‚ivist— st—li—n— di €—leontologi— e ƒtr—tigr—fi— volume IIW noF Q ppF PUSEPVS xovem˜er PHIQ

e xi‡ €i‚vishsh p‚yw „ri ƒ€e„rsex @yvixiusexD ie‚v‰ „‚seƒƒsgA

yp greyr D exr s €‚y†sxgiD grsxe

I P I Q

 y‰ ƒ x D exh‚ie „sx„y‚s D he‰yxq tsexq 8 ‚‰yƒ ui wy„exs

‚e™eivedX tuneD TD PHIQY —™™eptedX y™to˜er PSD PHIQ

uey wordsX xew t—xonD €erleidiformesD i—rly „ri—ssi™D ƒp—thi—nD dors—le ed —n—leF gh—ohuperleidus genF nF r—ppresent— il piuÁ —nti™o

ƒouth ghin—F perleidiforme ™onos™iuto e prov— l9origine dei perleidiformi gi—Á verso

l— fine del „ri—ssi™o snferioreF

e˜str—™tF e new —™tinopterygi—n genus —nd spe™iesD gh—ohuperE

leidus primus genF nF spF nFD is des™ri˜ed from the pper wem˜er of the

x—nlinghu porm—tion @ƒp—thi—nD yleneki—nD i—rly „ri—ssi™A —t w—ji—E

sntrodu™tion sh—n @w—ji— rillAD gh—ohu gityD enhui €rovin™eD ƒouthe—st ghin—F „he

new t—xon is —ssigned to the order €erleidiformes on the ˜—sis of —

„he wellEknown „ri—ssi™ i—rly pish p—un—s ™om˜in—tion of fe—turesX l—rge wedgeElike preoper™ulum with exp—nded

infr—or˜it—l pro™ess —nd the sensory ™—n—l running —lmost verti™—l —long @„ippD „intori et —lF PHIQA of w—d—g—s™—rD qreenl—ndD

the posterior m—rgin of the preoper™ulum itselfY fl—nk s™—les somewh—t

ƒpitz˜ergen —nd ‡est g—n—d— @entunes et —lF IWWHY vehE

higher th—n long in the trunk regionY presen™e of ep—xi—l r—ys in the

m—n IWSPY €ivete—u IWQRY ƒ™h—effer 8 w—ngus IWUTY

™—ud—l finF emong €erleidiformesD the new t—xon ˜elongs to the €erE

ƒtensio IWPID IWQPA —re not younger th—n the endE È leidid—eD ˜eing very simil—r to €erleidus in the gener—l ˜ody sh—peD

ƒmithi—nF „hose —ssem˜l—ges —re ™omposed of primitive p—ttern of the skull ˜onesD dentition m—de of pegElike m—rgin—l teethD

—nd num˜er of ep—xi—l r—ysF „he new genus is ™h—r—™terized ˜y the neopterygi—ns @€—r—semionotiformesA —nd ˜—s—l —™tiE

presen™e of — very l—rge —nteroEdors—l pro™ess on the su˜oper™ulumD

nopterygi—ns th—t in™lude some spe™i—lized t—x— @ƒ—urE

— high num˜er @—˜out SSA of tr—nsvers—l s™—le rows —nd r—ther posterior

i™hthys —nd fo˜—s—tr—ni—A —lre—dy —ppe—red in the l—test

dors—l —nd —n—l finsF h—ting ˜—™k to the ƒp—thi—nD gh—ohuperleidus genF

€ermi—n @viu 8 ‡ei IWVVY „intori 8 vom˜—rdo PHHUAF nF represents the oldest re™ord of the order €erleidiformesF

e few t—x— were —lso —s™ri˜ed to €erleidus following

‚i—ssuntoF †iene des™ritto il nuovo —ttinotterigio gh—ohuperleiE

ƒtensio @IWPIA @see entunes et —lF IWWHY vehm—n IWSPY

È

dus primus genF nF spF nF su esempl—ri rinvenuti nel wem˜ro medio dell—

€ivete—u IWQRAX thoughD —™™ording to vom˜—rdo @PHHIAD x—nlinghu porm—tion @ƒp—thi—noD yleneki—noD „ri—ssi™o snferioreA sulE

they l—™k the ep—xi—l r—ys ™h—r—™terizing the su˜holosE l— gollin— di w—ji— @w—ji—sh—nD gh—ohu gityD €rovin™i— di enhuiD giE

te—nsD so their —ttri˜ution to perleidiforms h—d therefore n—AF sl nuovo t—xon —pp—rtiene —i su˜holostei in qu—nto present— r—ggi

ep—xi—li nell— pinn— ™—ud—leF v— su— —pp—rtenenz— —i €erleidiformi eÁ to ˜e questionedF ell the „ipps r—nge from the sndu—n

™onferm—t— d— un insieme di ™—r—tteri qu—li i denti — pioloD un preoperE

to the end of the ƒmithi—nF „he youngest fishes —re

™olo —mpioD ™on un evidente pro™esso infr—or˜it—leD e ™on il ™—n—le

pro˜—˜ly those found in southe—stern ghin— —t insh—n sensori—le verti™—le lungo il m—rgine posteriore dell9ossoD le s™—glie dell—

in turong gounty @ti—ngsu €rovin™eA —nd w—ji—sh—n regione —nteriore del fi—n™o piuÁ—lte ™he lungheF v9—spetto gener—le del

™orpoD l— struttur— —n—tomi™— del derm—to™r—nioD l— dent—tur— non speE ne—r gh—ohu gity @enhui €rovin™eAX they ™—n ˜e pl—™ed

™i—lizz—t— e il numero di r—ggi ep—xi—li h— permesso l9—ttri˜uzione di

just ˜elow the ƒmithi—nGƒp—thi—n ˜ound—ry @„ong et —lF

gh—ohuperleidus primus genF nF spF nF —ll— f—migli— €erleidid—eF sl nuovo

PHHTAF e few t—x— h—ve ˜een des™ri˜ed from these two t—xon eÁ ™—r—tterizz—to d—ll— presenz— di un —mpio e —rrotond—to proE

m—jor ghinese sites —s well —s from other sp—rse lo™—lE ™esso —nteroEdors—le del su˜oper™oloD un numero elev—to difiledis™—E

glie @™ir™— SSA e d—ll— posizione rel—tiv—mente posteriore delle pinne ities in southern ghin— @ƒu IWVIY ƒu 8 vi IWVQY i—n et

I hep—rtment of qeology —nd qeologi™—l wuseumD €eking niversityD feijing IHHVUID €F ‚F ghin—F

iEm—ilX sunzuoyudpkuFeduF™nD dji—ngdpkuFeduF™nY

P hip—rtimento di ƒ™ienze dell— „err— –eFhesio9D niversit—Ádegli ƒtudi di wil—noD vi— w—ngi—g—lli QRD sEPHIQQ wil—noD st—lyF

iEm—ilX —ndre—FtintoridunimiFitD Q hep—rtment of qeologyD niversity of g—liforni—D h—visD g—liforni— WSTITD FƒFeF iEm—ilX mont—nidgeologyFu™d—visFedu

PUT ƒun FD „intori eFD ti—ng hF 8 wot—ni ‚F

˜een ™olle™ted during the qu—rrying worksF huring our —lF IWWUY viu et —lF PHHPY tin et —lF PHHQY „ong et —lF PHHTY

prelimin—ry PHIH field ™—mp—ign in w—ji—sh—n some fish h—o 8 vu PHHUAD with t—x— often ere™ted on single

rem—insD here des™ri˜edD were ™olle™ted together with spe™imensF „he gener—l —ssem˜l—ge in these ghinese loE

few fr—gment—ry spe™imens of gh—ohus—urusF „he m—E ™—lities is r—ther poor of spe™ies ™omp—red with the

jor field works were ™—rried out in PHII —nd PHIPD when other worldwide sitesF st ™omprises only — few p—r—seE

sever—l spe™imens of reptiles @ti—ng et —lF PHIPA —nd mionotiforms —nd the ™oel—™—nth gh—oui™hthys @„ong et

fishes were ™olle™ted —nd —re now under studyF wost —lF PHHTA other th—n — num˜er of spe™ies now —s™ri˜ed to

fishes ™ome from the uppermost ƒmithi—n nodule level €lesioperleidus ˜y „ong et —lF @PHHTAF „his poverty m—y

@top of the relongsh—n porm—tionAD ˜ut not —ll the ™olE ˜e —s™ri˜ed either to the f—™t th—t in the endEƒmithi—n —

le™ted fishes were preserved in nodulesF e few spe™iE new ™risis deeply —ffe™ted —t le—st the m—rine environE

mens were —lso found in the ƒp—thi—n x—nlinghu porE ment @q—lfettiet —lF PHHU—D ˜A or to — somewh—t different

m—tionD together with gh—ohus—urusF „his represents p—leoenvironmentD sin™e w—ji—sh—n is thought to h—ve

the first fish —ssem˜l—ge of this —geF „hough most m—E ˜een f—rther from the ™o—st th—n other sites likeD for

teri—l is poorly preserved —nd fr—gment—ryD we —re ™onE inst—n™eD the x‡ w—d—g—s™—r lo™—lities @felt—n IWWTAF

fident in —s™ri˜ing it to — very restri™ted num˜er of t—x—F et w—ji—sh—nD — ™omplete i—rly „ri—ssi™ sequen™e

„he —ssem˜l—ge thus —ppe—rs quite s™—ntyX so f—r only @pigF IA w—s exposed in — l—rge qu—rry exploited for ro—d

one perleidid genus —nd two or three nonEperleidid genE ™onstru™tions —nd ™ementEm—king limestone ro™ksF

er— h—ve ˜een prelimin—ry re™ognizedF „he perleidid efter the enhui €rovin™i—l —dministr—tion in PHII

genus is the o˜je™t of the present p—perF stopped the ™ommer™i—l qu—rrying to —llow det—iled

s™ientifi™ rese—r™hesD it w—s possi˜le to investig—te the

whole series ˜ed ˜y ˜edF ƒe—r™h for verte˜r—tes fo™used

w—teri—l —nd methods on the ƒmithi—nGƒp—thi—n ˜ound—ryD where the wellE

known level of fishE˜e—ring nodules ™rops out @„ong et

ƒpe™imens herein des™ri˜ed —re preserved in light grey ™—l™—rE

—lF PHHTAD —s well —s on the middleEupper ƒp—thi—nF rereD eous m—rls whose f—int l—min—tion h—rdly splitsD so th—t the fossils —re

usu—lly visi˜le in se™tion —s the ro™k is ˜rokenF purthermoreD in the —s first reported ˜y ghen @IWVSAD ƒ—uri™hthys —nd few

w—ji—sh—n qu—rry l—yers —re —lmost verti™—lly —rr—ngedY the w—ll left spe™imens of the ˜—s—l i™hthyos—ur gh—ohus—urus h—d

˜y the former ™ommer™i—l qu—rrying is

r—ther highD m—king the fossils diffi™ult

to re—™h —nd ™olle™tF „he m—teri—l used

for this study in™ludes only two ™omplete

spe™imens —nd four fr—gment—ry onesF

€relimin—ry prep—r—tion w—s done ˜y —irE

™hiselsY su˜sequentlyD thin steel tools unE

der stereomi™ros™ope were usedF €hotos

were t—ken ˜y me—ns of xikon hQIHH

™—mer— —nd xikkor pTH mi™rolensF hr—wE

ings were m—de through ™—mer— lu™id— on

— ‡svh wV stereomi™ros™ope —nd l—ter

digit—lized using edo˜e €hotoshop TFH

gr—phi™ softw—reF

e˜˜revi—tionsX qw€u E€D qeoE

logi™—l wuseum €eking niversity p—E

leontologi™—l ™olle™tionsY —ntD —ntor˜it—lY

˜rD ˜r—n™hiosteg—lY ™lD ™leithrumY dhyD derE

mohy—lY dptD dermopteroti™Y dsphD dermoE

sphenoti™Y exs™D extr—s™—pul—rY frD front—lY

ioD infr—or˜it—lY ioprD infr—or˜it—l pro™ess

of the preoper™ulumY mxD m—xill—Y mdD

m—ndi˜leY n—Dn—s—lY opD oper™ulumY p—D

p—riet—lY p™lD post™leithrumY pmxD prem—xE

ill—Y popD preoper™ulumY ptD posttempor—lY

roD rostr—lY s™D s™—leY s™lD supr—™leithrumY

pigF I E eD fA vo™—liz—tion of w—ji—sh—n qu—rry in gh—ohu gityD enhui €rovin™eD ƒouth ghin—F s˜oD su˜or˜it—lY sopD su˜oper™ulumF

~ m—kes the position of w—ji—sh—n u—rryF gA „he w—ji—sh—n u—rry seen from ‡est

tow—rd i—stF „he pi™ture w—s t—kes —lmost perpendi™ul—r to the verti™—l ˜eds of the

relongsh—n porm—tion @foregroundA —nd x—nlinghu porm—tion well visi˜le in the ˜—™kE

ƒystem—ti™ €—l—eontology

ground —nd m—king most of the qu—rry ˜ottomF €eople in the foreground —t left —re —t the

top of the relonsh—n porm—tion in the sm—ll tren™h ™orresponding to the level yielding

„he su˜holoste—ns —re

the fossiliferous nodules with fishesF sn the ˜—™kground —t right the w—ll rem—ining —fter

here ™onsidered —s spe™i—lized the ™ommer™i—l qu—rrying w—s stoppedX from th—t spot ™ome most of the gh—ohus—urus

—nd the —sso™i—ted fish rem—insD mostly gh—ouperleidus genF nF non neopterygi—n —™tinopteryE

e new perleidid from the i—rly „ri—ssi™ of southe—stern ghin— PUU

—nd r—ys of the dors—l —nd —n—l finsD needs to ˜e re™onE gi—ns following the e—rly interpret—tion ˜y frough

sideredF wost su˜holoste—n —nd ˜—s—l neopterygi—n t—x— @IWQID IWQWA —nd ƒ™h—effer @IWSSD IWUQAY thoughD they

whi™h show the endoskeleton of the medi—n finsD iFeF definitely need — ™omplete phylogeneti™ —n—lysisF „his

w—r™opoloi™hthysD qymnoi™hthysD uyphosi™hthysD €l—™oE is —n interesting fish group th—t ˜loomed in the „ri—ssi™

pleurusD „hor—™opterusD puyu—nperleidusD €eltopleurusD se—s sin™e the l—test i—rly „ri—ssi™D —s proved in this

indi™—te th—t the put—tive oneEtoEone rel—tionship is p—perF „hey show somewh—t derived ™h—r—™ters in the

not tot—lly —™hieved ˜e™—use —t le—st the l—st r—di—l of post™r—ni—l skeleton respe™t to the p—leonis™iforms for

˜oth fins —rti™ul—tes two to five r—ys @„intori et —lF inst—n™e @™—ud—l fin more or less symmetri™—l —nd reE

PHHVD PHIHD PHIPY ƒun et —lF PHIPY persF o˜sFAF purtherE du™ed num˜er of r—di—ls in the dors—l —nd —n—l finsA

moreD the su˜holoste—ns —nd the ˜—s—l non teleoste—n ˜eside — more ™onserv—tive derm—l skull p—ttern where

neopterygi—ns —™quire the extern—l symmetry of the the m—xill— rem—ined fixed to the preoper™ulum —nd the

™—ud—l fin through different w—ysF sn the former the interoper™ulum w—s still missingF sn the l—st ye—rsD su˜E

˜ody lo˜e ˜e™omes very shortD ˜ut not fully dors—l —s holoste—ns h—ve ˜een usu—lly ™onsidered either in this

the verte˜r—l ™olumn does not sh—rply ˜endD so th—t s—me w—y @furgin IWWPY vom˜—rdo PHHIY vom˜—rdo 8 È

ep—xi—l @dors—lly insertedA r—ys —re present @hemiheteE „intori PHHRY vom˜—rdo et —lF PHHVA or —s neopterygi—ns

ro™er™—l ™—ud—l finAF sn neopterygi—ns the ˜ody lo˜e ˜eE or stem group neopterygi—ns @q—rdiner et —lF PHHSY rurE

™omes shorter @—˜˜revi—ted hetero™er™—l ™—ud—l finA with ley et —lF PHHUY ˆu 8 ‡u PHIPAF „he l—tter hypothesis is

—ll the r—ys inserted ventr—l to the verte˜r—l ™olumnF m—inly due to the —lleged free m—xill— of vug—noi— @furE È

enyw—yD the thorough interpret—tion —nd phylogeneti™ gin IWWPAF roweverD vug—noi— h—s — r—ther spe™i—lized

—n—lysis of su˜holoste—ns —re f—r from ˜eing settledD —nd over—ll skull p—ttern —nd its m—xill— is not tot—lly free

—re ˜eyond the —im of this p—perY we refer to ƒun et —lF from the preoper™ulumD espe™i—lly if the postm—xill—ries

@PHIPA for the €erleidiformes interrel—tionshipsF of furgin @IWWPA —re ™onsidered —s —n—mesti™ p—rt of the È

€erleidiformesD together with €eltopleuriformesD preoper™ulumF woreoverD the sep—r—tion of the m—xill—

—re the l—rgest su˜holoste—n ordersF „hey —re ™h—r—™terE from the preoper™ulum —lso ™on™erns other ˜—s—l —™tiE

ized ˜y fl—nk s™—les deeper th—n longD verti™—l or —lmost nopterygi—nsD su™h —s fo˜—s—tr—ni— @xielsen IWSPA or

verti™—l preoper™ulumD usu—lly still ™onne™ted to the —nother su˜holoste—nD r—˜roi™hthys @vin et —lF PHIIAF

posterior end of the m—xill— ± ˜ut see vin et —lF @PHIIA roweverD —ll the —˜ove mentioned —™tinopterygi—ns l—™k

for r—˜roi™hthys —nd furgin @IWWPA for vug—noi— ± l—™k the interoper™ulumD whose —ppe—r—n™e w—s ™onsidered È

of the interoper™ulum —nd ˜y the presen™e of ep—xi—l ˜y €—tterson @IWUQA —s dire™tly rel—ted to the sep—r—tion

r—ys in the ™—ud—l finF „he me—ning of su˜holoste—n of the m—xill— from the preoper™ulum —mong neopteE

fishes is f—r from ˜eing fixed —nd no˜ody ever —ttempted rygi—nsF enother widelyE—™™epted neopterygi—n —poE

— gener—l phylogeneti™ —n—lysisF „hough — few different morphyD the oneEtoEone rel—tionship ˜etween r—di—ls

pigF P E gh—ohuperleidus primus genF

nF spF nF eA „he holotypeD

spe™imen qw€u E€EIIPHF

fA ƒpe™imen qw€u E€E

QHURF x—nlinghu porm—tion

@ƒp—thi—nD i—rly „ri—ssi™A of

w—ji—sh—n qu—rryD gh—ohu

gityD enhui €rovin™eD ghin—F ƒ™—le ˜—rs a IH mmF

PUV ƒun FD „intori eFD ti—ng hF 8 wot—ni ‚F

th—t looks slightly ™onvexF „he medi—n region is squ—rE orders h—ve ˜een —ssigned to the su˜holoste—ns during

the l—st UH ye—rsD we think —t present only €erleidiformes ishY —long the l—ter—l m—rgins there —re two not™hes for

—nd €eltopleuriformes —re thoroughly ™onsistent with the —nterior nostrilsF „he dors—lmost p—rt seems to ˜e

the —˜ove ™ited ™h—r—™ters —nd —re herein ™onsideredF n—rrower —nd the dors—l m—rgin —lmost semi™ir™ul—rF

„he rostr—l is well orn—mented ˜y round tu˜er™lesD of

whi™h those lo™—ted ˜etween the nostrils —re mu™h l—rE

ger th—n the othersF „he n—s—ls —re re™t—ngul—r —nd gl—ss ystei™hthyes ruxleyD IVVH

˜order the rostr—l —nd —re somewh—t shorterF „he senE

ƒu˜™l—ss e™tinopterygii gopeD IVVU

sory ™—n—l ™—n ˜e tr—™ed ˜y sever—l sm—ll poresX it enE

yrder €erleidiformes fergD IWQU

tered the n—s—l ™lose to the —nteroEmedi—l ™ornerD then

p—mily €erleidid—e froughD IWQI ™rossed the ˜one o˜liquely tow—rd the l—teroEdors—l ™orE

nerF elso the n—s—ls —re orn—mented ˜y s™—ttered tu˜erE qenus gh—ohuperleidus genF nF

™lesF

hi—gnosisX es for the type —nd only known spe™iesF „he p—ired front—ls —re re™t—ngul—r in sh—peD with

itymologyX prom gh—ohuD the ™ity whi™h the w—ji—sh—n qu—rE

— sh—llow em˜—yment —t the or˜it levelX their length is ry ˜elongs toD —nd where the studied spe™imens ™ome fromD —nd from

—˜out twi™e the m—ximum widthF „he medi—n suture

€erleidusD the type genus of the su˜holoste—n €erleidid—eF

seems str—ightD while the suture with the p—riet—ls is

quite indentedF „he supr—or˜it—l sensory ™—n—l w—s deeE

ply em˜edded in the ˜one —nd it g—ve rise to — ridge on gh—ohuperleidus primus genF nF spF nF

the inner surf—™eX it entered the front—ls —lmost in the pigs PEU

middle of e—™h ˜oneF „he surf—™e shows sm—ll —nd s™—tE

hi—gnosisX @e unique ™om˜in—tion of primitive —nd derived

tered tu˜er™lesF

™h—r—™tersY B—ut—pomorphiesFA wediumEsized perleidid @up to IPS mm

„he p—ired p—riet—ls —re squ—rishX they seem to ˜e in st—nd—rd lengthA with — quite elong—te fusiform ˜odyF €reoper™ulum

very l—rge l—ter—llyD so th—t it possi˜ly fused with the with n—rrowD elong—ted infr—or˜it—l pro™essD ™overed ˜y infr—or˜it—lsF

dermopteroti™sD even if we ™—n see no tr—™e of the infr—E ƒu˜oper™ulum with l—rgeD roundedD —nteroEdors—l pro™essBF herm—l

skull ˜ones orn—mented with s™—ttered sm—ll round tu˜er™lesD quite or˜it—l sensory ™—n—lF „he supr—or˜it—l sensory ™—n—ls

l—rger on the j—wsF e˜out SS tr—nsvers—l s™—leErows —long the l—ter—l

entered the ˜one —t —˜out IGQ from the l—ter—l m—rgin

lineBF enterior fl—nk s™—les —t le—st twi™e deeper th—n longF ƒ™—les denE

—nd re—™hed —t le—st the midElengthF xothing ™—n ˜e s—id ti™ul—ted on the posterior m—rginY —nterior s™—les showing f—int longE

—˜out the —™tu—l outline of the extr—s™—pul—rsD ex™ept itudin—l ridgesF v—rge s™—le in front of the —n—l finF pield of sm—ller

th—t their sh—pe w—s pro˜—˜ly su˜tri—ngul—rD —s shown s™—les in front of the ventr—l lo˜e of the ™—ud—l finBF ƒensory ™—n—l of

the l—ter—l line with few tr—nsvers—l pores opening in the dors—l h—lf of in spe™imen qw€u E€E QHUSF

the s™—lesF €e™tor—l fin m—de of IS r—ysD pelvi™ m—de of tenF hors—l fin

i—™h upper j—w is m—de of — m—xill— —ndD most

short —nd m—de of IP r—ysBF hors—l fin inserted in the posterior h—lf of

pro˜—˜lyD — prem—xill—Y thoughD this l—tter h—s never the ˜odyD st—rting —t —˜out UH7 of the sF lFBF en—l fin inserted —t —˜out

˜een found in —n—tomi™—l ™onne™tion ˜ut only —s UV7 of the sF lFBF „en to eleven ep—xi—l r—ysD seven of themD segmented

toothed fr—gments —round the —nterior end of the m—xE ˜ut un˜r—n™hedD ˜e—ring sm—ll fringing ful™r—BF €terygi—l formul—X

ill—F e possi˜le prem—xill—ry ˜one is visi˜le in spe™imens hQS

SS

qw€u E€E QHUSX there —re —t le—st six teeth ˜eing the € IUEIV e QHEQI g RU

s—me size —s those of the m—xill—F „he m—xill— shows —

itymologyX prom the v—tin word primusD me—ning firstD —s this

low —nterior p—rt ˜e—ring — minimum of IR pegElike is the first @—nd oldestA true perleidid des™ri˜ed from i—rly „ri—ssi™F

rolotypeX qw€u E€EIIPHD — ™omplete spe™imen with st—nE teethD ˜—™kw—rd de™re—sing in lengthD —nd — very high

d—rd length of IPS mmF

tri—ngul—r posterior pl—teD with —n —lmost str—ight or—l

€—r—typesX qw€u E€EIIPID QHURD QHUSD QHUTD QHUUF

m—rginF „he —nterior tip is smooth —nd — little depressedD

„ype horizon —nd lo™—lityX „he pper wem˜er of the x—nlinE

—s if it yielded the prem—xill—X six teeth —re ˜orne in this ghu porm—tion @ƒp—thi—nD i—rly „ri—ssi™A of w—ji—sh—n qu—rryD gh—ohu

region —nd they possi˜ly l—y medi—lly to the prem—xill— gityD enhui €rovin™eD ghin—F

teethF „he rem—ining p—rt of the m—xill— is ™overed ˜y

l—rge —nd dense tu˜er™lesF hes™riptionF „he st—nd—rd length of the ™omplete

„he lower j—ws —re elong—teD with —n —lmost spe™imens is IIS mm —nd IPS mm —nd the skull is QP

str—ight or—l m—rginF ywing to the poor preserv—tion —nd QS mm long respe™tivelyF elso the fr—gment—ry speE

—nd the dense orn—ment—tionD their single ˜ones ™—nnot ™imens should h—ve — simil—r size —s — ™omplete skull

˜e dete™tedF „eeth —re —pproxim—tely —ll the s—me @qw€u E€EQHUSA me—sures QT mm in lengthF „hey —re

lengthD ˜ut the posterior elements —re stouter th—n the r—ther elong—te fusiform fishesX the r—tio ˜etween the

—nterior onesF „hese l—tter —re simil—r in size —nd sh—pe m—ximum ˜ody depth —nd the tot—l length is —˜out

PS7F to the —nterior elements of the upper j—wsD ˜ut their tips

ƒkullF „he medi—n su˜Eex—gon—l rostr—l is someE ˜end slightly inw—rdD giving these teeth — sigmoid—l

wh—t longer th—n wideD with — rounded ventr—l p—rt sh—peF „here —re —t le—st PP pegElike teeth on e—™h side

e new perleidid from the i—rly „ri—ssi™ of southe—stern ghin— PUW

pigF Q E gh—ohuperleidus primus genF nF spF nF eA qw€u E€EQHUSD fish IF fA hr—wing of the skull of the s—meF ƒ™—le ˜—r a IH mmF

pigF R E gh—ohuperleidus primus genF nF spF nF eA ƒpe™imen qw€u E€EQHUTF fA hr—wing of the s—meF gA ƒpe™imen qw€u E€E QHUUF „he

—rrow points to the l—rge —nterodors—l pro™ess of the su˜oper™ulumF ƒ™—le ˜—rs a S mmF

of the lower j—wsD ˜ut further inner tooth rows were gin gently roundedF „he position of the sensory ™—n—l is

pro˜—˜ly presentX teeth fr—gments —re visi˜le in ˜etween m—rked ˜y very sm—ll pores —t the re—r edge of the ˜oneX

the teeth of the extern—l rowF it r—n verti™—l —nd gently ˜ent forw—rd in the dors—lmost

„he elements of the ™ir™umor˜it—l series —re p—rE p—rtF „he surf—™e is ™ompletely smooth ex™ept for very

ti—lly preserved on spe™imen qw€u E€E QHUT @pigF RA r—re sm—ll g—noine tu˜er™les

where — ™res™entEsh—ped jug—l infr—or˜it—l is lo™—ted —t „he oper™ulum is m—de of ˜ro—d oper™ul—r —nd

the posteroventr—l ™orner of the or˜itF „he sensory ™—n—l su˜oper™ul—r ˜onesD the former ˜eing only somewh—t

r—n ™lose to its —nterior m—rginF sn this spe™imen there l—rger th—n the l—tterF „he suture ˜etween the two eleE

—re three more ˜one fr—gments dors—lly to the jug—l ments is —nteroEventr—lly gently forw—rd in™lined —nd —

infr—or˜it—lX on the grounds of their position they m—y l—rge —nd rounded —nteroEdors—l pro™ess is well dete™tE

˜e reg—rded —s p—rts of the postor˜it—ls —ndGor the derE —˜leF „he surf—™e of the two ˜ones shows only very r—re

mosphenoti™F sm—ll g—noine tu˜er™lesF „en ˜r—n™hiosteg—l r—ys —re

„he preoper™ulum em˜r—™es the posterior pl—te of pl—™ed ventr—lly to the su˜oper™ulumF sn the first three

the m—xill—X its ventr—l region is tri—ngul—r —nd in spe™iE r—ys the —nteroEventr—l edge overl—ps the pre™eding r—yY

men qw€u E€EQHUT there is — wellEdeveloped infr—E the fourth r—y is overl—pped on ˜oth sides ˜e™—use there

or˜it—l pro™ess with — rounded —nterior tipF fe™—use is — revers—l of the overl—pping so th—t in the five —nteE

most of the pro™ess is smoothD it w—s pro˜—˜ly ™overed rior r—ys the most —nteroEventr—l edge is ™overed ˜y the

˜y the jug—l element of the infr—or˜it—ls seriesF „he dorE posterior m—rgin of the pre™eding r—yF xo ™le—r tr—™es of

s—l region is mu™h more exp—ndedD with the dors—l m—rE — gul—r element ™—n ˜e dete™tedF

PVH ƒun FD „intori eFD ti—ng hF 8 wot—ni ‚F

qirdlesF „he ™leithrum is —lmost tot—lly hidden ˜y

the oper™ul—r ˜onesD ex™ept for some —re—s ™overed ˜y

g—noine ridgesD dorsoEventr—lly elong—tedF ƒupr—™leiE

thrum is quite l—rge —nd there —re pro˜—˜ly two postE

™leithr—l ˜onesX —ll of them h—ve — few g—noine ridges

—nteroE™—ud—lly dire™tedF

€—ired —nd medi—n finsF ell the fins yield sm—ll

fringing ful™r—X those on the dors—l edge of the ™—ud—l fin

—re ˜orne ˜y —t le—st seven segmented r—ysD whi™h is —

quite unusu—l high num˜er of fringing ful™r— yielding

r—ysF

„he pe™tor—l fins h—ve — tri—ngul—r outlineY e—™h

fin is m—de of —˜out IS r—ysD showing — long unsegmenE

ted proxim—l ˜—se —nd sever—l dist—l segmentsF

„he pelvi™s —re sm—llD tri—ngul—r in outlineF „hey

—re m—de of only TEU r—ys —nd inserted —t —pproxim—tely

th

s™—le rowD —˜out h—lf of the ˜ody tot—l lengthF the IT

„he dors—l fin is inserted r—ther posteriorlyD well

in the se™ond h—lf of the ˜odyD —round the UH7 of the sF

th

s™—le rowF st is sm—llD its sh—pe is — lFD —t the QREQS

tri—ngle with — short ˜—seF st ™omprises only IHEII

r—ther short lepidotri™hi— whi™h ˜r—n™h on™e or twi™eF

„he —n—l fin is —lso sm—ll in size —nd tri—ngul—r in

sh—peF st h—s —lmost the s—me num˜er of r—ys —s the

dors—l oneD ˜ut they —ppe—r mu™h stouter —nd possi˜ly

˜r—n™h up to three timesF st is posterior to the dors—l —s

it inserts —round UV7 of the sF lFD even if it st—rts —t —˜out

th

s™—le rowF the QH

„he ™—ud—l fin is —lmost symmetri™—l extern—lly

—nd h—s —˜out QR segmented r—ysD of whi™h IHEII —re

™onsidered —s ep—xi—ls r—ys @pigF SAF „here —re —t le—st

seven short segmented fin r—ysD ˜oth dors—l —nd ventr—l

to the le—ding fin r—ysF ip—xi—l —nd m—jor ventr—l r—ys

—re —˜utting to e—™h otherD where—s two or three medi—n

r—ys —re ˜ro—dly sp—™edF „he g—noine ™overing of the

lepidotri™hi— is heterogeneousX p—t™hes —re present on

the proxim—l segments ˜ut not dist—llyF

ƒqu—m—tionF et le—st SS tr—nsvers—l s™—le rows —re

presentD ™ounted —long the l—ter—l line ™—n—lF „he —nteE

rior fl—nk s™—les —re twi™e —s deep —s long ˜oth dors—lly

—nd ventr—lly to the deepest s™—les of the row where the

l—ter—l line r—nF ell the s™—les h—ve — denti™ul—ted posterE

ior m—rginX —˜out IS denti™les in the highest —nterior

fl—nk s™—lesD —˜out five —t the ˜eginning of the ™—ud—l

region —nd one or two just ˜efore the ™—ud—l finF enteE

rior s™—les show f—int ridges in ™orresponden™e with

e—™h denti™leX thoughD the ridges dis—ppe—r —fter only

five to seven tr—nsvers—l rowsD —nd ˜eyond th—t point

most s™—les —re smoothD just ™overed ˜y — thin l—yer of

g—noineF e l—rge ov—l —n—l s™—le is present in front of the

—n—l finF e short field of sm—ller s™—lesD ™omp—red to

pigF S E gh—ohuperleidus primus genF nF spF nF g—ud—l finF eA „he

those —round themD is —t the ˜—se of the t—il ventr—l lo˜eF

holotype qw€u E€EIIPHF fA „he dr—wing of the s—meF

„he short ˜ody lo˜e ends with two rows of nine re™E gA ƒpe™imen qw€u E€EQHURF errows point to ep—xi—l

r—ysF ƒ™—le ˜—rs a S mmF t—ngul—r s™—les e—™hD well distin™t from the pre™eding

e new perleidid from the i—rly „ri—ssi™ of southe—stern ghin— PVI

rhom˜i™ onesF „he pores of the l—ter—l line sensory ™—n—l q—˜—nellid—e show very thin s™—les —rr—nged in

—re very few —nd they —ppe—r —s verti™—l slitsF very numerous rows —nd — pred—tory dentition @„intori

his™ussionF gh—ohuperleidus primus genF nF spF nF 8 vom˜—rdo IWWTAX ˜oth ™h—r—™ters —re —˜sent in the

displ—ys fe—tures typi™—l of the su˜holoste—ns —s —˜ove new t—xonF „he m—jor syn—pomorphy of vug—noid—e is

definedD su™h —s the presen™e of — verti™—l or —lmost — unique p—ttern of the ™heek regionD where the preoE

verti™—l preoper™ulum still ™onne™ted to the posterior per™ulum is still in ™ont—™t with the posterior region of

end of the m—xill—D the l—™k of —n interoper™ulumD the the m—xill—D even if for — short length @furgin IWWPAF È

presen™e of ep—xi—l r—ys in the ™—ud—l finD —nd fl—nk gh—ohuperleidus genF nF shows — wide ™ont—™t ˜etween

s™—les deeper th—n longF the preoper™ulum —nd the posterior edge of the m—xill—F

emong su˜holoste—nsD it is ™le—rly more simil—r to €seudo˜e—™oniid—e @voÂpezEer˜—rello 8 —v—ttieri

€erleidiformes th—n to €eltopleuriformesD owing to its PHHVA h—ve — gently rounded posterior edge of most

dentition m—de of pegElike m—rgin—l teethD l—rge —nd s™—les —nd spinous midEdors—l s™—lesD ™ontr—sting with

wedgeElike preoper™ulumD thi™k fl—nk s™—les just deeper the su˜re™t—ngul—r —nd pe™tin—ted s™—les of the new t—xE

th—n long in the trunk regionD numerous @—t le—st tenA on herein des™ri˜edF golo˜odontid—e —re represented ˜y

ep—xi—l r—ysF „he order €eltopleuriformes on the other l—rge fishesD even longer th—n SH ™mD with — r—ther ™onE

h—nd is ™h—r—™terized ˜y minute teeth or no teeth —t —llD serv—tive derm—l skull p—tternD su™h —s the presen™e of —

n—rrower —nd verti™—l preoper™ulumD fl—nk s™—les typiE sm—ll rostr—l followed ˜y — l—rge postrostr—l sep—r—ting

™—lly thinD n—rrow —nd mu™h deeper th—n long —long n—s—l ˜onesD —nd pe™uli—r s™—le orn—ment—tion @ƒun et —lF

most of the ˜ody fl—nkF PHHVAF xo postrostr—l h—s ˜een dete™ted in gh—ohuperE

„he f—milies ™urrently re™ognized in the order leidus genF nF

€erleidiformes —re gleithrolepid—eD €olz˜ergid—eD q—˜—E puyu—nperleidid—e ™omprise sm—ll perleidiforms

nellid—eD vug—noid—eD €seudo˜e—™oniid—eD golo˜odontiE with — pe™uli—r squ—m—tion p—tternX the num˜er of

d—eD €erleidid—e —nd puyu—nperleidid—e @vom˜—rdo et s™—les in e—™h verti™—l row in™re—ses ˜ehind the pelvi™

—lF PHIIY ƒun et —lF PHIPAF fins ˜y —ddition of sever—l s™—les ventr—l to the l—ter—l

„he new t—xon is different from gleithrolepid—e line s™—le row @ƒun et —lF PHIPAF gh—ohuperleidus genF nF

—nd €olz˜ergid—e @vom˜—rdo et —lF PHHVAD whi™h —re evidently differs from this l—st f—milyD —s it h—s no —ddiE

deepEfusiform or deepE˜odied perleidiformsF gleithroE tion—l longitudin—l s™—le rowsF

lepid—e —re ™h—r—™terized —lso ˜y — tri—ngul—r m—xill—D — €erleidid—e is ˜y f—r the most diverse f—mily withE

n—rrow lower j—w devoid of teeth —nd the —˜sen™e of in its orderF fesides the type genusD €erleidusD it ™ont—ins

posttempor—l @rut™hinson IWUQY q—rdiner IWVVAD while sever—l gener—D r—nging from the widdle „ri—ssi™ to the

€olz˜ergid—e show the prem—xill— fused to the —nteroE xori—nX weridensi—D eetheodontusD €eltoperleidusD hipE

dors—l edge of the m—xill— —nd the presen™e of long teronotusD gtenogn—thi™hthysD h—nini—D indenni— @furE È

pro™um˜ent —nterior teeth @qriffith IWUUY vom˜—rdo gin IWWPY wutter 8 rerzog PHHRY „intori IWWHY „intori

8 „intori PHHRY vom˜—rdo et —lF PHHVY ƒun et —lF IWWVY vom˜—rdo PHHIY vom˜—rdo 8 fr—m˜ill—s™—

PHHWAF xone of these ™h—r—™ters —re present in gh—ohuE PHHSAF „he new t—xon here des™ri˜ed is m—nifestly difE

perleidus genF nF ferent from —ll the ™ited t—x—F gomp—red to weridensi—D

pigF T E gh—ohuperleidus primus genF

nF spF nF ƒpe™imen qw€u E

€EQHUUF „he region ˜etween

pelvi™ fins @to the rightA —nd

—n—l fin @to the leftA showing

the —n—l s™—lesF ƒ™—le ˜—r a S mmF

PVP ƒun FD „intori eFD ti—ng hF 8 wot—ni ‚F

pigF U E gh—ohuperleidus primus genF

nF spF nF ƒpe™imen qw€u E

€EQHURF „he —nterior trunk

s™—les to show the posterior

denti™ul—tion —nd the thin

longitudin—l ridges on their

surf—™eF ƒ™—le ˜—r a S mmF

this genus rem—rk—˜ly differs in the sh—pe of ˜oth preE m—tely four more th—n €erleidus —ltolepisD the type speE

oper™ulum —nd m—xill—ryD —nd in the morphology of ™ies of the genus @vom˜—rdo PHHIAF

s™—les in the dors—l regionD —˜ove the deepest onesF e yther gener— des™ri˜ed —s perleididsD su™h —s €leE

p—rti™ul—r squ—m—tionD m—de of minute s™—lesD is ™h—E sioperleidus @ƒu 8 viD IWVQAD h—ngin— @viu et —lFD PHHPAD

r—™teristi™ —lso of eetheodontusD whi™h moreover h—s — €—r—perleidus @h—o 8 vuD PHHUA from the preEƒp—thi—n

pe™uli—r dentition m—de of very sm—ll teeth on the or—l i—rly „ri—ssi™D l—™k ep—xi—l r—ys in the ™—ud—l finF vomE

m—rgin of the j—ws —nd —lso on the p—l—t—l ˜onesD where ˜—rdo @PHHIA h—d ™le—rly shown th—t the —˜sen™e of ep—xE

they —re mostly low —nd stoutF €eltoperleidus is differE i—l r—ys prevent —ny t—xon from ˜eing —s™ri˜ed not only

ent in the skull roofD m—de of — single fused ˜one pl—teD to the €erleidid—e ˜ut —lso to the €erleidiformesY thus

in the p—ttern of ™heek ˜onesD in the sh—pe of the posterE we ™—nnot ™omp—re them with our new m—teri—lF

ior region of the m—xill— —nd in the dentition @furgin È ell these eviden™es support the ere™tion of — new

IWWPY wutter 8 rerzog PHHRAF iven more evident is the t—xon inside the €erleidid—eD ˜e™—use it is more simil—r

differen™e with hipteronotusD whose ˜ody is r—ther deep to €erleidus th—n to —ny other genus of the order €erE

—nd ˜e—ring — dors—l ridgeEs™—les row proje™ting —˜ove leidiformesF

the humpF foth gtenogn—thi™htys —nd indenni— present

— highly spe™i—lized dentitionD ˜esides rem—rk—˜le difE

feren™es in the skull p—ttern @espe™i—lly in the preoperE gon™lusions

™ul—r sh—peA @„intori IWWHY „intori IWWVY vom˜—rdo 8

„he su˜holoste—ns were previously supposed to fr—m˜ill—s™— PHHSAF elso the sm—ll —nd fusiform h—niE

˜e the first —™tinopterygi—ns to r—di—te intensively —fter ni— is different in the gener—l ˜ody morphology —nd

the endE€ermi—n ™risisD in v—te enisi—nGi—rly v—dini—n squ—m—tion —nd in the kind of dentition @vom˜—rdo

of widdle „ri—ssi™D followed ˜y the neopterygi—ns l—ter PHHIAF

onD in the xori—n @„intori IWWVAF ividen™e to this hyE yn the ™ontr—ryD gh—ohuperleidus spF nF is simil—r

pothesis m—inly ™—me from the i™hthyof—un—s of the to €erleidusY the two t—x— show — ™omp—r—˜le p—ttern of

western „ethysD espe™i—lly those yielded ˜y the fes—no the derm—l skull ˜ones —nd — simil—r dentitionD whi™h is

porm—tion —nd the weride vimestoneD in the wonte ƒ—n of — nonEspe™i—lized kind in ˜oth ™—sesF ƒ™—les sh—re —

qiorgio —re—F ‚e™entlyD new finds in the €—nxi—n @quizE well denti™ul—ted posterior m—rgin —nd f—int longitudiE

hou €rovin™eA —nd vuoping @‰unn—n €rovin™eA gounE n—l ridges on the s™—les of the most —nterior rowsF

tiesD indi™—te th—t ˜oth neopterygi—ns —nd su˜holoste—ns xeverthelessD we ™—n o˜serve some differen™es in the

intensively r—di—ted during the widdleEv—te enisi—n skullD m—inly in the su˜oper™ulumD whi™h h—s — l—rge

@ƒun et —lF PHHVD PHHWD PHIPY „intori et —lF PHHVD PHIHY —nd rounded —nteroEdors—l pro™essD tot—lly l—™king in

vin et —lF PHIIY voÂpezEer˜—rello et —lF PHIIY vom˜—rdo €erleidusF „he m—jor differen™es ™on™ern the ˜odyF

et —lF PHIIY ‡en et —lF PHIPY ˆu 8 ‡u PHIPA even with „he gener—l outlineD in f—™tD is more elong—ted in the

sever—l morphologi™—lly spe™i—lized formsD on the whole new t—xon th—n in €erleidusD —nd its dors—l fin is mu™h

m—king the „ri—ssi™ widdle pish p—un— @„wppD „intori further posteriorly inserted @UH7 of the sFlF vs SU7AF

et —lF PHIQAF roweverD we do not know of —ny good l—te „he —n—l fin st—rts —˜out five s™—le rows ˜efore the ˜eE

i—rly „ri—ssi™ or i—rly enisi—n fishEf—un—sX this timeE ginning of the dors—l finD while in €erleidus they —re

g—p needs to ˜e fully understood in order to definitely inserted more or less —t the s—me rowF pin—llyD the new

st—te when the „wpp —™tu—lly ˜loomed @„intori et —lF t—xon h—s IHEII ep—xi—l r—ys in the ™—ud—l finD —pproxiE

e new perleidid from the i—rly „ri—ssi™ of southe—stern ghin— PVQ

the new ˜ed ˜y ˜ed ex™—v—tion in the w—ji—sh—n qu—rryD PHIQA —fter its —ppe—r—n™e in the ƒp—thi—nD proved ˜y this

is fund—ment—l to improve the p—leontologi™—l re™ordF new €erleidiformes genusF gert—inlyD the gh—ohu fish

pin—llyD we w—nt to emph—size th—t the rising of —ssem˜l—ge from the ƒp—thi—n x—nlinghu porm—tion is

the „wpp h—ppened together with the —ppe—r—n™e of still poorly knownF purther field ™olle™ting will ˜e neE

sever—l m—rine reptile groupsD just —˜out Q million ye—rs ™ess—ry to le—rn the —™tu—l ™omposition of this fish f—un—F

—fter the €G„r ™risisF yn™e moreD our models of r—di—tion —ndGor evoE

lution inside — group su™h —s the —™tinopterygi—ns fishes e™knowledgmentsF „his study w—s fin—n™i—lly supported ˜y

@„intori IWWVY „intori et —lF PHIQA prove to ˜e deeply xƒpg @€roje™t RHUHPHHID RHWPHIPRHHPAD ƒt—te uey v—˜or—tory of €—E

l—eo˜iology —nd ƒtr—tigr—phy @x—njing snstitute of qeology —nd €—E influen™ed ˜y the preserv—tion windows —nd ˜y the disE

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