAD the whole ™—rin—e —re fiE

IIUPH in the over—ll sh—pe of the ™rown in l—˜iolingu—l nely serr—tedD with V denti™lesGmmD ˜ut the ™rowns —re

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viewD tri—ngul—r —nd high @gr‚ equ—ls PFHRD ge is UP the mesi—l ™—rin—D whi™h h—s —lso wellEdefinedD l—rgeD

—nd ™hiselEsh—ped denti™lesD —t le—st —t midE™rownF degreesAD the slight l—˜iolingu—l ™ompressionD the ™onE

prom the photogr—ph of the ™rown @™ourtesy of wF stri™tion —t the ˜—se of the ™rown only visi˜le dist—lly

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itudin—l relief with — fl—t surf—™e is present —long the ˜—s—llingu—ltwistingof the dist—l™—rin—F roweverD it

lingu—l side or notF „his spe™imen resem˜les the spe™iE differs from w€gw IIUPH inX the sh—pe of the dentiE

men pqq f ‚FIWQW figured ˜y w—rtin et —lF @PHHTA @™fF ™lesD h—ving the s—me denti™les morphology mesi—lly

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hor—todonF „his ™—sts sever—ldou˜ts on the referr—lof this p—rt —nd the ™—rin—e —re not mentioned in the

isol—ted ziphodont teeth from the pper gret—™eous of des™riptionF

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hor—todon is des™ri˜ed in det—ilin the liter—tureDthe hF †FA sh—res with w€gw IIUPH — t—ll @gr‚ equ—ls

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m—ni— @dr—wn from w—rtin et —lF PHHTD plF ID figFP—AY iA IIth left m—ndi˜ul—r tooth of

er—ripesu™hus wegeneriD lingu—l viewD wxx qehPHD ilrh—z porm—tionD q—dof—ou—D

xiger @from ƒereno 8 v—rsson PHHWD figF PIgAF „he s™—le ˜—r equ—ls I mmF e˜˜revi—E

tionsX ˜pD –˜ump9Y ˜rD ˜—s—lresorptionY d˜™D dist—l˜—s—l™onstri™tionY d™D dist—l™—rin—Y m™D mesi—l™—rin—Y pmD pun™ture m—rkF

QIV h—ll— †e™™hi— pFwF 8 g—u eF

™odyliform genus reported to d—te from the iurope—n the tooth from €ol—zzo shows differen™es with —ll the

teeth reported from eustri—D ƒp—inD —nd ‚om—ni—F „he pper gret—™eousF st is noteworthy th—t the €ol—zzo

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™—rin— —nd —pi™—lh—lfof the mesi—l™—rin— without ™omE ™hi—n er—ripesu™husF „his genus h—s — p—nEqondw—n—n

pletely individu—lized denti™lesF distri˜utionY its monophyly h—s ˜een questioned ˜y yrE

teg— et —lF @PHHHA —nd „urner 8 ƒerti™h @PHIHAD ˜ut w—s

purther ™omp—risons supported ˜y €ol8 epestegui— @PHHSA —nd ƒereno 8

emong gro™odyliformesD — str—ight ™—rin— runE v—rsson @PHHWAF „he sh—red presen™e of m—ny ™h—r—™tersD

ning —pi™o˜—s—lly —long the mesi—l m—rgin of the teeth in™luding —n —ut—pomorphyD suggesting — stri™t —ffinity

is — plesiomorphi™ fe—ture sh—red ˜y most t—x—D where—s

˜etween w€gw IIUPH —nd — qondw—n—n t—xonD h—s

— lingu—l shifting of the mesi—l ™—rin— is — poorly menE

evident p—leo˜iogeogr—phi™ impli™—tionsF roweverD

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sin™e the distri˜ution of — lingu—lly shifted mesi—l ™—rin—

present in the dyros—urid rypos—urus rogersii from the

—mong ™ro™odyliforms m—y h—ve ˜een overlooked in

w——stri™hti—n of xew terseyD ˜ut not in other dyros—urE

liter—tureD the phylogeneti™ me—ning of th—t ™h—r—™ter

idsY henton et —lF IWWUAF e lingu—l shifting of the mesi—l

@for ex—mpleD its in™lusiveness —mong er—ripesu™hus speE

™—rin— o™™urs —lso in ‚—z—n—ndrongo˜e s—k—l—v—e @ƒiE

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mone w—g—nu™oD persF ™omF to eF gFAD —n enigm—ti™

™l—disti™ fr—meworkF purthermoreD the €ol—zzo ™rown

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™—r with possi˜le ™ro™odylomorph —ffinities @w—g—nu™o

the mesi—l ™—rin— without ™ompletely individu—lized

et —lF PHHTAF ‚e™entlyD ƒereno 8 v—rsson @PHHWX QSA reE

denti™lesD — fe—ture yet unreported in other ™ro™odyliE ported ––lingu—lly defle™ted mesi—l ™—rin— —nd —sso™i—ted

formsD —lthough —lso pro˜—˜ly overlooked in liter—tureF trough99 —s —n —ut—pomorphy of the l—rgest post™—niniE

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nitiesF porm—tion @epti—nEel˜i—nA of xigerF „he IIth dent—ry

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figlio PHHSA —nd from the edri—ti™Ehin—ri™ g—r˜on—te mesi—l —nd dist—l ™—rin—eD —nd — ™entr—lD longitudin—l reE

€l—tform of the iurope—n —r™hipel—go —s well @ƒteel lief with — fl—t surf—™e on the lingu—l side @ƒereno 8

IWUQY €ered—Eƒu˜er˜iol— PHHWAF es notosu™hi—ns were v—rsson PHHWD figsF PHgD PIeD PIgAF st differs from

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