‚ivist— st—li—n— di €—leontologi— e ƒtr—tigr—fi— volume IIV noF Q ppF QSWEQUQ xovem˜er PHIP

p ‰ ex€i‚vish ƒ hixqs qixq i„ evFD PHIP @yƒ„isgr„r‰iƒD

eg„sxy€„i‚‰qssA p‚yw „ri wshhvi „‚seƒƒsg yp ‰ xxex €‚y†sxgiD

ƒy „r grsxe

I P P I

 y‰ ƒ x D g‚sƒ„sxe vywfe‚hy D exh‚ie „sx„y‚s D he‰yxq tsexq D

I I I

‡isgrixq rey D ‰ exvsx ƒ x 8 r xsx vsx

‚e™eivedX w—y PID PHIPY —™™eptedX tuly QD PHIP

ro˜usti denti — piolo ™on un —pi™e —™umin—toA d—lle —lte s™—glie medioE uey wordsX xew t—xonD e™tinopterygi—nsD €erleidiformesD widE

l—ter—li e l— pinn— ™—ud—le ™he present— r—ggi ep—xi—liF uesto t—xon dle „ri—ssi™D ƒouth ghin—F

mostr— tutt—vi— un— disposizione uni™— delle s™—glie @qui ™onsider—t—

e˜str—™tF „wo well preserved spe™imens of puyu—nperleidus ™ome —ut—pomorfi—AD d—t— d—ll9inserzione di s™—glie supplement—ri in

dengi qeng et —lFD PHIP —re des™ri˜ed from the pper wem˜er of the ™i—s™un— fil— verti™—le posteriore —lle pinne pelvi™heD —ppen— sotto le

qu—nling porm—tion @€elsoni—nD enisi—nD widdle „ri—ssi™A in vuoping s™—glie dell— line— l—ter—leD dove l— loro —ltezz— diminuis™e ˜rus™—menteF

gountyD ‰unn—n €rovin™eD ƒouth ghin—F „he —ttri˜ution to €erleidiE €er questo ™—r—ttere presente solo in questo t—xonD e nonost—nte l—

formes is ™onfirmed —™™ording to — ™om˜in—tion of fe—tures su™h —s the re™ente des™rizione del genereD si eÁritenut— ne™ess—ri— l9istituzione dell—

gener—l skull p—tternD with the l—rge preoper™ulum showing — promiE nuov— f—migli— puyu—nperleidid—eY vengono inoltre fornite nuove indiE

nent infr—or˜it—l pro™essD the m—xill— with deep posterior region —nd — ™—zioni sulle tendenze evolutive e le spe™i—lizz—zioni morfologi™he dei

dentition m—de of long —nd stout pegElike teeth with —n —™umin—te —pexD ––su˜olostei99F n9—n—lisi ™l—disti™— dei perleidiformi gi—Á notiD in™luso il

the deep midEl—ter—l fl—nk s™—les —nd the ™—ud—l fin with ep—xi—l fin r—ysF t—xon in oggettoD support— un— monofili— possi˜ile dell9ordineD e si

roweverD this t—xon shows —n unique —rr—ngement of the squ—m—tion —d—tt— perfett—mente —lle rel—zioni p—leo˜iogeogr—fi™he dei t—x— d9—™qu—

@here ™onsidered —s —ut—pomorphyA given ˜y the insertion of sever—l dol™e del qondw—n—F s perleidiformi spe™i—lizz—ti — ™orpo —lto del

s™—les ine—™h verti™—l row posterior to the pelvi™ finDjust ˜elow the qondw—n— e dell— „etide si sono evoluti in modo indipendenteD ™ome

s™—les ™—rrying the l—ter—l lineD when their depth de™re—ses —˜ruptlyF —™™ert—to d—ll9—n—lisi di p—rsimoni—F

ywing to this unique ™h—r—™terD —nd in spite of the very re™ent des™ripE

tion of this genusD the —uthors ™onsidered ne™ess—ry the ere™tion of —

new f—milyD puyu—nperleidid—e offering —lso new insights on the evoluE

sntrodu™tion tion—ry trends —nd morphologi™—l spe™i—liz—tions of the ––su˜holosE

te—ns99F e ™l—disti™ —n—lysis of the well known perleidiformesD with this

„he term ––su˜holoste—ns99 w—s ever ™onsidered —s t—xon in™ludedD supports — possi˜le monophyly of the orderD —nd

—n inform—l t—xonomi™—l n—me or gr—de levelD sin™e m—t™hes well the p—leo˜iogeogr—phi™ rel—tionships of the qondw—n—

freshEw—ter t—x—F „he spe™i—lized deepE˜odied qondw—n— —nd „ethy—n frough @IWQTA first ™oined itD to lodge — possi˜le unn—E

perleidiformes —re independently evolvedD —s™ert—ined ˜y the p—rsiE

tur—l —ssem˜l—ge of fossil fishes whi™h possess mos—i™

mony —n—lysisF

™h—r—™ters of p—l—eonis™oids —nd higher —™tinopteryE

gi—ns eFgF holoste—ns @or ˜—s—l neopterygi—nsY frough ‚i—ssuntoF †engono des™ritti due esempl—ri ˜en ™onserv—ti del

perleidiforme puyu—nperleidus dengi qeng et —lFD PHIPD provenienti d—l IWQWY ƒ™h—effer IWSSD IWUQAF ƒevenorders @€—r—semionoE

wem˜ro ƒuperiore dell— qu—nling porm—tion @€elsoni™oD enisi™oD „ri—s tiformesD ‚edfieldiiformesD €holidopleuriformesD vug—E

wedioA nell— gonte— di vuoping @€rovin™i— dello ‰unn—nD gin— weriE

noiiformesD geph—loxeniformesD €erleidiformesD €eltoE dion—leAF v9—pp—rtenenz— —i €erleidiformes viene ™onferm—t— gr—zie —ll—

pleuriformesA h—ve previously ˜eenproposed @ƒ™h—effer

™om˜in—zione di ™—r—tteri d—t— d—ll— struttur— gener—le del ™r—nio @—mE

IWUQAF rowever further t—xonomi™ revisions proved pio preoper™olo ™he mostr— unpro™esso infr—or˜it—le˜ensvilupp—toD

m—s™ell—re ™on un9—lt— regione posteriore e dent—tur— f—tt— di lunghi e th—t only if the ––su˜holoste—ns99 is severely restri™tedD

I hep—rtment of qeology —nd qeologi™—l wuseumD €eking niversityD feijing IHHVUID €eople9s ‚epu˜li™ of ghin—F

iEm—ilX sunzuoyudpkuFeduF™nY dji—ngdpkuFeduF™nY wh—odpkuFeduF™nY ylsundpkuFeduF™nY lins—n™—idITQF™om

P hip—rtimento di ƒ™ienze dell— „err—D niversit—Ádegli ƒtudi di wil—noD vi— w—ngi—g—lli QRD sEPHIQQ wil—noD st—lyF iEm—ilX ™ristin—Flom˜—rdodunimiFitY —ndre—FtintoridunimiFit

QTH ƒun FD vom˜—rdo gFD „intori eFD ti—ng hFD r—o ‡FD ƒun ‰F 8 vin rF

™—n possi˜ly syn—pomorphies ˜e identified @furgin ghin— iFeF the enisi—n €—nxi—n p—un—D the v—te v—dini—n È

ˆingyi p—un—D —nd the i—rly g—rni—n qu—nling p—un— IWWPY q—rdiner et —lF IWWTAF xow—d—ysD the ––su˜holosE

@ti—ng et —lF PHHSY vi PHHTY wot—ni et —lF PHHVAD is th—t it te—ns99D restri™ted to peltopleuriforms —nd perleidiformsD

yieldsD ˜esides m—rine reptiles —nd inverte˜r—tesD — represent — morphologi™—lly distin™t group whi™h is

highly —˜und—nt fish —ssem˜l—geD pro˜—˜ly with more ™h—r—™terized ˜y the ™om˜in—tion of — primitive skullE

th—n PS t—x— of —™tinopterygi—ns —nd ™rossopterygi—nsF p—tternwith the m—xill— stri™tly —rti™ul—ted to the preE

„he good preserv—tion —nd the high num˜er of spe™iE oper™ulumD — pe™uli—r squ—m—tionm—de of deep —nterior

mensD m—ke the vuoping p—un— one of the most imporE fl—nk s™—lesD —n —dv—n™ed dors—l —nd —n—l fin stru™ture

t—nt finds in the l—st de™—des for widdle „ri—ssi™ m—rine with lepidotri™hi— equ—l in num˜er to their endoskelet—l

i™hthyof—un—s @„intori et —lF PHIHAF ƒo f—rD —lre—dy IP supports ex™ept for the l—st r—di—l supporting more th—n

spe™ies of W gener— of fishes h—ve ˜een n—med —nd deE one lepidotri™hium @—s for the ˜—s—l neopterygi—nsD see

s™ri˜ed @w—r™opoloi™hthys —ni „intori et —lF PHHVY vuoE „intori et —lF PHIHAD —nd the hemihetero™er™—l ™—ud—l fin

pingi™hthys ˜ergi ƒunet —lF PHHWY ƒ—uri™hthys d—w—zienE ne—rly re—™hing —n extern—l symmetry with the ep—xi—l

sis ‡u et —lF PHHWY ƒ—uri™hthys yunn—nensis h—ng et —lF finr—ys @„intori8 vom˜—rdo PHHSY vom˜—rdo et —lF

PHIHY qymnoi™hthys inopin—tus „intori et —lF PHIHY r—E PHIIAF

˜roi™hthys ˜roughi vinet —lF PHIIY €erleidus sinensis ––ƒu˜holoste—ns99D so f—r well re™orded from the

vom˜—rdo et —lF PHIIY ƒ—ngiorgioi™hthys sui voÂpezEerE widdle „ri—ssi™ of xorthernst—lyD ƒwitzerl—ndDƒp—in

˜—rello et —lF PHIIY ƒinos—uri™hthys longipe™tor—lis ‡u et —nd ghin—D —re gre—tly morphologi™—lly spe™i—lized

—lF PHIIY ƒinos—uri™hthys longimedi—lis ‡u et —lF PHIIY @furginIWWPD IWWTY vom˜—rdo IWWWD PHHIY vom˜—rdo È

ƒinos—uri™hthys minut— ‡u et —lF PHIIY uyphosi™hthys 8 „intori PHHRY vom˜—rdo et —lF PHHVY „intori 8 vomE

gr—ndei ˆu 8 ‡u PHIPAF ˜—rdo PHHSY ƒunet —lF PHHVD PHHWA ˜e™—use of their noE

„he qu—nling porm—tion of the vuoping gounty t—˜ly differenti—ted trophi™ —d—pt—tionsF sn the v—te

gener—lly represents the enisi—n deposits of the southE „ri—ssi™D ––su˜holoste—ns99 re—™hed —g—in — r—ther high

western ‰—ngtze €l—tform in the ghin— ƒouthern flo™kF level of morphologi™—l spe™i—liz—tionsD su™h —s fins —nd

„he fossiliferous l—yers yielding the vuoping f—un— ˜eE verte˜r—l ™olumn—d—pted to gliding@ „hor—™opterusD

long to the pper wem˜er —nd were su˜divided into „intori 8 ƒ—ssi IWWPAD powerful pred—tory dentition

two horizonsD n—mely vower —nd pper †erte˜r—te @q—˜—nelli—D „intori 8 vom˜—rdo IWWTA or fully duroE

rorizons @v†r —nd †rD see —lso ƒun et —lF PHHWA ph—gous one @indenni—D vom˜—rdo 8 fr—m˜ill—s™—

sep—r—ted ˜y IFV m thi™kD ˜—rrendolomiti™ limestones PHHSAF

with ™hert nodules @pigF IhAF „he ™—r˜on—te mi™rof—™ies „he two f—irly wellEpreserved spe™imens of

—n—lysis suggests th—t the pper wem˜er of the qu—nE puyu—nperleidus dengi from the widdle enisi—n @widE

ling porm—tion represents — sh—llowingEupw—rd @regresE dle „ri—ssi™A of the vuoping p—un—D ‰unn—n €rovin™eD

siveA deposition—l sequen™es in —n interpl—tform ˜—sin ƒouthern ghin—D suggest to ere™t — new perleidiform

@ru—ng et —lF PHHWAF f—mily for this genusD —nd to provide further insights

„he vuoping p—un— is supposed to ˜e ™oev—l to onthe rel—tionshipsof perleidiformsD espe™i—lly reg—rdE

the €—nxi—n p—un— @ƒun et —lF PHHTD PHHWY ru—ng et —lF ing the —rr—ngement of the fl—nk s™—lesF st must ˜e

PHHWAF e™tu—llyD ™onodont ˜iostr—tigr—phy of the pper pointed out th—t spe™imens des™ri˜ed herein —re from

wem˜er of the qu—nling porm—tionD exposed in ˜oth — quite different fossil —ssem˜l—ge in respe™t to the supE

vuoping gounty @‰unn—n €rovin™eA —nd —dj—™ent €—nE posed one for the holotype of qeng et —lF @PHIPAF

xi—n gounty @quizhou €rovin™eAD —s™ert—ined the fossiE

liferous l—yers of these two m—rine verte˜r—te f—un—s

within the s—me ™onodont xi™or—ell— ko™keli zoneD — qeologi™—l setting —nd —ge

definitive indi™—tor of the €elsoni—n in the „ethy—n

„he vuoping p—un—D from the pper wem˜er of re—lmF roweverD it is —lmost impossi˜le to ™omp—re in

the qu—nling porm—tion @€elsoni—nD widdle „ri—ssi™A of more det—iled w—y the re—l rel—tive —ge of the two sitesF

‰unn—n €rovin™e in ƒouth ghin— @™orresponding to the

wem˜er ss in h—ng et —lF PHHWAD is — re™ently dis™overed ‚em—rksF sn this p—perD following the €rin™iple of

fossilEv—gerst—tteD whi™h yields —˜und—nt —nd very well È €riorityD we use the nomen™l—ture —dopted ˜y qeng et

preserved fishesD m—rine reptilesD different —rthropods —lF @PHIPAD ˜ut some ™l—rifi™—tions —re neededF

@isopodsD de™—podsD horseshoe ™r—˜sD mysidi—™e—nsD snere™tingthe t—xon puyu—nperleidus dengi qeng

et™FAD mollus™s @˜iv—lvesD g—stropodsD ™eph—lopodsAD et —lFD PHIPD the euthors indi™—te —s vo™—lity —nd rorE

e™hinodermsD ˜r—™hiopods —nd ™onodontsD —s well —s izonX ––tiy—ngsh—nD ƒhi˜—li—nsh—nD puyu—n gountyD ‰unE

for—minifersD m—king it — ™omplex m—rine e™osystemF n—n €rovin™eY widdle „ri—ssi™ @ pper v—dini—nA huE

„he striking fe—ture of the vuoping p—un—D in ™ontr—st g—npo wem˜er of the p—l—ng porm—tion99 @qeng et —lF

to other „ri—ssi™ m—rine verte˜r—te f—un—s of ƒouthern PHIPX WITAF pollowing this indi™—tionD we ™—st dou˜t

xew perleidiform f—mily @ystei™hthyesD e™tinopterygiiA from widdle „ri—ssi™ of ƒouth ghin— QTI

IA the holotype of puyu—nperleidus dengi qeng et

—lFD PHIPD h—s the s—me kind of preserv—tion shown ˜y

the m—teri—l ™oming from the fossiliferous l—yers of the

vuoping p—un—D —s the spe™imens here des™ri˜edF „he

field experien™e of some of the euthors of this p—per

@FƒFD eF„FD hFtFD ‰FƒFA suggests th—t the fossils ™oming

from puyu—nh—ve — kindof preserv—tion™ompletely

differentX therefore the provenien™e of the holotype of

puyu—nperleidus dengi qeng et —lFD PHIPD from th—t loE

™—lity —nd horizon is highly questionedF

PA „he holotype of the spe™ies isD —™™ording to the

euthors who ere™ted itD v—te v—dini—n in —geD while the

spe™imens des™ri˜ed in this p—per ™ome from widdle

enisi—n levelsF „he time sp—n dividing these spe™imensD

whi™h undou˜tedly ˜elong to the s—me spe™iesD is —t

le—st suspi™iousD in p—rti™ul—r if we t—ke into ™onsider—E

tionthe explosive evolution—ryr—di—tionshown˜y the

widdle enisi—n i™hthyof—un—sD —nd the wellEknown exE

tr—ordin—ry v—ri—˜ility of the represent—tives of ––su˜hoE

loste—ns99 @vom˜—rdo et —lF PHHVD PHIIY ƒun et —lF PHHWY

vinet —lF PHIIY qenget —lF PHIPAF purthermoreD followE

ing the l—test version of the sntern—tion—l ghronostr—tiE

gr—phi™ gh—rt @sntern—tion—l gommission on ƒtr—tigr—E

phy ± s qƒD tuly PHIPAD the time sp—n˜etweenwiddle

enisi—n —nd pper v—dini—n m—y ˜e up to IH wyX this

m—kes unlikelyD —™™ording to usD the presen™e of the

s—me spe™ies inlevels sever—l millionsye—rs —p—rtF woreE

overD the v—te v—dini—n fish f—un— from hinxi—oE‡ush—E

puyu—nis quite poor in––su˜holoste—ns99 ™omp—red to

the enisi—n vuoping p—un—D ˜eing so f—r known only

r—˜roi™hthys orient—lis @vinetF —lF PHIIA —nd €eripeltoE

pleurus spF nF @tin PHHT —nd person—l o˜serv—tionsAD ˜oth

of them representing the youngest represent—tives for

these gener—F

w—teri—l —nd methods

sn the se™tion through the qu—nling porm—tionD hwz P —nd

hwz R ™omprise the fossiliferous l—yers of the vuoping p—un— @pigF

IgD hAY the spe™imens des™ri˜ed herein were ™olle™ted from the hwz

RD —nd —re now stored —t the qeologi™—l wuseum —nd hep—rtment of

qeologyD €eking niversityD feijingD ghin— @qw€u AF „hey were

prep—red me™h—ni™—lly ˜y fine steelEneedles under the ˜ino™ul—r mi™roE

s™opes ‡svh w VF hr—wings were m—de ˜y one of ™o—uthors @gFvFAFD

with the help of — ™—mer— lu™id— —nd illustr—tion softw—re for digit—liz—E

tionF

„he —n—tomi™—l nomen™l—ture of the skull —re used —™™ording to

furgin @IWWPAF gon™erning the terminology of the elements of the È

pigF I E eD fD simplified m—ps indi™—ting the site of the vuoping

™—ud—l finD we follow rut™hinson @IWUQ—AD q—rdiner @IWVVAD qr—nde

p—un—Y gD hD simplified str—tigr—phi™ ™h—rt indi™—ting the

8 femis @IWWVA —nd vom˜—rdo @PHHIAD referring the term ––ep—xi—l fin

str—tigr—phi™ position of the vuoping p—un— —nd det—iled

r—ys99 to the ™—ud—l finr—ys dors—l to the noto™hordor whenthe upper

sequen™e of the v†rD respe™tively @modified from ƒun et

™—ud—l fin r—ys elong—te ˜eyond the termin—tion of the ˜ody —xis @for

—lF PHHWAF

further dis™ussionsee vom˜—rdo PHHIAF

e˜˜revi—tions used in figuresX —flsD —n—l fin l—ter—l s™uteY —ntD

—˜out the provenien™eD —nd therefore the d—tingD of the

—ntor˜it—lY ˜fD ˜—s—l ful™r—Y ˜rD ˜r—n™hiosteg—l r—ysY dptD dermopteroti™Y

m—teri—l des™ri˜ed ˜y qeng et —lF @PHIPAD on the ˜—sis of drD dist—l r—di—lsY ™lD ™leithrumY dsphD dermosphenoti™Y efrD ep—xi—l fin

two m—ino˜serv—tionsX r—ysY exs™D extr—s™—pul—rY frD front—lY hyD hypur—lsY ifoD infr—or˜it—lsY ldrD

QTP ƒun FD vom˜—rdo gFD „intori eFD ti—ng hFD r—o ‡FD ƒun ‰F 8 vin rF

l—st dist—l r—di—lY lljD left lower j—wY llsD l—ter—l line s™—lesY mxD m—xill—Y thi™kD somewh—t higher th—n longer only in the —nterior

n—D n—s—lY opD oper™ulumY p—D p—riet—lY p—sD p—r—sphenoidY p™lD post™leiE

fl—nkF xeverthelessD morphologi™—l v—ri—˜ility of the

thrumY pfrD pelvi™ finr—ysY pmxD prem—xill—Y popD preoper™ulumY prD

fl—nk s™—les @rut™hinson IWUQ—Y furginIWWPY „intori È

proxim—l r—di—lsY ptD posttempor—lY rD r—di—lsY rljD right lower j—wY roD

8 vom˜—rdo IWWTY vom˜—rdo IWWWY wutter 8 rerzog rostr—lY s˜oD su˜or˜it—lY s™D s™—lesY s™lD supr—™leithrumY soD supr—or˜it—lY

PHHRA hinders the t—xonomi™ signifi™—n™e of squ—m—tion sopD su˜oper™ulumY vlrD ventr—l lo˜e r—ys of the ™—ud—l finF

for ––su˜holoste—n99 t—x—F ispe™i—lly in peltopleuriformsD

the depth of the midEl—ter—l s™—les is highly v—ri—˜leD

r—nging from — depth equ—ling th—t of the whole fl—nk ƒystem—ti™ p—leontology

@r—˜roi™hthysD frough IWQWY furginIWWHD IWWPY vinet È

—lF PHIIA to — ™omplete —˜sen™e @„hor—™opterus m—gniE

ruxleyD IVVH ystei™hthyes

fi™usD „intori 8 ƒ—ssi IWWPAD through v—rious intermediE

e™tinopterygii gopeD IVVU

—te st—ges @vom˜—rdo IWWWAF „he new t—xon h—s two

€erleidiformes fergD IWQU horizont—l rows of high fl—nk s™—les so th—t reg—rding

the sh—pe of the fl—nk s™—les it is seemingly more simil—r puyu—nperleidid—e f—mF nF

to peltopleuriforms th—nto perleidiformsD however they

hi—gnosisX ƒm—ll €erleidiformes with pe™uli—r squ—m—tionp—tE —re thi™k with g—noine r—ther th—n thin —s th—t of peltoE

ternX the num˜er of s™—les in e—™h verti™—l row in™re—ses ˜ehind the

pleuriformsF woreoverD fl—nk s™—les —re sometimes sigE

pelvi™ fins ˜y —ddition of sever—l s™—les ventr—l to the l—ter—l lines s™—le

nifi™—ntly higher —lso in the perleidiforms eltisolepis —nd rowF

vug—noi— @furginIWWPD PHHSY wutter 8 rerzog PHHRAF È

„he presen™e of ep—xi—l fin r—ys in the t—il is —

his™ussionF „he morphologi™—l ™h—r—™ters of the

™h—r—™ter sh—red ˜y perleidiforms —nd peltopleuriformsD

––su˜holoste—ns99 —mong perleidiforms —nd peltopleuriE

however it is highly v—ri—˜le in num˜erF emong perleiE

forms h—s —lre—dy ˜eenrem—rked @fu rginIWWPY „intori È

diformes the num˜er of ep—xi—l fin r—ys r—nges from

8 vom˜—rdo IWWTY vom˜—rdo 8 „intori PHHRY wutter

—˜out six @€erleidusAtoIH@gleithrolepisAorIP@q—˜—E

8 rerzog PHHRD —mong othersAF puyu—nperleidus fits

nelli—A @q—rdiner IWVVY q—rdiner 8 ƒ™h—effer IWVWY „inE well with ––ƒu˜holoste—n99 morphologiesX moreoverD —

tori 8 vom˜—rdo IWWTY vom˜—rdo PHHIAD —s inthe ™—se ™om˜in—tion of fe—tures th—t ™le—rly m—kes it dis™rimiE

of puyu—nperleidus @IHEIP ep—xi—l finr—ysAF n—te from the known represent—tives of the group ™—n

emong the ™h—r—™ters sh—red ˜y ––su˜holoste—ns99D ˜e proposedF

one of the most pe™uli—r is the presen™e of sexu—l diE „he dentition —nd the sh—pe of the preoper™ulum

morphism usu—lly shown ˜y modified —n—l fins in m—le —re differenti—ted ˜etween perleidiforms —nd peltoE

spe™imens @furginIWWH—Y vom˜—rdo IWWWAF es it is for È pleuriformsF „he dentition of perleidiforms is gener—lly

the p—ttern of squ—m—tionD there is — v—riety of —n—l fin m—de of pegElike m—rgin—l teeth —nd ™rushing inner ones

from — simple thinning of r—ys with sm—ll modifi™—tionsD @evenif nothighly spe™i—lizedA @fu rginIWWPY vom˜—rdo È

hooklets onthe posterior p—rt of the fin@eFgF in r—˜E PHHIA ˜ut deepE˜odied forms show — more spe™i—lized

roi™hthysDfurginIWWH or „hor—™opterusD gFvF persF È dentition su™h —s in pel˜eri— @vom˜—rdo 8 „intori

o˜sFA to ™omplex stru™turesD simil—r to the gonopodi— PHHRA —nd ƒtopp—ni— @vom˜—rdo et —lF PHHVA from the

of some living gyprinodontid— teleostsD —s in €eripeltoE western„ethysF €eltopleuriforms —re often™h—r—™terE

pleurus —nd €eltopleurus @furginIWWH˜D IWWPY vom˜—rE È ized ˜y minute teeth or no tooth —t —ll @eFgF r—E

do IWWWAF xeverthelessD sexu—l dimorphism is —lso preE ˜roi™hthysDfurginIWWPY €eltopleurusD vom˜—rdo IWWWAF È

sent in some perleididD q—˜—nelli— —gilis possessing — €erleidiforms oftenh—s — l—rge —ndwedgeEsh—ped preE

strongly dimorphi™ —n—l fin @„intori 8 vom˜—rdo oper™ulumD while the preoper™ulum is mu™h n—rrower

IWWTAF fesides the modified finr—ysD the m—le —n—lfin —nd more verti™—lly —rr—nged —mong peltopleuriformsD

of sexu—lly dimorphi™ ––su˜holoste—ns99 is often ™h—r—™E ex™ept few gener—D eFgF weridensi— or eltisolepisD whose

terized ˜y the presen™e of — l—rge s™ute whi™h ™overs the preoper™ulum is quite n—rrow —nd verti™—l @furginIWWPY È

proxim—l ends of the lepidotri™hi—F e l—rge s™ute present wutter 8 rerzog PHHRAF „he new t—xon shows — l—rge

onspe™imenqw€u E€EQHHW of puyu—nperleidus is preoper™ulum with — well developed infr—or˜it—l proE

thus interpreted —s rel—ted to the sexu—l dimorphism ™essD —nd strong upper —nd lower j—ws ˜e—ring long

r—ther th—n— simple preE—n—ls™—le due to its sh—pe pegElike teeth with —n —™rodine ™—pD whi™h is reminisE

—nd typi™—l positionF „he other spe™imen qw€u E€E ™ent of perleidiformsF

QHIH l—™ks this l—rge s™uteD ˜ut inste—d its ventr—l s™—les yf the post™r—ni—l —n—tomyD differen™es ˜etween

h—s denti™ul—tions on the posterior m—rgin while those perleidiforms —nd peltopleuriforms in™lude the sh—pe

of qw€u E€EQHHW h—ve — smooth oneF „he mutu—lly —nd —rr—ngement of the fl—nk s™—lesD —nd the stru™ture

ex™lusive presen™e of — l—rge —n—l s™ute —nd denti™les on of the ™—ud—l finF „he fl—nk s™—les —re typi™—lly thinD

n—rrow —nd high to very high —long the ˜ody in peltoE posterior m—rginof the s™—les ™—n˜e therefore rel—ted to

pleuriformsD while those of perleidiforms —re usu—lly sexu—l dimorphism r—ther th—nto t—xonomyF

xew perleidiform f—mily @ystei™hthyesD e™tinopterygiiA from widdle „ri—ssi™ of ƒouth ghin— QTQ

„—king —ll ™h—r—™ters mentioned —˜ove into ™onE from the perleidiform f—milies mentioned —˜oveX for this

sider—tionsD the t—xon here des™ri˜ed is in™luded within re—son it is ne™ess—ry to ere™t — new f—mily ˜—sed on this

unique —ut—pomorphy to —™™ommod—te itF perleidiforms r—ther th—npeltopleuriformsD inp—rti™ul—r

˜—sed onthe fe—tures of the skull p—tternF

‚em—rksF qeng et —lF @PHIPAD ere™ted the new es — very su™™essful orderD the €erleidiformes ™urE

genus puyu—nperleidusD on the ˜—sis of — singleD —ndD rently in™lude seven f—miliesX q—˜—nellid—eD gleithroleE

—™™ording to the euthorsD only known spe™imen —nd pid—eD €olz˜ergiid—eD golo˜odontid—eD €erleidid—eD

they put it withinthe f—mily €erleidid—e D together with €seudo˜e—™oniid—e —nd vug—noiid—e @frough IWQIY

other new t—x— of perleidiformsD vuopingperleidus —nd rut™hinson IWUQ—Y q—rdiner IWVVY furginIWWPY „intori È

hi—ndongperleidus ˜—sed onfollowing™h—r—™tersX 8 vom˜—rdo IWWTY wutter PHHRY vom˜—rdo 8 „intori

––fishes of sm—ll sizeD the preoper™ulum ne—rly verti™—lD PHHRY voÁpezEer˜—rello 8 —v—ttieri PHHVAF

most of skull ˜ones orn—mented with tu˜er™les or „he q—˜—nellid—eD — monotypi™ f—mily repreE

ridgesD m—xill— —s in€—l—eonis™iformesDj—ws toothed sented ˜y — single spe™iesD q—˜—nelli— —gilisD is typi™—l

with ™oni™—l teethD ne—rly symmetri™—l ™—ud—l fin99 for its numerous rows @—˜out WSA of very thin s™—les —nd

@qeng et —lF PHIPX WPSAF e™™ording to usD with this the powerful pred—tory dentition m—de of ™oni™—lD r—E

™hoi™e qeng et —lF @PHIPA underestim—ted the rem—rk—˜le di—lly stri—tedD uneven teeth @„intori 8 vom˜—rdo

differen™es on ˜ody morphologyD skull p—ttern —nd IWWTAF ell represent—tives of the deepE˜odied perleidiE

—˜ove —ll squ—m—tion˜etweenpuyu—nperleidus —nd form f—mily gleithrolepidiid—e —re ™h—r—™terized for

the other gener— des™ri˜ed in the s—me p—per @—™tu—lly their tri—ngul—r m—xill—D — n—rrow lower j—w devoid of

the three generi™ di—gnosis —re very simil—r to e—™h other tooth —nd the —˜sen™e of posttempor—l @rut™hinson

—nd quite uninform—tiveAD whi™hD owing to their —™tu—l IWUQ—Y q—rdiner IWVVAF „he €seudo˜e—™oniid—e —re ™h—rE

™h—r—™tersD ™—nnot ˜e —s™ri˜ed to the s—me f—milyF „—kE —™terized ˜y the presen™e of —n in™omplete dors—l ridge

ing into ™onsider—tion the ™omplexity of the orderD —lso of spineElike s™—les ˜etween the skull —nd the dors—l fin

the ™omp—rison of the genus with the represent—tives of —nd s™—les with str—ight posterior ˜orderD elev—ted ™enE

e—™h f—mily of €erleidiformes —ppe—rs simplisti™ —nd tr—l region —nd m—rgin—l ™on™entri™ ridges of g—noineF

hurriedF puyu—nperleidus is unique for the insertion of „he f—mily €olz˜ergiid—e in™ludes deepEfusiform to

s™—les ine—™h verti™—l row —fter the pelvi™ finsXsimil—r deepE˜odied represent—tivesD whi™h sh—re the gener—l

—ddition —mong perleidiforms is known @—lthough to — skull p—tternwith —noper™ulum sm—ller th—nthe su˜E

lesser degreeA only on vug—noi— @see —˜oveAD ˜ut inthe oper™ulumD the prem—xill— fused to the —nteroEdors—l

dis™ussion qeng et —lF @PHIPA do not mention itD limiting edge of the m—xill—D —nd the presen™e of the pro™um˜ent

to ™ite — different s™—le p—ttern for the two t—x—F e™™ordE long —nterior teeth @qriffith IWUUY vom˜—rdo 8 „intori

ing to the qeng et —lF @PHIPAD ™h—r—™teristi™ of puyu—nE PHHRY vom˜—rdo et —lF PHHVY ƒunet —lF PHHWAF golo˜oE

given ˜y the deepened fl—nk s™—lesY for this perleidus is dontid—e —re represented ˜y l—rgeEsized perleidiforms

re—sonthey ™omp—re their newt—xonwith €eltoperleiE whi™h yield — r—ther ™onserv—tive skull osteologyD su™h

dus —nd eltisolepisD from whi™h puyu—nperleidus differsD

—s the presen™e of — sm—ll rostr—l followed ˜y — l—rge

—mong the other ™h—r—™tersD —lso for ––lower deepened postrostr—l sep—r—ting n—s—l ˜onesD —nd —mong other

fl—nk s™—le row ™ontinued ˜y five horizont—l s™—le rows

™h—r—™tersD — hemiduroph—gous dentition —nd well orn—E

posterior to level of —n—l fin insertion99 @qeng et —lF PHIPX

mented s™—les @furginIWWTY wutter PHHRY ‚us™oniet —lF È

WPSAD negle™ting the import—n™e of the pe™uli—r —ddition

PHHUY ƒunet —lF PHHVAF „he genus vug—noi—D the only

of s™—lesF represent—tive of the f—mily vug—noiid—eD shows the —dE

dition of one or two s™—le rows in ™orresponden™e of the

m—in midEl—ter—l fl—nk s™—lesD the ones ˜e—ring the l—tE

puyu—nperleidus qengD tinD ‡u 8 ‡—ngD PHIP

er—l line ™—n—lF vug—noi— is however very derived for its

„ypeEspe™iesX puyu—nperleidus dengi qengD tinD ‡u 8 ‡—ngD PHIP

skull p—tternD for inst—n™e the m—xill— is ™ompletely free

from the l—rge —nd ˜ro—d preoper™ulum —nd the eleE hi—gnosisX es for the single spe™iesF

qeogr—phi™ distri˜utionX es for the spe™ies ments of the ™heek show very distin™tive morphology

—nd —rr—ngement @frough IWQWY furginIWWPAF elthough È

with — skull p—tternvery simil—r to th—t of the new puyu—nperleidus dengi qengD tinD ‡u 8 ‡—ngD PHIP

t—xonD —mong known €erleidid—e gener—D none of them pigs PET

is ™h—r—™terized ˜y — squ—m—tionwith very deep midE

hi—gnosis @emendedAX emong the following ™om˜in—tion of l—ter—l s™—les —s well —s the insertion of —ddition—l s™—le

™h—r—™tersD —n—sterisk @BA indi™—tespossi˜le —pomorphiesX deepEfusiE

rows posterior to the pelvi™ finsF

form perleidiform up to VS mm in st—nd—rd length @ƒvAY oper™ulum

„he unique p—ttern of squ—m—tion with the inserE

slightly l—rger ˜ut n—rrower th—n the su˜oper™ulumY s™—les of the

tionof sever—l s™—les ine—™h verti™—l row —fter the pelvi™ midEl—ter—l fl—nk rowD ™—rrying the l—ter—l lineD highly deepened up to

the level of the pelvi™ finsY B—ddition of up to four s™—les in the verti™—l finsD definitively dis™rimin—te the genus puyu—nperleidus

QTR ƒun FD vom˜—rdo gFD „intori eFD ti—ng hFD r—o ‡FD ƒun ‰F 8 vin rF

pigF P E puyu—nperleidus dengi qengD tinD ‡u 8 ‡—ngD PHIPX eD qw€u E€EQHHWY fD qw€u E€EQHIHY gD dr—wing of qw€u E€EQHHWF

ƒ™—les visi˜le in inner side —re sh—dedF

rows ˜ehind the pelvi™ finsY ventr—l s™—les ˜etween the pe™tor—l —nd rolotypeX e ne—rly ™omplete spe™imenD s†€€ †ITSIUD stored

pelvi™ fins very lowD mu™h longer th—n deepY Bs™—les ™overing the ™—ud—l —t the snstitute of †erte˜r—te €—leontology —nd €—leo—nthropologyD

pedun™le irregul—rly sh—pedF „ri—ngul—r dors—l fin well posterior to the ghinese e™—demy of ƒ™ien™esD €ekingF

h—lfElength of the ˜odyY Bdist—l r—di—ls of the dors—l fin enl—rgedD with w—teri—lX qw€u E€EQHHW @pigF PeD gAX —n—lmost ™omplete

the l—st one ˜e—ring four lepidotri™hi—Y ™—ud—l fin wideD with more th—n spe™imen of TQ mm in st—nd—rd length @ƒvAF „he snout region is missing

RH ™losely —rr—nged lepidotri™hi— of whi™h IH to IP —re referred to —s —nd the skull roof is fr—gment—ryF qw€u E€EQHIH @pigF PfAX — f—irly

ep—xi—l finr—ysF wellEpreserved fish of VS mm in ƒvD with elements of the snout region

xew perleidiform f—mily @ystei™hthyesD e™tinopterygiiA from widdle „ri—ssi™ of ƒouth ghin— QTS

pigF Q E puyu—nperleidus dengi qengD

tinD ‡u 8 ‡—ngD PHIPX dr—wE

ings of the skull in eD

qw€u E€EQHHWY fD qw€E

u E€EQHIHF

—nd the skull roof p—rti—lly preservedF „he s™—les —long the ˜elly —nd the @pigF QeD fAF „he n—s—ls —re re™t—ngul—r in sh—peD dorE

dors—l ridge —re s™—ttered —nd two or three rows of deepened s™—les on

soEventr—lly elong—tedY the opening for the —nterior nosE

the —nterior fl—nk —re missingF

tril is well dete™t—˜le onspe™imenqw€u E€EQHIHF

vo™—lity —nd horizonX es mentioned —˜oveD the distri˜ution of

„he pores of supr—or˜it—l sensory ™—n—l —re longitudinE the spe™ies is limited to —round h—w—zi vill—ge of vuoping gountyD

—lly —rr—nged with respe™t to the longest —xis of the ‰unn—n €rovin™eD ƒouth ghin—X vower †erte˜r—te rorizon @v†rA of

the vuoping p—un—D in the pper wem˜er of the qu—nling porm—tionD ˜oneF e l—rge ™onvex element visi˜le from the inside

€elsoni—n @widdle enisi—nD widdle „ri—ssi™AF „he presen™e of the speE

—nd p—rtly ™overed ˜y other ˜ones is interpreted —s

™ies —t tiy—ngsh—nD ƒhi˜—li—nsh—nD puyu—n gountyD ‰unn—n €rovin™e

the rostr—lF „he front—l is p—rti—lly preserved on ˜oth from the pper v—dini—n hug—npo wem˜er of the ––p—l—ng porm—E

spe™imensX it is quite shortD with — supr—or˜it—l n—rrow tion99 —s supposed ˜y qeng et —lF @PHIPA is not ™onsideredD even if the

region longer th—n th—t of the exp—nded postor˜it—l oneF genus @—nd ™onsequently the new f—milyA took the n—me from th—t

lo™—lityF „he pores of supr—or˜it—l sensory ™—n—l —long the l—ter—l

m—rgin—re visi˜le onqw€u E€EQHIHY the postor˜it—l

hes™riptionF „he two f—irly wellEpreserved spe™iE region is ˜ro—d ˜ut its outline is not ™le—rly dete™t—˜leD

mens —re deepEfusiform in sh—peD with tot—l lengthGm—xE —s well —s the ™ourse of the interfront—l sutureF „he surE

im—l ˜ody depth ˜eing —˜out QEQFSY spe™imen qw€u E f—™e of the front—l is orn—mented ˜y sm—ll tu˜er™les —nd

€EQHHW seems to ˜e — little deeper th—nqw€u E€E p—t™hes of g—noineF „he p—riet—l is qu—dr—ngul—rD joined

QHIHF „hey —re sm—llEsizedD with the ƒv up to VS mmF to the front—l through — sinuous suture —nd to the exE

„he he—d length is —˜out PW7 of the ƒvF „he pedun™le is tr—s™—pul—r ˜one in — rounded oneD well visi˜le on

moder—tely deepD ˜eing RH7 of the m—ximum ˜ody qw€u E€EQHHWF „wo pit lines —re dete™t—˜le on the

depthF „he tri—ngul—r dors—l fin is pl—™ed well posterior posterior h—lf of the ˜oneF yn the p—riet—l the orn—menE

to the h—lfElength of the ˜odyD four or five s™—leErows t—tion is m—de of g—noine p—t™hes whi™h tend to mergeD

˜ehind th—t of the —n—l fin @pigF PAF ell skull ˜ones —re m—king — quite homogeneous surf—™eF „he dermopteroE

orn—mented with sm—ll g—noine tu˜er™lesD somewh—t ti™ h—s — tr—pezoid—l sh—peD —nteroEposteriorlyY the

homogeneously —rr—nged on e—™h elementF strong orn—ment—tion is m—de of fl—t g—noine tu˜er™les

ƒkullF „he ˜ones in the ethmoid—l region —re ™omE —nd sm—ll pores of the tempor—l sensory ™—n—l —re visi˜le

pletely missing on qw€u E€EQHHWD while they —re p—rE on the ventr—l region of this ˜one on qw€u E€EQHIHF

ti—lly —nd s™—tteredly preserved on qw€u E€EQHIH „he extr—s™—pul—r ˜ones were displ—™ed —nd p—rti—lly

QTT ƒun FD vom˜—rdo gFD „intori eFD ti—ng hFD r—o ‡FD ƒun ‰F 8 vin rF

overl—pped ˜y the oper™ulum onqw€u E€EQHHWF „he wellEdeveloped infr—or˜it—l pro™ess extending up to

tr—je™tory of the supr—tempor—l ™ommissure ™—n˜e —˜out the middle of the or˜itF sts —nteroEventr—l m—rgin

identified ˜y — groove ™rossing the ventr—l p—rt of the is ™on™—veD where it widely em˜r—™es the posterior edge

extr—s™—pul—r ˜one on qw€u E€EQHIHF of the m—xill—F „he oper™ulum is — shieldElike elementD

gir™umor˜it—l seriesF et le—st two l—rge supr—orE its depth ˜eing IFS times of the m—ximum widthF st is

˜it—l ˜ones —re ™le—rly visi˜le on the dors—l m—rgin of slightly l—rger ˜ut n—rrower th—n the su˜oper™ulum —nd

the or˜it of ˜oth spe™imensD ˜ut their sh—pe —nd size —re ™ont—™ts the l—tter with — str—ight sutureF „he su˜operE

well dete™t—˜le only on qw€u E€EQHIHF „he —nterioE ™ulum is squ—rishD with — slightly rounded ventr—l m—rE

most one is tri—ngul—r —nd very l—rgeX it pro˜—˜ly forms ginF foth preoper™ulum —nd oper™ulum —re ™h—r—™terE

the —nterior m—rgin of the or˜itD ™ont—™ting the n—s—l ized ˜y —n orn—ment—tion m—de of g—noine tu˜er™lesD

˜onesF „he se™ond is re™t—ngul—r —nd o™™upies the —nteE more densely —rr—nged on preoper™ulumF fetween the

roEl—ter—l m—rgin of the front—l ˜oneD for h—lf of its posteroEdors—l ™orner of preoper™ulum —nd —nteroEdorE

lengthF foth —re elong—ted —nteroEposteriorly —nd —re s—l m—rginof oper™ulum there is enoughroom to —™E

strongly orn—mented ˜y sm—ll tu˜er™lesF „wo —ddition—l ™ommod—te some ˜ones @—nteoper™ulumD dermohy—lAD

sm—llD re™t—ngul—r fr—gments —re visi˜le on qw€u E€E howeverD owing to the st—te of preserv—tionD it is not

QHIH ˜ehind the se™ond element —ndD —™™ording to their possi˜le to ˜e ™onfidently —s™ert—inedF et le—st seven

positionD they ™ould represent — third —nd fourth eleE ˜r—n™hiosteg—l r—ys —re presentF „hey —re tri—ngul—r in

ment of the seriesF e sm—ll hourgl—ss elementD visi˜le sh—pe —nd ˜e—r fl—t g—noine tu˜er™lesF

on qw€u E€EQHHW —nd lying ˜ehind the infr—or˜it—l t—wsF „he upper j—w ™onsists of — m—xill— with —n

˜ones —nd on su˜or˜it—l is interpreted —s the dermoE exp—nded —nd rounded posterior regionD —nd — n—rrower

sphenoti™D whi™h is ™le—rly displ—™ed in respe™t to its —nterior one whi™h is nevertheless stoutF yn qw€u E

origin—l positionF „he orn—ment—tion is m—de of sm—ll €EQHIH — medi—n tr—pezoid—l ˜oneD ™—rrying two l—rge

tu˜er™les —rr—nged longitudin—lly to the elementF „here teeth th—t —re simil—r in size to those ˜orne ˜y the —nteE

is —t le—st one su˜or˜it—lD —lso visi˜le on qw€u E€E rior regionof the upper j—wD is interpreted—s the preE

QHHWD pl—™ed ˜etween—nteroEdors—lm—rginof preoperE m—xill—F „he dors—l ˜order of the upper j—w is slightly

™ulum —nd posteroEdors—l ˜order of the l—rgest infr—orE ™on™—veF „here —re nine long —nd strong pegElike teeth

˜it—l —nd p—rtly ™overed ˜y dermosphenoti™F „he st—te with —n —™umin—te —™rodine —pexD ˜orne on the —nterior

of preserv—tion —nd the strong orn—ment—tion do not h—lf of the or—l m—rginF „hey —re gr—du—lly de™re—sing in

—llow to st—te the pre™ise outline of this element or if length posteriorlyF „he lower j—w is — stout wedgeElike

other simil—r ˜ones were presentF yf the infr—or˜it—l ˜oneD with — ™onvex ventr—l m—rginF yn qw€u E€E

series only two elementsD whi™h ˜order the posteroE QHHW the or—l m—rginof the lower j—w is onlyp—rti—lly

ventr—l edge of or˜itD —re well dete™t—˜leF en elong—te visi˜leD ˜eing ™overed ˜y the ventr—l m—rgin of m—xill—D

elementD overl—pping the dors—l m—rgin of the m—xill—D is m—king impossi˜le to pre™isely des™ri˜e the det—ils of

interpreted —s the first infr—or˜it—lY even if its ventr—l teethF ynqw€u E€EQHIH there —re —t le—st six pegElike

suture is not ™le—rly definedD its sh—pe ™—n ˜e outlined teeth with —n—™rodine™—p visi˜le onthe or—l m—rginof

due to to the orn—ment—tion p—tternD with longitudin—l the lower j—wF

g—noine ridges following the dors—l m—rgin of the eleE €e™tor—l girdleF „he posttempor—l is roundedD

ment —nd ™le—rly different from th—t of the upper j—wD with — smooth dors—l region —nd — slightly orn—mented

whi™h ™onsists of rounded g—noine tu˜er™lesF yn ventr—l oneD where the pores of the sensory ™—n—l ™omE

qw€u E€EQHIH the s—me element seems to h—ve — very ing from the extr—s™—pul—r ˜ones —re dete™t—˜le on

l—rge ventr—l regionD ™overing most of the —nterior p—rt qw€u E€EQHHWF „he supr—™leithrum is n—rrow —nd

of the upper j—wD even if —lso in this spe™imenD its ventr—l deepD extending for PGQ of the height of oper™ulumY its

m—rginis notdete™t—˜leF xevertheless the ™ourse of the surf—™e is smooth —nd the p—ss—ge of the l—ter—l line is

pores of the infr—or˜it—l sensory ™—n—l whi™h tends to well dete™t—˜le onits dors—l regionF„he post™leithrum

˜end ™le—rly downw—rdsD is well definedF „he other inE h—s the s—me sh—pe —nd size of the supr—™leithrum —nd it

fr—or˜it—l visi˜leD ˜ordering the whole posterior m—rgin is pl—™ed —t the level of the suture ˜etwenoper™ulum

of the or˜itD is si™kleEsh—pedD with the dors—l lim˜ —nd su˜oper™ulumF „he ™leithrum is only visi˜le ˜y p—rt

slightly enl—rgedF „he infr—or˜it—l sensory ™—n—l is visiE of its ventr—l regionD ˜eing ™overed ˜y the su˜oper™uE

˜le —long the medi—n region of the elementF e n—rrow lumY its surf—™e is orn—mented ˜y sm—ll g—noine tu˜erE

elong—ted ˜one lying on the supposed rostr—l —nd ™—rryE ™lesF

ing the pores of — sensory ™—n—lD pro˜—˜ly represents the €—ired finsF „he p—ired fins —re in™ompletely preE

—ntor˜it—lF servedD m—king it impossi˜le to des™ri˜e the det—iled

gheekD oper™ul—r —nd ˜r—n™hiosteg—l seriesF „he stru™ture of r—ysF ‚emn—nts of the pe™tor—l fins in™lude

l—rge —nd verti™—l preoper™ulum h—s its dors—l region the proxim—l p—rts of the lepidotri™hi— of the right fin

ne—rly equ—l to the ventr—l one in sizeF st ˜e—rs — very —nd the very dist—l region of those of the left oneF „en

xew perleidiform f—mily @ystei™hthyesD e™tinopterygiiA from widdle „ri—ssi™ of ƒouth ghin— QTU

pigF R E puyu—nperleidus dengi qengD

tinD ‡u 8 ‡—ngD PHIPX dr—wE

ings of the medi—n fins in e

@dors—l finA —nd g @—n—l finAD

qw€u E€EQHHWY f @dors—l

finA —nd h @—n—l finAD

qw€u E€EQHIHF „he enE

doskelet—l elements —re

sh—ded @light greyX r—di—ls

—nd dist—l r—di—lsY d—rk greyX

proxim—l r—di—lsAF

r—ys ™—n ˜e ™ounted in the right pe™tor—l finF „he pelvi™ in front of the dors—l —nd ventr—l lo˜es —re segmented

th

s™—le rowD —re finsD pl—™ed —t the level of the IUGIV from their ˜—sesD while ˜etweenthe two lo˜es r—ys

preserved only in p—rt —nd with the dist—l region s™—tE show — slighly longer —nd slender proxim—l ˜—sesF „he

teredF yn ˜oth pe™tor—l —nd pelvi™ finsD the proxim—l pre™ise num˜er of ˜r—n™hing is un™le—r due to the inE

˜—ses —re very long —nd the r—ys —re pre™eded ˜y few ™omplete preserv—tionD howeverD the prin™ip—l r—ys

˜—s—l ful™r—F ˜r—n™h —t le—st twi™eF q—noine ™overing is present on

np—ired finsF „he dors—l fin@pigF ReD fA is l—rgeD ˜oth ep—xi—l —nd hyp—xi—l m—rgin—l r—ys —nd de™re—ses

th

tri—ngul—r in sh—peF st is pl—™ed —t the level of the PV gr—du—lly tow—rds the ™entr—lD where there —re —˜out ten

s™—le rowsF „here —re —˜out PH lepidotri™hi— with long ™entr—l lepidotri™hi— devoid of —ny g—noineF „he tip of

unsegmented proxim—l ˜—ses —nd the following sm—ll the finis deli™—te —ndp—rti—lly preserved ˜e™—use the

squ—red segmentsD pre™eded ˜y three ˜—s—l ful™r—F „he dist—l ends of —ll r—ys —re m—de of thinner —nd more

first lepidotri™hiumD segmented ˜ut un˜r—n™hedD is ruE sp—™ed elements whi™h l—™k of g—noineF yn qw€u E

diment—rily developedD ™onsisting of —˜out ten segments €EQHHW only one ˜—s—l ful™rum in front of e—™h lo˜e is

onqw€u E€EQHHW —ndthree onqw€u E€EQHIHD present —nd fringing ful™r— —re —˜sentD while on

—nd it is followed ˜y r—ys whi™h —pp—rently in™re—se qw€u E€EQHIH ˜oth lo˜es —re pre™eded ˜y strong ˜—E

—˜ruptly in lengthF i—™h r—y ˜r—n™hes —t le—st twi™eD s—l ful™r—F feing in™omplete in its posterior p—rtD it is not

˜eginning from the third oneF q—noine ™overing is reE possi˜le to verify if fringing ful™r— were presentF

stri™ted to the proxim—l elements of the —nteriormost indoskelet—l elementsF „he ˜—ses of the medi—n

r—ysD —nd to the medi—n region of the dist—l onesY on —nd ™—ud—l fins is p—rtly un™overed ˜y s™—les so th—t

these l—tterD g—noine —ppe—rs —s minute p—t™hes or tu˜erE the endoskelet—l elements ™—n ˜e o˜servedF „he medi—n

™lesF xo fringing ful™r— —re dete™t—˜leF fins show dist—l —nd proxim—l r—di—ls @pigF ReAX in the

„he tri—ngul—r —n—l fin @pigF ReD fA is pl—™ed —t the dors—l finD the dist—l r—di—ls —re l—rge —nd re™t—ngul—rD

th

s™—le rowsY there —re —˜out IP lepiE level of the PQGPR with thi™kened m—rginsD of whi™h e—™h r—di—l joins to

dotri™hi— ˜r—n™hing —t le—st twi™eF „he g—noine ™overing one lepidotri™hiumD ex™ept th—t the l—st one supports —t

is not homogeneousD ˜eing limited to the medi—n p—rt of le—st four r—ysF „he dors—l end of this l—st element is

the segments of the —nteriormost lepidotri™hi—D like th—t ˜ending posteriorlyF †entr—llyD these elements ™ont—™t

of the dors—l finF xeither ˜—s—l nor fringing ful™r— on the the proxim—l r—di—lsD visi˜le only in their dors—lmost

—nterior m—rgin of the fin —re visi˜leF regionF sn the —n—l finD the dist—l r—di—ls seem to ˜e

„he ™—ud—l fin@pigF SeD fA is hemihetero™er™—lD sm—ller th—nthose of the dors—l oneD˜ut they show their

with dors—l —nd ventr—l ˜ody lo˜es ne—rly symmetri™—l s—me stru™tureF snthe ™—ud—l finonly— sm—ll region

extern—llyF st is wideD slightly forked —nd h—s —˜out RQE ˜etween the two lo˜es is l—™king s™—lesD showing the

RT very ™losely —rr—nged lepidotri™hi—D of whi™h IH to IP dist—l ends of few hypur—lsF

™—n˜e ™onsidered—s ep—xi—l finr—ysF „here —re —t le—st T ƒqu—m—tionF p to RP tr—nsvers—l s™—le rows h—ve

unsegmented —nd un˜r—n™hed lepidotri™hi— on the dorE ˜een ™ountedD ˜ut the s™—les ™overing the ™—ud—l pedunE

s—l —nd five on ventr—l lo˜e on qw€u E€EQHHWF ‚—ys ™le seems irregul—rly —rr—ngedD so th—t it is not possi˜le

QTV ƒun FD vom˜—rdo gFD „intori eFD ti—ng hFD r—o ‡FD ƒun ‰F 8 vin rF

iorlyD e—™h s™—le qui™kly de™re—sing in height @pigF TeD

fAF wore ventr—llyD there —re —t le—st five horizont—l

rowsD whi™h —re mu™h longer th—n deep ˜etween the

pe™tor—l —nd pelvi™ fins @pigF PgAF efter theseD —ll the

s™—les ˜e™ome sm—ll —nd rhom˜oid—l in sh—peF „he

s™—les on the dors—l regionD —lthough less evidentD seem

to ˜e present the s—me horizont—l multipli™—tion of the

s™—les seen for the ventr—l fl—nk s™—leErowY —lthough the

—nteriormost p—rt of the dors—l region is not ™ompletely

preservedD onqw€u E€EQHHW there is —nevidentdisE

pl—™ement —nd insertion of —ddition—l s™—les rows ˜eginE

ning from the middle of the ˜ody to the insertion of the

dors—l finF feginning from this regionD the squ—m—tion

—ppe—rs homogeneous for sh—peD size —nd —rr—ngement

of the s™—lesD ˜eing rhom˜oid—lD slightly longer th—n

deeperD ex™ept for those ™overing the ™—ud—l pedun™leF

‡ithinthe ™—ud—l pedun™leDthe s™—les of the medi—n

p—rt —re sm—ll —nd roundedD while those ˜ordering the

dors—l —nd ventr—l s™—ly lo˜es of the fin —re —nteroEposE

teriorly elong—tedF yn qw€u E€EQHHW these s™—les —re

regul—rly —rr—nged on — single rowD while on qw€u E

€EQHIH they seem to ˜e more irregul—rly set due to st—te

of preserv—tionF „wo s™utes ™over the tip of the —xi—l

˜ody lo˜eX the dors—lmost is tri—ngul—rD while the venE

tr—l one h—s —n —xe sh—peF „he ™ourse of the l—ter—l line is

visi˜le tr—nsp—rently on the dors—l region of the high

fl—nk s™—lesD tr—™ed ˜y verti™—l slits on sever—l s™—les of

the posterior p—rt of the ˜odyF

en —n—l s™uteD ™overing the proxim—l p—rt of the

lepidotri™hi— of the —n—lfinonqw€u E€EQHHW @pigF

ReAD is visi˜leX it is r—ther l—rge —nd ov—l in sh—peD with

few g—noine on the ™entr—l p—rt of the elementF yn

qw€u E€EQHIH there is no —n—l s™ute where they —re

—ltern—ted ˜y the equiv—lents of few horizont—l rows of

pigF S E puyu—nperleidus dengi qengD tinD ‡u 8 ‡—ngD PHIPX very low s™—lesF „his differen™e is interpreted —s rel—ted

dr—wings of the ™—ud—l fins in eD qw€u E€EQHHWY fD

to the sexu—l dimorphismD hereinF ell s™—les —re thi™k

qw€u E€EQHIHF por — ˜etter lo™—liz—tionD the hypur—l

—nd smoothX on qw€u E€EQHHW their posterior m—rE

finr—ys @light greyA —ndthe ep—xi—l finr—ys @d—rk greyA

gins —re str—ightD while on qw€u E€EQHIH they —re h—ve ˜eensh—dedF

v—ri—˜le —nd ˜e—r sever—l sm—ll denti™ul—tionsD espe™i—lly

on s™—les of the ventr—l fl—nk s™—leErowD where denti™les to pre™isely dete™t e—™h verti™—l row inthe posteriorE

—re ˜orne ˜y most of the m—rgin even if they —re mu™h most p—rt of the ˜odyF „he —rr—ngement of the s™—les

denser in the ventr—l h—lfF ƒp—rse denti™les —re present of this new t—xon is pe™uli—rX the s™—les on the midE

—lso onthe s™—les of most of the ™—ud—l regionon l—ter—l fl—nk @whi™h ˜e—r the l—ter—l lineA —re re™t—ngul—rD

qw€u E€EQHIHF highly deepenedY their depth is quite the s—me until the

midElength of the ˜odyD where their depth —˜ruptly deE

™re—sesF „he ventr—l fl—nk s™—leErow whi™h lies immediE

€hylogeneti™ —n—lysis

—tely ˜elow the midEl—ter—l row —re m—de —lso ˜y highly

deepened s™—lesD ˜eing —˜out IGQ of those of the midE es —lre—dy pointed out ˜y voÂpezEer˜—rello 8

l—ter—l row indepth @pigF PD TAF elongthis s™—le rowD the —v—ttieri @PHHVD pF IHRUAD perleidiforms h—d — wide str—E

multipli™—tionof the s™—les st—rts —t the l—st two high tigr—phi™—l r—nge ™overing most of the „ri—ssi™D even if

deepened s™—lesX on qw€u E€EQHHW the first one douE they ™on™entr—te mostly in the widdle —nd v—te „ri—ssi™F

˜lesD thentriples pro™eedingposteriorlyD while on st must ˜e pointed out th—t wh—t is ™onsidered ˜y voÂE

qw€u E€EQHIH the first one dou˜les —nd keep this pezEer˜—rello 8 —v—ttieri @PHHVA —s ––the most ˜—s—l

st—te for one s™—leErowD then triples pro™eeding posterE monophyleti™ group99 m—de ˜y €erleidusD weridensi—

xew perleidiform f—mily @ystei™hthyesD e™tinopterygiiA from widdle „ri—ssi™ of ƒouth ghin— QTW

pigF T E puyu—nperleidus dengi qengD

tinD ‡u 8 ‡—ngD PHIPX det—ils

of the —rr—ngement of the

fl—nk s™—les in eD qw€u E

€EQHHWY fD qw€u E€EQHIHF

—nd €eltoperleidus is not —™tu—lly from the v—te „ri—ssi™ seems to ˜e — ™le—r su˜division˜etweenthe freshEw—ter

of gentr—l iurope ˜ut it is yielded m—inly ˜y the „ethE dweller of the qondw—n— —nd the m—rine „ethy—n genE

y—nwiddle „ri—ssi™ sequen™einthe ƒouthernelpsD in er—D with only hipteronotus @„intori IWWHA th—t ™ould ˜e

xorthern st—ly —nd ƒouthern ƒwitzerl—ndD —s well —s in ™onsidered — possi˜le ™ontinent—l t—xon —long the southE

ƒouthernghin—@vom˜—rdo PHHIY vom˜—rdo et —lF ern m—rgin of the then northern ™ontinentF „husD it is

PHIIAF not ™orre™t to —s™ri˜e the ––iurope—n99 gener— to the

voÂpezEer˜—rello 8 —v—ttieri @PHHVA dis™ussed v—ur—si— @voÂpezEer˜—rello 8 —v—ttieri PHHVX IHRRA or

some of the €erleidiformes ˜y performing — ™l—disti™ ™onsidering the group —s widely distri˜uted in €—nge—

—n—lysis to enlighten the interrel—tionships —mong themF @voÂpezEer˜—rello 8 —v—ttieri PHHVX IHRVAD —s those

pollowing their t—x— —nd ™h—r—™ter sele™tionD — further were ™ontinent—l regions while —lmost —ll the nonE

™l—disti™ —n—lysis w—s ™ondu™ted to —ssess the phylogeE qondw—ni—n t—x— —re fully m—rineF purthermoreD in

neti™ rel—tionships of puyu—nperleidus with other ––perE the light of the new finds in ƒouthern ghin—D —t the level

leidiforms99F snthe newd—t— m—trix we est—˜lishedD sevE of the genus the put—tive „ethy—n t—x— —re no more

er—l ™h—r—™ter st—tements were revised ˜—sed on the diE limited to ‡estern„ethys @nowiuropeA ˜ut —re ˜e™omE

re™t o˜serv—tions on spe™imens of €eltoperleidusD hipE ing known —lso in the i—stern „ethysD espe™i—lly the

teronotus —nd gtenogn—thi™hthys @eF„F —nd gFvF persF ƒouthernghin—flo™kF „hereD —p—rt from the t—xondeE

o˜serFAD —s well —s w—nliett— following ‡—de @IWQSA s™ri˜ed hereinD we —re —lso —w—re of the presen™e of

—nd rut™hinson @IWUQ—A ™ontri˜utionsF golo˜odus @ƒunet —lF PHHVAD vuopingi™hthys @ƒunet —lF

„he new d—t— m—trix ™onsists of IU t—x— —nd PU PHHWA —nd €erleidus @vom˜—rdo et —lF PHIIA while €elE

™h—r—™ters @„—˜F IA —nd the p—rsimony —n—lysis w—s perE toperleidus —nd other undes™ri˜ed perleidiform t—x— —re

formed using the exh—ustive se—r™h ™omm—nd @fr—n™hE still under study ˜y the rese—r™h group —t €u @feijE

—ndE˜oundA in €e €B version RFH˜IH for ‡indows ingAF st must —lso ˜e pointed out th—t while the „ri—ssi™

@ƒwofford PHHPAF ell ™h—r—™ters were equ—lly weightedF qondw—n— fish f—un—s —re somewh—t rel—ted to the

wultist—te ™h—r—™ters were tre—ted —s unordered to —llow „ethy—n m—rine f—un—s @€erleidiformes —nd m—y˜e ‚edE

—ll possi˜le hypotheses of evolution—ry rel—tionship fieldiformes if the l—tter will prove to ˜e rel—ted to €erE

—mong the ™h—r—™ter st—tesF „he se—r™h resulted in three leidiformes themselvesA the v—ur—si— f—un—s —re quite

most p—rsimonious trees @w€„A @tree lengthD „va TV different @see for inst—n™e ˆu—nd q—o PHIIAF „h—t there

stepsY ™onsisten™y indexD gsa HFRUY retention indexD —re ™lose rel—tionships ˜etween some of the „ethy—n

‚sa HFSWAF „he stri™t ™onsensus tree of these tree equ—lly f—un—s —nd the qondw—n— freshEw—ter environments is

p—rsimonious trees w—s shown in pigF UF „o —ssess ™l—de —lso supported ˜y the ™ommonpresen™eof some isopod

supportsD — non p—r—metri™ ˜ootstr—p —n—lysis w—s ™—rE ™rust—™e—ns in the widdle „ri—ssi™ of southern ghin—

ried out in€e €B usingSHH repli™—tesF @vuoping p—un—A —nd eustr—li— @pu et —lF PHIHAF

‚eg—rding the p—leoenvironment—l —nd p—leo˜ioE yur —n—lysis m—t™hes well the p—leo˜iogeogr—phi™

geogr—phi™ distri˜utionof the perleidiformsD there rel—tionships of the qondw—n— freshEw—ter t—x— @pigF UAD

QUH ƒun FD vom˜—rdo gFD „intori eFD ti—ng hFD r—o ‡FD ƒun ‰F 8 vin rF

pigF U E gonsensus of Q most p—rsiE

monious trees of TV steps

with — gonsisten™y sndex of

HFRU —nd — ‚etention sndex

of HFSWF

—n—lysis is ˜—sed on voÂpezEer˜—rello 8 —v—ttieri —nd the hypothesis of their monophyly —ppe—rs pl—usiE

˜le onthe ˜—sis of our resultsD ˜ut the low ˜ootstr—p @PHHVA oneF e ™omplete revision of €erleidiformesD —s

support v—lue @IV7A requires further v—lid—tionsF well —s the se—r™h to ™l—rify the rel—tionships with other

emong the „ethy—n t—x—D the very spe™i—lized gtenogE ––su˜holoste—n99 groups is ˜eyond the s™ope of this

n—thi™hthys —nd pel˜eri— forms — monophyleti™ group —s workF es we —re working on further perleidiform m—E

the sister group of —ll the rem—ining perleidiformsY teri—l from the widdle „ri—ssi™ of ƒouthernghin—Dwe

puyu—nperleidus is more ™losely rel—ted to the „ethy—n will ™onsider for — ™omplete phylogeneti™ —n—lysis —fter

t—x— hipteronotusD €erleidusD —nd the ™l—de ™ont—ining

the des™riptionof those newt—x—F „his will imply —lso

weridensi— —nd €eltoperleidusD th—nthe other perleidiE

™h—nges in the ™h—r—™ter settingF

formsF rowever these t—x— form — polytomyD showing —

unresolved rel—tionshipF „he qondw—n— deepE˜ody e™knowledgmentsF ‡e —re deeply inde˜ted to h—vide ƒ—ssi for

running the €e € —n—lysis —nd for dis™ussing the resultsF „h—nks —lso perleidiforms @iFeF gleithrolepisD gleithrolepidin—D ryE

go to those who helped in™olle™tingthe fossilsD inp—rti™ul—r wrF hu—n dropessumA —nd the „ethys t—x— @pel˜eri— —nd hipteroE

@hu—n ‡eig—ngAF ‡e —™knowledge —lso two —nonimous referees for

notusD for inst—n™eA does not form — sister group —s in

revising — previous version of his p—per —nd „F furginfor his helpful È

voÂpezEer˜—rello 8 —v—ttieri @PHHVA —nd the sh—red

—dvi™eF „his study w—s fin—n™i—lly supported ˜y €eking niversityD

™h—r—™ters reg—rding the ˜ody depth should ˜e — ™onE

xƒpg @gr—nt RHUHPHHID RHWPHIPRHHPAD ƒt—te uey v—˜or—tory of €—l—eoE

vergent —™quisition r—ther th—n — phyleti™ re—sonF ˜iology —nd ƒtr—tigr—phy @x—njing snstitute of qeology —nd €—l—eonE

‡e must point out th—t sever—l other des™ri˜ed tologyD geƒA @xoF IPQIHUAD —nd — spe™i—l gr—nt from the he—n of niE

versit—Á degli ƒtudi of wil—no for the st—li—n p—rti™ip—tionF perleidiforms h—ve not ˜een ™onsidered hereD —s this

xew perleidiform f—mily @ystei™hthyesD e™tinopterygiiA from widdle „ri—ssi™ of ƒouth ghin— QUI

‚ipi‚ixgiƒ

ferg vF ƒF @IWQUA E e ™l—ssifi™—tionof fishElike verte˜r—tesF q—rdiner fF qFD w—isey tF qF 8 vittlewood hF „F @IWWTA E

fullF e™—dF ƒ™iF ‚ƒƒD RX IPUUEIPVH ‘sn‚ussi—n“F snterrel—tionships of ˜—s—l neopterygi—nsF snX ƒti—ssny

frough tF @IWQIA E ynfossil fishes from the u—roo ƒystem vF tFD €—renti vF ‚F 8 tohnson qF hF @idsA E snterreE

—nd some gener—l ™onsider—tions on the ˜ony fishes of l—tionships of fishesX IIUEIRTD e™—demi™ €ressD vonE

the „ri—ssi™ periodF €ro™F oolF ƒo™F vondFD IWQIX PQSE donF

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