‚ivist— st—li—n— di €—leontologi— e ƒtr—tigr—fi— volume IPI noF I IP plsF ppF TIEIHI w—r™h PHIS

„ri qixi‚e qv‰g‰wi‚sƒD ei s€ig„ix exh e‚g„sgeD exh eƒƒygse„ih

wyvv ƒu pe xe yp „ri vy‡i‚ €visƒ„ygixi e‚he ‚s†i‚ ƒig„syx

@xy‚„ri‚x s„ev‰A

I P

qese g‚s€€e 8 qsexv ge ‚esxi‚s

‚e™eivedX he™em˜er IUD PHIRY —™™eptedX pe˜ru—ry TD PHIS

uey wordsX vower €leisto™eneD erd— ‚iverD fiv—lve system—ti™sD in f—™t its fossiliferous sediments were soon —dopted —s

fiostr—tigr—phyD €—l—eo™lim—tologyF

the ˜ound—ry str—totype for the €i—™enzi—n ƒt—ge @eFgF

€—reto IVTSY f—r˜ieri IWTUAF ƒu˜sequentlyD integr—ted

e˜str—™tF „he vower €leisto™ene m—rine su™™ession of the erd—

™—l™—reous pl—nkton ˜iostr—tigr—phi™ studies ˜y ‚io et ‚iverD ™ropping out —t g—stell9erqu—to @xorthern st—lyAD is well exE

—lF @IWVVA —nd ˜y ‚—ffi et —lF @IWVWA demonstr—ted th—t — posedD ™ontinue —nd ri™hly fossiliferousF st ™onsists of s—ndstonesD siltE

stones —nd ™l—ystones —nd it is topped ˜y ™ontinent—l ™onglomer—tes hi—tus w—s present right —t the ˜—se of the str—totypeD

whi™h indi™—te — m—jor se— level dropF sn the present p—per qly™ymeris

m—king thus this —re— unsuit—˜le to define — str—totypeF gly™ymerisD qly™ymeris infl—t—D qly™ymeris insu˜ri™—D eequipe™ten s™—˜E

elthough numerous studies h—ve ˜een performed on

rell—D eequipe™ten oper™ul—ris —nd er™ti™— isl—ndi™— —re des™ri˜ed —nd

this p—rt of the erd— ‚iver se™tionD less h—s ˜een done revisedF sn —dditionD the —sso™i—ted f—un— is illustr—tedD —nd its ˜iostr—tiE

on the upper p—rt of the su™™essionD whi™h extends gr—phi™ —nd p—l—eo™lim—ti™ signifi™—n™e is —n—lyzedF „his study points

out th—tX IA the f—un— h—s — high ˜iodiversityD ™omprising ISW t—x—Y downstre—m the ˜ridge lo™—ted —t the entr—n™e of the

˜iv—lves —re domin—ntD followed ˜y g—stropodsD ™or—lsD serpulidsD ˜r—E

town of g—stell9erqu—to @homini™i PHHID PHHRY woneE

™hiopodsD e™hinoidsD —rthropods @eFgF ˜—rn—™lesAD ˜ryozo—ns —nd s™—E

g—tti et —lF PHHIAF „his p—rt of the su™™ession is very ri™h

phopodsY PA the l—st o™™urren™es of the ˜iv—lves eequipe™ten s™—˜rell—D

in m—™rofossils —nd it ™overs without signifi™—nt g—ps qly™ymeris infl—t— —nd gh—m— pl—™entin—D the first o™™urren™e of er™E

the i—rly €leisto™eneD — keyEtime interv—l to underst—nd ti™— isl—ndi™— —nd the l—st o™™urren™es of the g—stropods „urritell— triE

™—rin—t— pliore™ens —nd x—ss—rius prysm—ti™us suggest — l—te qel—si—nE the ™lim—te ™h—nge le—ding to the widdle €leisto™ene

g—l—˜ri—n —ge @i—rly €leisto™eneA for the investig—ted m—rine su™™esE

™ontinent—l gl—™i—tionF „he i—rly €leisto™ene w—s ™h—rE

sionY QA the o™™urren™e of ˜ore—l guestsD su™h —s er™ti™— isl—ndi™—D €seuE

—™terized ˜y sever—l ™lim—ti™ os™ill—tions linked to gl—E

d—mussium peslutr—e —nd possi˜ly wytilus edulisD testify to — ™lim—ti™

™i—lGintergl—™i—l ™y™lesD whi™h —ffe™ted —lso the wediterE ™h—nge with — shift from w—rm to ™old se—w—ter temper—turesF

r—ne—n —re— —nd ™ulmin—ted —t the end of the i—rly

€leisto™ene with the onset of the northern hemisphere

™ontinent—l gl—™i—tions @eFgF fertini PHHIY gl—rk et —lF sntrodu™tion

PHHTAF

„he erd— ‚iver m—rine su™™essionD ™ropping out „his work represents the first of — series of p—persD

in xorthern st—lyD is well exposedD ™ontinue —nd very whi™h will ˜e —imed to des™ri˜e —nd revise the most

™ontroversi—l —nd question—˜le €lio™eneE€leisto™ene ri™h in fossil inverte˜r—tesF

mollusk spe™ies found —long the erd— se™tionF rereD „he str—tigr—phi™—lly lower p—rt of the su™™ession

we fo™us on the des™riptions —nd revisions of six spe™ies @—n™le—nE€i—™enzi—n in —geA ™ropping out ˜etween the

˜elonging to the gener— qly™ymerisD eequipe™ten —nd town of vug—gn—no †—l d9erd— —nd g—stell9erqu—to

er™ti™—F „he —sso™i—ted mollusk f—un— is —lso illustr—ted @xorthern st—lyA w—s the su˜je™t of sever—l rese—r™hesY

I hip—rtimento di ƒ™ienze dell— „err— ––erdito hesio99D niversit—Á degli ƒtudi di wil—noD †i— w—ngi—g—lli QRD PHIQQ wil—noD st—lyF

iEm—ilX g—i—F™ripp—dunimiFit

P ‚iserv— x—tur—le qeologi™— del €i—™enzi—noD †i— ƒ™—lin—t— ysped—le RGTD g—stell9erqu—toD PWHIRD st—lyF iEm—ilX gFr—ineridp—r™hiemili—o™™ident—leFit

TP gripp— qF 8 ‚—ineri qF

˜y ™ontinent—l ™onglomer—tes indi™—ting — m—jor se— leE

vel drop@pigF PAF

„he —ltern—tions of thi™k siltyEmuddy interv—ls

@hemipel—gi™ suspension settlingA —nd s—ndstones @pigF

PA whi™h ™h—r—™terize the over—ll m—rine su™™ession

™—n ˜e interpreted —s floodEgener—ted delt—Efront lo˜es

re™ording the fin—l s—ndy deposition of highEdensity

flows triggered ˜y river floods @homini™i PHHIY gripp—

et —lF in progressAF „his suggests th—t it w—s deposited in

pigF I E ƒket™h m—pwith the positionof the vower €leisto™ene

— te™toni™—lly —™tive setting during ph—ses of —dv—n™e of

g—stell9erqu—to porm—tion ™ropping out —long the erd—

f—n delt—s @homini™i PHHIY pellettiD persF ™ommFAF

‚iverD ‡estern imili—D st—ly @modified —fter €ervesler et —lF

PHII —nd gripp— PHIQAF f—se of the se™tion —t

RRëSI9IPFS99xY WëSP9PPFR99iF

w—teri—l —nd methods

possil spe™imens were ™olle™ted using — ˜ed ˜y ˜ed —ppro—™h

—nd the ˜iostr—tigr—phi™ —nd p—l—eo™lim—ti™ signifi™—n™e

from PHH fossiliferous ˜eds @one s—mple for e—™h ˜edAD positioned on the

of some of its t—x— is —n—lyzed with the —im of underE erd— ‚iver log meter ˜y meter @pigF PY eppendixAF e ˜ulk s—mple of —t

le—st HFS liters w—s s—mpledY s—mpling w—s then ™ontinued until new st—nding how the erd— ˜iot— responded to the ™lim—ti™

spe™ies were foundF „he ™olle™ted spe™imens were then prep—redD

™h—nges of the i—rly €leisto™eneF

w—shed —nd ™le—ned from the sediment using —n —ir drillD in ™—se of h—rd

sedimentD or — s™—lpelD in ™—se of soft sedimentF „he length —nd the

width of fossil spe™imens were me—sured using — ™—liper @to the ne—rest

HFI mmA —nd —t e—™h of them h—s ˜een —ssigned — unique sh to identify qeologi™—l setting

it @sh usedX egqY progressive num˜ers from the ˜—se to the top of the

se™tionD see eppendixAF ivery fossil spe™imen w—s identified —t generi™ „he studied m—rine su™™ession ˜elongs to the

—nd spe™ifi™ level using the ™ommon te™hniques of identifi™—tion —nd upper p—rt of the g—stell9erqu—to porm—tionD ˜elonging

™l—ssifi™—tionD in order to o˜t—in — ™onsistent t—xonomy @see next p—r—E

to the –u—tern—rio w—rino ƒupersynthem9 @qel—si—nE

gr—phAF e ™omplete list of the spe™ies found in e—™h str—tigr—phi™ ˜ed is

g—l—˜ri—nY g—l—˜rese 8 hi hio PHHWAD whi™h ™rops out

reported in the eppendixF

in xorthern st—ly —long the erd— ‚iver —t g—stell9erE „he fossil preserv—tion is gener—lly goodY —rti™ul—ted ˜iv—lve

spe™imens in life position —re present @—uto™hthonous —ssem˜l—gesAD qu—to @‡estern imili—D xorthern st—lyA @pigF IAY it exE

˜ut usu—lly the —ssem˜l—ges —re domin—ted ˜y dis—rti™ul—ted v—lvesF tends downstre—m the ˜ridge present —t the entr—n™e of

xotwithst—nding dis—rti™ul—tionD the shells —re fresh —nd preserve the

the town tow—rd xExi for ne—rly P km @˜—se —t

fr—gile spiny orn—ment—tion —nd in some ™—ses the ™olor p—tternD indiE

RRëSI9IPFS99xY WëSP9PPFR99iAF

™—ting in situ dis—rti™ul—tion @˜y ˜iotur˜—tionA or shortEdist—n™e tr—nsE

„he se™tion ˜elongs to the g—stell9erqu—to piggyE port @see pigF P for the position of the tr—nsported ˜edsAF ƒhells —re

usu—lly whiteD r—rely ˜l—™k or greyD showing little —˜r—sion @see t—˜le ˜—™k ˜—sin @‚overi 8 „—vi—ni PHHQAD whi™h developed

PD pF QRP in homini™i PHHI for t—phonomi™ sign—tures —nd interpret—E

—fter the fr—gment—tion of the €o €l—in foredeepin the

tionsAF „he t—phonomi™ sign—tures —nd the ˜iof—˜ri™ thus suggest the

l—te wessini—n @g—l—˜rese 8 hi hio PHHWAF „he ˜—sin is

presen™e in some ˜eds of tr—nsport or winnowingD ˜ut not over dist—n™e

filled ˜y — sediment—ry su™™ession of l—te wessini—n to

@mostly p—r—uto™hthonous —ssem˜l—gesAF purthermoreD in the m—jority

rolo™ene —geD org—nized in — l—rge s™—le tr—nsgressiveE of ™—sesD shells do not show the presen™e of mi™ro˜oringD whi™hD in

sh—llow w—terD usu—lly indi™—tes — prolonged exposure to ˜iologi™—l or regressive ™y™le ™ontrolled ˜y te™toni™s @woneg—tti et —lF

physi™—l f—™tors —fter de—thF „his l—™k of mi™ro˜oring thus suggests —

PHHIAF feds form — regul—r mono™line dipping tow—rds

qui™k ˜uri—l —nd short residen™e time on the se— floor th—t h—ve preE

the xExi —nd no m—jor f—ults o™™ur in this —re— @woneE

served the shells fresh —nd their orn—ment—tion int—™t @€—rsonsEru˜E

g—tti et —lF PHHIAF „he m—rine l—yers —t the ˜—se of the ˜—rd et —lF PHIRAF

g—stell9erqu—to ˜—sin ™onsist in deepse— sediments deE

posited —fter the end of the wessini—n s—linity ™risis

@g—l—˜rese 8 hi hio PHHWAD when m—rine ™onditions

possil —ssem˜l—ges

were restored in the wediterr—ne—n se—Y upw—rd they

p—ss to slope —nd shelf f—™ies —nd then through — regresE „he f—un— is ™h—r—™terized ˜y — high ˜iodiversityD

sive trend to the widdle €leisto™ene —lluvi—l ™ontinent—l in™luding ˜oth inf—un—l —nd epif—un—l spe™iesD of sh—lE

depositsD whi™h represent the fin—l retre—t of the se— in low —nd deepinfr—littor—l —nd ™ir™—littor—l environE

this —re— —nd the est—˜lishment of — ™ontinent—l environE mentsY — det—iled p—l—eoe™ologi™—l —n—lysis of the erd—

ment with verte˜r—te f—un—s —nd fresh w—ter mollusks ˜iot— is in progress —nd will ˜e the su˜je™t of — sep—r—te

@gig—l— pulgosi IWUTY €elosio 8 ‚—ffi IWUUY gi—ngheE p—perF

rotti et —lF IWWUAF „he f—un—D here ex—minedD is ™omposed ˜y ISW

„he studied se™tion is PRH mEthi™k —nd ™onsists of t—x— @„—˜F IY €ls IEISA ™oming from PHH shell ˜edsD of

whi™h ˜iv—lves —re domin—nt with IHS t—x—D followed ˜y s—ndstonesD siltstones —nd ™l—ystones ˜ounded —t the top

wollusk f—un— of the vower €leisto™ene erd— ‚iver se™tion @xorthern st—lyA TQ

g—stropods @RR t—x—A —nd — few ™or—ls @Q t—x—A —nd serE

pulids @P t—x—AY ˜r—™hiopodsD e™hinoidsD —rthropods @eFgF

˜—rn—™lesAD ˜ryozo—ns —nd s™—phopods do —lso o™™ur in

the f—un—D testifying th—t it is ™h—r—™terized ˜y — high

˜iodiversityF

es s—id —˜oveD ˜iv—lves represent the domin—nt

t—xon in the erd— f—un—Y in p—rti™ul—rD here we fo™us

on the des™ription —nd revision of three ˜iv—lve gener—

@qly™ymerisD eequipe™ten —nd er™ti™—AF por these re—E

sonsD we m—inly de—l with the pro˜lem—ti™s th—t ™h—rE

—™terize ˜iv—lve system—ti™sF „he ™l—ssifi™—tion of most

spe™imens w—s m—de diffi™ult ˜y the f—™t th—t ˜iv—lve

t—xonomy is — ™ompli™—ted topi™ in ™onst—nt evolutionY

in f—™tD —s fieler 8 wikkelsen @PHHTD pF PPRA o˜served

––mu™h of the t—xonomi™ inst—˜ility in ˜iv—lve rese—r™h is

not — result of ™onfli™ting hypotheses of rel—tionshipsD

˜ut one of —n over—˜und—n™e of —v—il—˜le n—mes99 —nd

this is the m—in pro˜lem we de—lt with the ˜iv—lve ™l—sE

sifi™—tion of the erd— —ssem˜l—gesF e™™ording to timeE

nez et —lF @PHHWA the st—tus of m—ny gener— is still unE

™ert—in —nd spe™ies —re —ssigned to different gener— deE

pending on the —uthorsY nonethelessD there is — ™ert—in

st—˜ility in the spe™ies ™on™ept th—t —llows —dequ—te

identifi™—tion —t the spe™ies levelsF „he „re—tise on snE

verte˜r—te €—leontology E wollus™—D p—rt x @gox et —lF

IWTWD IWUIA h—s —lw—ys ˜een used —s — st—rting point for

˜iv—lve system—ti™sD ˜ut it is now outd—ted —nd hen™e do

not in™lude re™ent findingsF

sn the l—st few ye—rsD ˜iv—lve workers h—ve ˜een

in™re—singly involved in l—rgerEs™—le phylogeneti™ —n—E

lysesD using — ™om˜in—tion of morphologi™—lD p—l—eonE

tologi™—l —nd mole™ul—r d—t— sour™es to investig—te evoE

lution—ry p—tterns —nd refine system—ti™s for v—rious

p—rts of the ˜iv—lve treeF sn the me—ntimeD — revised

volume of „re—tise on snverte˜r—te €—leontology ± wolE

lus™—D p—rt x is in prep—r—tion @v—wren™eD niversity of

u—ns—sD €—leontologi™—l instituteAF

„he ‡orld ‚egister of w—rine ƒpe™ies @‡o‚wƒA

h—s ˜een ™onsulted for resolving ™riti™—l issues of noE

men™l—ture @v—lid n—me —nd so onAF sn some ™—sesD howE

everD we preferred to m—int—in the spe™ifi™ n—me used

for fossil spe™imens inste—d of the one used for ext—nt

spe™ies —s the link with modern spe™ies h—s not —lw—ys

˜een verified —nd the l—tter m—y differ — lot from the

™orresponding fossil ones ‘„ellin— ™or˜is @fossilA r—ther

th—n „ellin— ™—rni™olor @re™entAY e™teon semistri—tus @fosE

silA inste—d of e™teon torn—tilis @re™entA“F

pigF P E ƒtr—tigr—phi™ log of the erd— se™tion in whi™h —re lo™—ted

the most signifi™—nt first —nd l—st o™™urren™esY — simplified

sedimentologi™—l des™ription is —lso shown @gourtesy of hrF pF pellettiAF

TR gripp— qF 8 ‚—ineri qF

List of taxa occurring in the Arda marine section Bivalves Nucula placentina (Lamarck, 1819) Lutraria angustior Philippi, 1844 Nucula sulcata Bronn, 1831 Lutraria oblonga (Gmelin, 1791) Nucula sp. Lutraria sp. Nuculana pella (Linnaeus, 1758) Ensis ensis (Linnaeus, 1758) 3 Saccella commutata (Philippi, 1844) Tellina albicans Gmelin, 1791 Arcoidea indet. Tellina corbis Sowerby, 1867 Arca noae Linnaeus, 1758 Tellina incarnata Linnaeus, 1758 Arca tetragona Poli, 1795 Tellina pulchella Lamarck, 1818 Arca sp. Tellina serrata Brocchi, 1814 Barbatia mytiloides (Brocchi, 1814) Tellina tenuis da Costa, 1778 Anadara cf. A.diluvii (Lamarck, 1805) Tellina sp. 4 Striarca lactea (Linnaeus, 1758) Moerella distorta (Poli, 1791) Bathyarca sp. Donax cf. D.trunculus Linnaeus, 1758 Glycymeris glycymeris (Linnaeus, 1758) Donax cf. D.venustus Poli, 1795 Glycymeris inflata (Brocchi, 1814) Donax sp. Glycymeris insubrica (Brocchi, 1814)1 Solecurtidae indet. Glycymeris sp. Solecurtus scopula (Turton, 1822) Mytilus edulis Linnaeus, 1758 Azorinus chamasolen (da Costa, 1778) Mytilus galloprovincialis Lamarck, 1819 Arctica islandica (Linnaeus, 1767) Mytilus sp. Glossus humanus (Linnaeus, 1758) 5 Pinna sp. Venus nux Gmelin, 1791 Amusium cristatum (Bronn, 1827) Venus sp. Pseudamussium peslutrae (Linnaeus, 1771)2 Pitar rudis (Poli, 1795) Aequipecten opercularis (Linnaeus, 1758) Callista chione (Linnaeus, 1758) Aequipecten scabrella (Lamarck, 1819) Pelecyora brocchi (Deshayes, 1836) Mimachlamys varia (Linnaeus, 1758) Chamelea gallina (Linnaeus, 1758) Talochlamys cf. T.multistriata (Poli, 1795) Clausinella fasciata (da Costa, 1778) Flexopecten flexuosus (Poli, 1795) Clausinella sp. Flexopecten glaber (Linnaeus, 1758) Dosinia lupinus (Linnaeus, 1758) Pecten flabelliformis (Brocchi, 1814) Dosinia sp. Pecten jacobaeus (Linnaeus, 1758) Polititapes cf. P.rhomboides (Pennant, 1777) Anomia ephippium Linnaeus, 1758 Polititapes rhomboides (Pennant, 1777) Anomia sp. Polititapes senescens (Cocconi, 1873) Monia patelliformis (Linnaeus, 1761) Polititapes sp. Loripes lacteus (Linnaeus, 1758) Timoclea ovata (Pennant, 1777) Lucinoma borealis (Linnaeus, 1767) Corbula gibba (Olivi, 1792) Loripinus fragilis (Philippi, 1836) Hiatella rugosa (Linnaeus, 1767) Lucinella divaricata (Linnaeus, 1758) Panopea glycymeris (Born, 1778) Chama gryphoides Linnaeus, 1758 Panopea sp. Chama placentina (Defrance, 1817) Pholas dactylus Linnaeus, 1758 Carditidae indet. Teredo sp. Venericardia sp. Pandora inaequivalvis (Linnaeus, 1758) Cardites antiquatus (Linnaeus, 1758) Thracia pubescens (Pulteney, 1799) Astarte fusca (Poli, 1791) Thracia sp. Astarte sp. Clavagella sp. Cardium indicum Lamarck, 1819 Ostrea edulis Linnaeus, 1758 Acanthocardia aculeata (Linnaeus, 1758) Ostrea sp. Acanthocardia echinata (Linnaeus, 1758) Saccostrea cf. S.cucullata (Born, 1778) Acanthocardia paucicostata (G. B. Sowerby II, 1834) Acanthocardia tubercolata (Linnaeus, 1758) Acanthocardia sp. Papillicardium papillosum (Poli, 1791) Papillicardium sp. Parvicardium exiguum (Gmelin, 1791) Laevicardium crassum (Gmelin, 1791) Laevicardium oblongum (Gmelin, 1791) Laevicardium sp. Mactra stultorum (Linnaeus, 1758) Spisula subtruncata (da Costa, 1778)

wollusk f—un— of the vower €leisto™ene erd— ‚iver se™tion @xorthern st—lyA TS

Gastropods Diodora graeca (Linnaeus, 1758) Nassarius musivus (Brocchi, 1814) Trochidae indet. Nassarius mutabilis (Linnaeus, 1758) Calliostoma cf. C.conulus (Linnaeus, 1758) Nassarius obliquatus (Brocchi, 1814) Jujubinus striatus (Linnaeus, 1758) Nassarius prysmaticus (Brocchi, 1814) Jujubinus sp. Nassarius semistriatus (Brocchi, 1814) Diloma patulum (Brocchi, 1814) Nassarius sp. Turritella tricarinata (Brocchi, 1814) Mitra sp. Turritella sp. Acteon semistriatus Glibert, 1952 Turridae indet. Acteon sp. Aporrhais uttingeriana (Risso, 1826) Ringicula auriculata (Ménard de la Groye, 1811) Aporrhais pespelecani (Linnaeus, 1758) Ringicula sp. Aporrhais sp. Cylichna cylindracea (Pennant, 1777) Calyptraea chinensis (Linnaeus, 1758) Conus ventricosus Gmelin, 1791 Calyptraea sp. Conus sp. Capulus ungaricus (Linnaeus, 1758) Pyramidella sp. Xenophora crispa (König, 1825) Naticarius stercusmuscarum (Gmelin, 1791) Other macroinvertebrates Naticarius sp. Cladocora sp. Euspira sp. Flabellum sp. Neverita josephinia Risso, 1826 Corals indet. Neverita sp. Terebratula sp. Galeodea echinophora (Linnaeus, 1758) Echinoids indet. Epitonium tiberii (de Boury, 1890) Dentalium sp. Epitonium turtonis (Turton, 1819) Serpulorbis sp. Bolinus sp. Ditrupa sp. Murex sp. Barnacle indet. Nassarius cf. N.gibbosulus (Linnaeus, 1758) Bryozoa indet.

Nassarius cf. N.clathratus (Born, 1778)

I

„—˜F I E vist of t—x— o™™urring in the erd— m—rine se™tionF elso known —s qly™ymeris numm—ri— @vinn—eusD IUSVA or qly™ymeris viol—™es™ens

P Q R

elso known —s €seud—mussium septemr—di—tum @wullerD IUUTAY elso known —s „ellin— nitid— @€oliD IUWIAY elso @v—m—r™kD IVIWAY È

S

elso known —s †enus multil—mell— @v—m—r™kD IVIVAF known —s „ellin— distort— @€oliD IUWIAY

fiostr—tigr—phi™ impli™—tions „he —n—lysis of the ˜iostr—tigr—phi™ signifi™—n™e of

the r—nge of the spe™ies of the erd— f—un— is m—de diffiE

™ult ˜y two m—in pro˜lemsX IA ev—il—˜le liter—tureX d—t— „he erd— f—un— is m—inly ™omposed of spe™ies

on r—nges of extin™t spe™ies —re very fr—gment—ry —nd th—t m—de their first —ppe—r—n™e in the wio™ene @„ortoE

™onfusing —s often sever—l —uthors give — different time ni—nA or in the i—rly €lio™ene @‚—ffi et —lF IWVSAY this

of dis—ppe—r—n™e for the s—me spe™ies @often referring to f—un— est—˜lished in the wediterr—ne—n ƒe— —fter the

lo™—l dis—ppe—r—n™eAY see for ex—mple the ™—se of the wessini—n s—linity ™risisD when norm—l m—rine w—ters

˜iv—lves gh—m— pl—™entin— —nd qly™ymeris infl—t— deE inv—ded the ˜—sin from the etl—nti™ @‚—ffi et —lF IWVSAF

s™ri˜ed ˜elowF PA ƒ—mpling ˜i—sX m—™rofossils —re disE „he m—jority of the spe™ies found in the erd— ‚iver

™ontinuously o™™urring —long the se™tionY when r—re su™™ession ˜elongs to the modern mollusk f—un— ™urE

they m—y h—ve not ˜een found during s—mple ™olle™tion rently thriving in our se—sY thusD they do not provide

@ƒignorEvipps effe™tA @ƒignor 8 vipps IWVPAF sn —ddiE ˜iostr—tigr—phi™ inform—tionsF

tionD the l—st o™™urren™es of some spe™ies m—y ˜e the emong the erd— ˜iot—D there —re however spe™ies

result of — ™h—nge in the p—l—eoe™ologi™—l ™onditionsD th—t m—de their first —ppe—r—n™e or ˜e™ome extin™t durE

thus representing — lo™—l extin™tionF gonsidering these ing the time of deposition of the se™tionD thus representE

pro˜lems —nd using — ™—utious —ppro—™hD it h—s ˜een ing — useful tool to —ssess the —ge of the m—rine su™™esE

possi˜le to identify fifteen spe™ies of ˜iv—lves —nd g—sE sionD t—king into —™™ount the pro˜lems des™ri˜ed ˜elowF

tropods firstly or l—stly o™™urring —long the se™tionF „he €revious works @eFgF homini™i PHHIY woneg—tti et —lF

most signifi™—nt of these first —nd l—st o™™urren™es —re PHHIA inferred —n i—rly €leisto™ene —ge for the f—un—

here des™ri˜ed @pigF PAF under ex—mD mostly ˜—sed on the ™omp—rison with the

eequipe™ten s™—˜rell— l—stly o™™urs in the su™™esE ˜etter known ƒtirone ‚iver se™tion @homini™i PHHIY

qunderson et —lF PHIPAD ™ropping out IH km ƒi from sion —t RSFTS mY —™™ording to woneg—tti 8 ‚—ffi @PHHIA

the erd— ‚iver ƒe™tionD whi™h h—s very simil—r ˜iot—F it ˜e™—me extin™t —t PFI w—F

TT gripp— qF 8 ‚—ineri qF

—˜out PFHH w— —go in the ƒ—nterno †—lley @xorthern gh—m— pl—™entin— —nd qly™ymeris infl—t— show

their l—st o™™urren™e in the se™tion well ˜efore the py st—lyAD ˜—sed on m—gnetostr—togr—phi™ eviden™e —nd

of eF isl—ndi™— @respe™tively —t QUFHS m —nd RP mAF „his r—diometri™ —ge o˜t—ined ˜y ™or—lsF

is in —greement with the findings of ‚—ffi @IWVTA —nd e™™ording to ‚io et —lF in †—n gouvering @IWWUAD

woneg—tti 8 ‚—ffi @PHHIA who suggested th—t these speE the str—tigr—phi™—lly lowest level where eF isl—ndi™— o™E

™ies got extin™t in the wediterr—ne—n ƒe— —t IFV w—Y ™urs seems to ˜e right in the erd— —nd the ƒtirone se™E

howeverD some —uthors o˜served th—t they survived —lso tionsY the first —ppe—r—n™e of eF isl—ndi™— in the erd—

—fter this time interv—l —nd they pro˜—˜ly died out —t the se™tion should thus ˜e ˜r—™keted ˜etween IFUUEPFHH w—

end of the i—rly €leisto™ene @qre™o IWUHY g—protti depending on the different interpret—tion dis™ussed

IWUPAF —˜oveF

x—ss—rius prysm—ti™us l—stly o™™urs —t PPIFRH m in „his spe™ies ˜e™—me extin™t in the wediterr—ne—n

the se™tionF e™™ording to sever—l —uthors @ƒo™iet—Á st—li—E ƒe— —round WFV k—D ˜ut it is still living now—d—ys in the

n— di w—l—™ologi—D ™orresponding we˜ p—geAD this t—xon etl—nti™ y™e—n —long the emeri™—n ™o—st —nd in iurope

got extin™t —t the end of the i—rly €leisto™ene —nd this —long the ™o—sts of s™el—ndD qre—t frit—inD ƒ™—ndin—vi—n

seems to ˜e in —greement with our findingsF peninsul—D henm—rkD xetherl—ndsD felgium up to the

„urritell— tri™—rin—t— h—s is l—st o™™urren™e —t northern ™o—st of pr—n™e @fritt—ny —nd xorm—ndyA

PPRFPH m in the se™tionF „his spe™ies h—s three su˜EspeE @h—hlgren et —lF PHHHAF

™iesX „F tri™—rin—t— tri™—rin—t—D „F tri™—rin—t— pliore™ens „he presen™e of mollusk spe™ies th—t —re known

—nd „F tri™—rin—t— ™ommunisD whi™h differ ˜y few ™h—rE to dis—ppe—r —t the end of the i—rly €leisto™ene in the

—™ters in the orn—ment—tion —nd ˜y the outline of the upper p—rt of the se™tion @t—king into —™™ount the proE

—perture @see forghi 8 †e™™hi PHHSAF „he erd— spe™iE ˜lem—ti™s expl—ined —˜oveA —nd the l—™k of t—x— of

mens —re mostly simil—r to „F tri™—rin—t— tri™—rin—t— —nd stri™tly widdle €leisto™ene —geD —llow ex™luding —

„F tri™—rin—t— pliore™ens r—ther th—n to „F tri™—rin—t— younger —ge for the erd— se™tion ™ropping out downE

™ommunisY —lthough it is diffi™ult to dis™ern ˜etween stre—m the ˜ridge of g—stell9erqu—toF „he str—tigr—phi™

the former two spe™iesD the m—jority of the erd— spe™iE r—nges of the ˜iv—lves —nd g—stropods suggest —n i—rly

mens ˜elonging to „urritell— seem to ˜e more simil—r to €leisto™ene —geD more ex—™tly l—te qel—si—nEg—l—˜ri—nD

„F tri™—rin—t— pliore™ensF sn p—rti™ul—rD „F tri™—rin—t— —nd this is supported ˜y prelimin—ry d—t— on n—nnofosE

™ommunis is ™onsidered — ‚e™ent su˜spe™iesD where—s silsF purther —n—lyses —re now in progress ˜—sed on forE

„F tri™—rin—t— tri™—rin—t— —nd „F tri™—rin—t— pliore™ens —minifers —nd m—gnetostr—tigr—phy to ˜etter ™onstr—in

—re respe™tively wio™eneE€lio™ene —nd vower €leistoE the —ge of the se™tionF

™ene su˜spe™iesF „he erd— spe™imens thus ˜elong to „F

tri™—rin—t— pliore™ensD the spe™ies th—t got extin™t —t the

end of the i—rly €leisto™eneF €—l—eo™lim—ti™ impli™—tions

elthoug the l—st o™™urren™es of xF prysm—ti™us

„he evolution of the ˜iot— of the wediterr—ne—n —nd „F tri™—rin—t— pliore™ens in the se™tion m—y ˜e due

˜—sin h—s ˜een strongly —ffe™ted ˜y €lio™eneE€leisto™ene to — p—l—eoe™ologi™—l ™ontrol @pigF PAD prelimin—ry d—t—

™lim—ti™ ™h—nges @‚—ffi et —lF IWVSAF sn the erd— se™tion ˜—sed on n—nnofossil seem to ˜e in —greement with the

the most import—nt ˜ioti™ event from — p—l—eo™lim—ti™ —ge given ˜y these two t—x— @fottini 8 ‚—ffiD persF

perspe™tive is the —ppe—r—n™e of the ––fore—l @or xorthE ™ommFAF

ernA quests99 @fqAF „hese spe™iesD m—inly mollusksD osE e very import—nt ˜ioti™ event in the erd— su™™esE

tr—™ods —nd for—minifersD migr—ted into the wediterr—E sion is represented ˜y the first o™™urren™e of the ˜iv—lve

ne—n ƒe— through the ƒtr—it of qi˜r—lt—r from higher er™ti™— isl—ndi™— —t IHQFUH m from the ˜—se of the se™E

northern l—titudesD in ™onsequen™e of the ™lim—ti™ ™oolE tionD whi™h until PHIH h—s ˜een used to m—rk the €lioE

ing ˜eginning in the g—l—˜ri—n @q—rilli PHIIAF w—l—test— ™eneE€leisto™ene ˜ound—ry @€elosio 8 ‚—ffi IWURY ‚—ffi

8 —rleng— @IWVTA —nd ‚—ffi @IWVTA developed —n exE IWVTAY now—d—ys it m—rks the qel—si—nEg—l—˜ri—n

h—ustive dis™ussion on ˜ore—l guestsD their —rriv—l —nd ˜ound—ryF ‚—ffi @IWVTA d—ted the first —ppe—r—n™e of

their distri˜ution in the wediterr—ne—n fossil ™ommuE er™ti™— isl—ndi™— into the wediterr—ne—n ƒe— —t the top

nities —nd in the present se—sF of the ylduv—i m—gneti™ su˜™hron —t IFTU w—Y howeverD

er™ti™— isl—ndi™— is the most f—mous fqD ˜ut other this —ge h—s to ˜e ™orre™ted —sD —fter IWWSD — new —stroE

interesting ™lim—ti™ indi™—tors h—ve ˜een found in the nomi™—lly tuned time s™—le ˜eg—n to ˜e used to define

se™tionX wytilus edulis —nd €seud—mussium peslutr—e the m—gnetostr—tigr—phi™ ˜ound—ries @†—n gouvering

˜oth —ppe—ring str—tigr—phi™—lly —˜ove the eF isl—ndi™— IWWUAF „he new —ge for the topof the ylduv—i su˜™hron

first o™™urren™eF eF isl—ndi™— is found now—d—ys —t high —nd the ™orresponding first —ppe—r—n™e d—tum for eF

isl—ndi™— in the wediterr—ne—n ƒe— is thus IFUU w—F e™E l—titudes in the etl—nti™ y™e—n —long the emeri™—n

™ording to uukl— et —lF @IWUWA eF isl—ndi™— first —ppe—rs ™o—st —nd in iurope —long the ™o—sts of s™el—ndD qre—t

wollusk f—un— of the vower €leisto™ene erd— ‚iver se™tion @xorthern st—lyA TU

˜ore—l guestsD there is — gr—du—l de™re—se of w—rmEtemE frit—in —nd the ƒ™—ndin—vi—n peninsul— @h—hlgren et —lF

per—te f—un—sD from the pper €lio™ene to the widdle PHHHAY —™™ording to w—l—test— 8 —rleng— @IWVTA wytilus

€leisto™eneD whi™h h—s ˜een ™onsidered — ™onsequen™e edulis h—s — simil—r geogr—phi™ distri˜utionD where—s

of —n in™re—se in se—son—lity —nd — de™re—se in pre™ipit—E €seud—mussium peslutr—e is now widespre—d in the e—stE

tion —nd in temper—ture @€elosio 8 ‚—ffi IWUUY ‚—ffi ern p—rt of the etl—nti™ o™e—n from s™el—nd to woro™™o

IWVTY homini™i PHHIAF ™o—stsF roweverD their interpret—tion —s ˜ore—l guests is

„he —rriv—l of ˜ore—l guestsD the ™h—nge in the not —lw—ys str—ightforw—rd —nd in f—™tD only the ˜iv—lves

flor— —nd the dis—ppe—r—n™e of w—rmEtemper—te t—x— er™ti™— isl—ndi™— —nd €seud—mussium peslutr—e ™—n ˜e

thus indi™—te — progressive ™lim—te deterior—tion during ™onsidered good m—rkers in this reg—rdF

the i—rly €leisto™eneD whi™h w—s pro˜—˜ly ™—used ˜y — e™™ording to w—l—test— 8 —rleng— @IWVTAD wytiE

™omplex interpl—y of f—™tors su™h —s ™older se—w—ter lus edulis —ppe—rs in the wediterr—ne—n ƒe— in the g—E

temper—turesD in™re—se in se—son—lity —nd —ridityF rowE l—˜ri—nF roweverD it is often diffi™ult to distinguish it

everD the import—n™e —nd timing of e—™h of these envirE from wytilus g—lloprovin™i—lisD whi™h is — typi™—l wedE

onment—l v—ri—˜les in driving the o˜served ™h—nges reE iterr—ne—n form —nd thus not — ˜ore—l guestD —s they ˜oth

m—ins unsolvedF show gre—t v—ri—tion in shell sh—pe due to environment—l

™onditions @ƒeed IWWPAY they ™—n —lso ˜e hy˜ridized

@ƒki˜inski et —lF IWUVAF qosling @IWWPA —nd ƒeed @IWWPD

gon™lusions IWWSA o˜served th—t not — single morphologi™—l ™h—r—™E

teristi™ ™—n ˜e used to distinguish these wytilus spe™iesF

„he vower €leisto™ene p—rt of the erd— ‚iver por these re—sons it h—s ˜een very diffi™ultD if not imE

su™™ession h—s proved to ˜e —n interesting se™tion for possi˜leD to distinguish the two spe™ies in the erd— se™E

its ™ontinuity —nd its well preserved —nd very ri™h fossil tion —nd it is thus pro˜lem—ti™ to ™onsider the o™™urE

spe™imens whi™h —llow to ™onstr—in its —ge —nd infer its ren™e of this t—xon —s — ˜ore—l guestF

p—l—eo™lim—ti™ evolutionF

e™—ntho™—rdi— e™hin—t— does —lso o™™ur in the f—uE

„he system—ti™ study of the erd— f—un— led to

n—Y it w—s ™onsidered ˜y sever—l —uthors — fq @eFgF w—E

™on™lude th—tX

l—test— 8 —rleng— IWVTY ‚—ffi IWVTAD —ppe—ring in the

IF st is ™h—r—™terized ˜y — high ˜iodiversity with

g—l—˜ri—n in the wediterr—ne—n ƒe—F roweverD some

ISW t—x—Y ˜iv—lves —re the domin—nt t—xonD followed ˜y

—uthors @eFgF woneg—tti 8 ‚—ffi PHHIA reported th—t it

g—stropodsD ™or—lsD serpulidsD ˜r—™hiopodsD e™hinoidsD

w—s present in the wediterr—ne—n ƒe— —lre—dy sin™e the

—rthropods @eFgF ˜—rn—™lesAD ˜ryozo—ns —nd s™—phopodsF

€lio™eneF st is not ™le—r though if the —uthors here refer

PF „he l—st o™™urren™es of the ˜iv—lves eequipe™E

to eF e™hin—t— e™hin—t— or to eF e™hin—t— mu™ron—t—Yin

ten s™—˜rell—D qly™ymeris infl—t— —nd gh—m— pl—™entin—D

f—™t the first su˜spe™ies is —n etl—nti™ restri™ted t—xon

the first o™™urren™e of er™ti™— isl—ndi™— —nd the l—st o™E

@the ˜ore—l guest of w—l—test— 8 —rleng— IWVT —nd

™urren™es of the g—stropods „urritell— tri™—rin—t— plioE

‚—ffi IWVTAD where—s the l—tter is the typi™—l wediterr—E

re™ens —nd x—ss—rius prysm—ti™us suggest — l—te qel—si—nE

ne—n su˜spe™iesF xow—d—ys they —re ™onsidered synoE

g—l—˜ri—n —ge @i—rly €leisto™eneA for the su™™essionF

nyms —nd this m—y —dd pro˜lems to the interpret—tionF

QF „he o™™urren™e of ˜ore—l guests —long the se™E

v— €ern— 8 h9e˜r—mo @PHHWA g—ve — ™l—rifying dis™usE

tionD su™h —s er™ti™— isl—ndi™—D €seud—mussium peslutr—e

sion on this topi™ o˜serving th—t —lso e™—ntho™—rdi—

—nd possi˜ly wytilus edulisD suggests th—t — ™lim—ti™

e™hin—t— e™hin—t— w—s —lre—dy present in the €lio™ene

™h—nge o™™urred in the se™tion with — shift to ™older

—s shown in sever—l re™ords @€—ll— IWTTY g—v—llo 8 ‚eE

se—w—ter temper—turesF „he evolution of this ™lim—ti™

petto IWWPAF „husD —™™ording to v— €ern— 8 h9e˜r—mo

deterior—tion is ™omplexD ˜ut it prep—res the ground

@PHHWA e™—ntho™—rdi— e™hin—t— should not ˜e ™onsidered

for the onset of the ™ontinent—l gl—™i—tion of the widdle

—s — ˜ore—l guestY here we follow this interpret—tionF

—nd v—te €leisto™eneF

eside from the o™™urren™e of ˜ore—l guestsD the

—n—lysis of the ™omposition of the f—un— does not —llow

h—ving more pre™ise p—l—eo™lim—ti™ inform—tionY in f—™tD

ƒystem—ti™s —nd des™riptions

it is m—inly domin—ted ˜y eurytherm—l spe™ies h—ving —

™osmopolit—n distri˜utionD in™luding ˜ore—l to w—rm „he des™ription of the spe™ies ˜elonging to the

temper—te w—ter spe™iesY the stri™tly —r™ti™ or tropi™—l genus qly™ymeris h— gost—D IUUVD eequipe™ten pis™herD

t—x— —re inste—d —˜sent from the —sso™i—tionsF IVVT —nd er™ti™— ƒ™hum—™herD IVIU —re here presentedF

sn the wediterr—ne—n ˜—sinsD other ˜ioevents in „hese spe™ies —re very —˜und—nt —nd regul—rly present

—ddition to the —ppe—r—n™e of ˜ore—l guests testify to — —long —ll the erd— ‚iver m—rine su™™essionD for these

shift tow—rds ™old ™lim—te ™onditionsD —s for inst—n™e — re—sons they —re here des™ri˜edF ‡—iting for the pu˜liE

signifi™—nt ™h—nge in the pollen flor— @von— 8 fertoldi ™—tion of the revised volume of the ––„re—tise on snverE

IWUPAF elsoD in ™on™omit—n™e with the —ppe—r—n™e of the te˜r—te €—leontology99 the des™riptions ˜elow follow the

TV gripp— qF 8 ‚—ineri qF

™l—ssifi™—tion of gox et —lF @IWTWA —nd yliver 8 rolmes hes™riptionF v—rge to medium sized ˜i™onvex

shellD equiv—lveD slightly inequil—ter—l with ™ir™ul—r outE in fieler 8 wikkelsen @PHHTA for the er™oide—D of ‡—ller

lineY shell su˜st—n™e thi™kY v—lves slightly infl—ted with et —lF in fieler 8 wikkelsen @PHHTA for the €e™tinoide—

low —nd not very prominent orthogyr—te to slightly —nd of gox et —lF @IWTWA —nd fieler 8 wikkelsen @PHHTA

opisthogyr—te um˜oY dorsoEventr—l di—meter ne—rly for the er™ti™oide—F

equ—l to —nteroEposterior oneF

yrn—ment—tion of ˜oth v—lves ™onsisting of thin

r—di—l ™ostell—eD —nd irregul—r ™on™entri™ growth linesY

gl—ss fiv—lvi— vinn—eusD IUSV @fuon—nniD ITVIA

the l—tter p—ss to ™o—rse growth l—mell—e in the posteroE

ƒu˜™l—ss €teriomorphi— feurlenD IWRR

ventr—l p—rt of the v—lveY ventr—l intern—l m—rgin reguE

yrder er™oid— ƒtoli™zk—D IVUI l—rly flutedF

snterior of ˜oth v—lves with prionodont dentition ƒuperf—mily er™oide— v—m—r™kD IVHW

with teeth rel—tively shortD l—ter—lly † sh—pedD o˜solesE

p—mily qly™ymeridid—e h—llD IWHV

™ent in the ™entr—l p—rt —nd more evident l—ter—llyD deE

termining — division in two p—rt of the dentitionY lig—E

ment —re— highD n—rrowD tri—ngul—r —nd fl—t or slightly qenus qly™ymeris h— gost—D IUUV

™on™—veY extern—l lig—ment —mphideti™ duplivin™ul—r

„ypeEspe™iesX qly™ymeris gly™ymeris @vinn—eusD IUSVA

well preservedF from the ‚e™ent of the wediterr—ne—n ƒe—F

himy—ri—n —nisomy—ri—n —nd integrip—lli—te shellY

™ir™ul—r p—lli—l lineY —nterior —ddu™tor mus™le s™—r with ‚em—rksF gox et —lF @IWTWA identified two su˜geE

tri—ngul—r sh—pe —nd l—rger th—n the posterior su˜™ir™uE ner— for qly™ymeris h— gost—D IUUVX qly™ymeris sFsF —nd

l—r —nd su˜ov—l mus™le s™—rY ˜oth —ddu™tor mus™le s™—rs qly™ymerit—F rereD we follow ƒquires @PHIHA in —˜—nE

—re well impressed on — low myophori™ pl—tformY s™—rs doning su˜gener— definition for qly™ymeris —nd in ™onE

of r—di—lly dire™ted fi˜ers of mus™le m—ntle —long the sidering qly™ymerit— —s — distin™t genus @ƒquires PHIHAD

p—lli—l lineF —s its shell surf—™e h—s m—ny prominently r—ised r—di—l

himensionsF ƒee pigs Q —nd RF ri˜s with well defined n—rrow intersp—™es in ™ontr—st to

his™ussionF qly™ymeris gly™ymeris h—s — ™ir™ul—r the m—inly smooth surf—™e of qly™ymerisF

outline —nd poorly infl—ted v—lvesY for these ™h—r—™ters it qly™ymerisD —s other qly™ymeridid—eD h—s — deE

differs from qly™ymeris ˜im—™ul—t— @€oliD IUWSAD qly™yE ™eptive v—lve orient—tionD whi™h do not follow the ™l—sE

meris infl—t— @fro™™hiD IVIRA —nd qly™ymeris insu˜ri™— si™—l rulesX the l—rger —ddu™tor s™—r is the —nterior one

@fro™™hiD IVIRAF purthermoreD the lig—ment —re— is

—nd not the posterior —s usu—lly h—ppens in other ˜iv—lve

slightly different from the other spe™ies of qly™ymerisD groups @see xi™ol 8 tones IWVRAF

˜e™—use it is high ˜ut not l—ter—lly exp—ndedF

qly™ymeris gly™ymeris together with qly™ymeris

qly™ymeris gly™ymeris @vinn—eusD IUSVA

infl—t—D represents the l—rger spe™ies @pigF QA with — rG

€lF ID figs ID PY €lF PD figs ID Q v r—tio ne—rly equ—l I @pigF RAD whi™h —llow to distinE

guish it from the other qly™ymeris spe™iesF

IUSV er™— gly™ymeris vinn—eusD pF TWS

ƒtr—tigr—phi™ —nd geogr—phi™ o™™urren™eF qlyE

IWTT qly™ymeris gly™ymeris @vinn—eusA ± „e˜˜leD pF QQD plF PD

™ymeris gly™ymeris is known from the wio™ene to the figsF —E™

‚e™ent from the ‡estern etl—nti™ ™o—st to the wediterE IWUH qly™ymeris gly™ymeris @vinn—eusA ± fu™™heriD pF PST

IWUR qly™ymeris gly™ymeris @vinn—eusA ± €—renz—nD pF RUD plF SD r—ne—n ƒe— @w—r—sti 8 ‚—ffi IWVHAF

figF RH

IWVH qly™ymeris @qly™ymerisA gly™ymeris @vinn—eusA ± enfossi

8 fr—m˜ill—D pF SVD plF S @IAD figsF U—E™ qly™ymeris infl—t— @fro™™hiD IVIRA

IWVT qly™ymeris @qly™ymerisA —ffF qF @qFA gly™ymeris @vinn—eusA €lF ID figF QY €lF PD figs RD SY €lF QD figF I

± homeÁne™hD pF IQSD figsF RsEt @™um synFA

IWVT qly™ymeris gly™ymeris @vinn—eusA ± ‚iedlD pF QSWD plF IPP IVIR er™— infl—t— fro™™hiD pF RWRD plF IID figF U

IWVV qly™ymeris gly™ymeris @vinn—eusA ± qomezEel˜—D pF ITVD IVUU €e™tun™ulus infl—tus @fro™™hiA ± ƒeguenz—D pF PVPEPVQ

plF VQD figF II IVUWEIVVP €e™tun™ulus insu˜ri™us v—rF @fro™™hiA ± pont—nnesD pF

IWWQ qly™ymeris @qly™ymerisA gly™ymeris @vinn—eusA ± voz—no IUSD plF IID figF Q

pr—n™is™o et —lFD pF IUPD plF SD figsF IDPF IVWV €e™tun™ulus infl—tus @fro™™hiA ± elmer— 8 fofillD pF IPT

IVWV exine— infl—t— @fro™™hiA ± ƒ—™™o in fell—rdi 8 ƒ—™™oD pF QPD

w—teri—lX ƒix right v—lveX egqIREIPEPPEPRD egqPU˜isEUD plF †sssD figsF IEIH @™um synFA

egqPRQEIEPY twelve left v—lvesX egqIREIEPEQERESEIIEIQEPQD IWHU €e™tun™ulus infl—tus @fro™™hiA ± gerulli srelliD pF IIWD plF WD

egqPTERD egqPU˜isEVD egqQSEID egqPRIEIY one undetermined figF UY plF IHD figsF ID W

v—lveX egqPVEIF IWIRc €e™tun™ulus @exin—e—A insu˜ri™us @fro™™hiA ± fongoD pF

y™™urren™eX eqgIRD egqPTD egqPU˜isD egqPVD egqQSD RUS

IWQQ €e™tun™ulus @exine—A infl—tus @fro™™hiA ± foniD pF IRT egqPRID egqPRQD erd— ‚iver se™tionD g—stell9erqu—toD st—ly

wollusk f—un— of the vower €leisto™ene erd— ‚iver se™tion @xorthern st—lyA TW

pigF Q E reight —nd length v—lues of

spe™ies of qly™ymerisF

IWQS €e™tun™ulus @exine—A infl—tus @fro™™hiA ± gowper ‚eedD pF y™™urren™eX egqWD egqIHD egqIRD egqPRD egqPW˜isD

SIH erd— river se™tionD g—stell9erqu—toD st—lyF

IWQU €e™tun™ulus infl—tus @fro™™hiA ± hu˜ertret et —lFD pF IHH

IWQV €e™tun™ulus @exine—A infl—tus @fro™™hiA ± „—v—niD pF IRUD

hes™riptionF v—rge sizedD ˜i™onvex shell slightly plF SD figF P

inequiv—lve —nd inequil—ter—l with su˜™ir™ul—rD slightly

IWSH €e™tun™ulus @exine—A infl—tus @fro™™hiA ± pest—D pF WT

elong—ted outlineY shell su˜st—n™e thi™kY v—lves strongly IWSI qly™ymeris infl—t— @fro™™hiA ± sm˜esiD pF II

IWSI qly™ymeris @qly™ymerisA infl—tus @fro™™hiA ± ‚ossi ‚on™hE infl—ted with prominent orthogyr—te to slightly opisthoE

ettiD pF PHD figsF S—Ef

gyr—te um˜oY dorsoEventr—l di—meter l—rger th—n —nteroE

IWSP €e™tun™ulus infl—tus @fro™™hiA ± ve™ointreD pF TI

posterior oneF foth v—lves show — ridge in their posterE

IWSS €e™tun™ulus @exine—A infl—tus @fro™™hiA ± w—l—test— 8 xiE

ior p—rtD extending from the um˜o to the ventr—l m—rgin ™osi—D pF IUT

—nd forming —n o˜tuse —ngle in the shell outlineF IWSU qly™ymeris @qly™ymerisA infl—tus @fro™™hiA ± xi™osi—D pF

IUT yrn—ment—tion of ˜oth v—lves ™onsisting of very

IWSU qly™ymeris @qly™ymerisA infl—tus @fro™™hiA ± w—l—test—D pF

fineD regul—r —nd dense r—di—l ™ostell—e —nd in numerous

PP

™on™entri™ growth linesY the l—tter ˜e™ome growth l—E

IWSV qly™ymeris @qly™ymerisA infl—tus @fro™™hiA ± irun—l irenE

mell—e irregul—rly sp—™ed in the posteroEventr—l p—rt of tozD pF IRRD plF PPD figsF IWDPH

the v—lveY ventr—l interior m—rgin regul—rly flutedF IWTP qly™ymeris @qly™ymerisA infl—tus @fro™™hiA ± €—p—ni 8

€elosioD ppF QHTD QITD plF PD figF IP snterior of ˜oth v—lves with prionodont dentition

IWTQ qly™ymeris @qly™ymerisA infl—t— @fro™™hiA ± w—l—test—D pF

with teeth rel—tively shortD l—ter—lly † sh—pedD ˜e™oming

PQQD plF ˆsssD figF Q

o˜soles™ent or verti™—lly dire™ted in the middle p—rtY

IWTV qly™ymeris @qly™ymerisA infl—tus @fro™™hiA ± fon—donn—D

lig—ment —re— highD l—ter—lly exp—ndedD tri—ngul—r —nd pF PVT

IWUI qly™ymeris @qly™ymerisA infl—t— @fro™™hiA ± €es™e 8 ‚—E fl—t or slightly ™on™—veY extern—l lig—ment —mphideti™

pettiD pF IPRD plF ID figF Q duplivin™ul—r well preservedF

IWUP qly™ymeris @qly™ymerisA infl—t— @fro™™hiA ± g—prottiD pF

himy—ri—n —nisomy—ri—n —nd integrip—lli—te shellY

STD plF QD figF IW

™ir™ul—r p—lli—l lineY —nterior —ddu™tor mus™le s™—r with

IWUQ qly™ymeris @qly™ymerisA infl—tus @fro™™hiA ± w—r—stiD pF

tri—ngul—r sh—pe —nd slightly l—rger th—n the posterior IHH

IWUR qly™ymeris @qly™ymerisA infl—t— @fro™™hiA ± w—l—test—D pF su˜™ir™ul—r to su˜ov—l mus™le s™—rY ˜oth —ddu™tor musE

QHD plF PD figsF P—D˜

™le s™—rs —re well impressed on — myophori™ pl—tform

IWUT qly™ymeris infl—t— @fro™™hiA ± g—prottiD pF RD plF TD figF IW

@the posterior one is more elev—tedAY s™—rs of r—di—lly

IWVH qly™ymeris @qly™ymerisA infl—t— @fro™™hiA ± enfossi 8

dire™ted fi˜ers of mus™le m—ntle —long the p—lli—l lineF fr—m˜ill—D pF SVD plF †@sAD figsF V—D˜

€osterior ped—l retr—™tor s™—r with — sm—ll ™—l™ified pl—tE IWWQ qly™ymeris @qly™ymerisA infl—t— @fro™™hiA ± voz—no pr—nE

™is™o et —lFD pF IURD plF SD figsF SDT formF

himensionsF ƒee pigs Q —nd RF

w—teri—lX „hree —rti™ul—ted spe™imensX egqIHERD egqIREPSD

his™ussionF qly™ymeris infl—t— m—inly differs egqPREQQY twelve right v—lvesX egqWEQD egqIREIRD egqPW˜isEIESE

from qly™ymeris ˜im—™ul—t— @€oliD IUWSAD qly™ymeris UEWEPVEPWEQHEQQEQREQTY nine left v—lvesX egqWERD egqIRETD egqPWE

gly™ymeris @vinn—eusD IUSVA —nd qly™ymeris insu˜ri™— ˜isEPEQETEQIEQPEQSEQUF

UH gripp— qF 8 ‚—ineri qF

pigF R E reightGlength r—tio of spe™ies

of qly™ymerisF

@fro™™hiD IVIRA for its more elong—ted outline —nd for its

pe™uli—r ridge in the posterior p—rt of the v—lveD whi™h

€ve„i I

defines —n o˜tuse —ngle in the shell outlineF „he heightG

ell spe™imens —re xIY —A extern—l viewD ˜A intern—l viewD ex™ept when

length r—tio is —lw—ys higher th—n IF purthermore it

indi™—tedF

differs from qly™ymeris gly™ymeris @vinn—eusD IUSVA ˜eE I—E˜ E qly™ymeris gly™ymerisD left v—lve @egqIREQAY P—E˜ E qly™yE

meris gly™ymerisD right v—lve @egqIREPPAY Q—E˜E qly™ymeris infl—t—D ™—use it is strongly infl—ted —nd from qly™ymeris insuE

right —A —nd left ˜A v—lves of —n —rti™ul—ted spe™imen @egqPREQQAY R—E ˜ri™— @fro™™hiD IVIRA ˜e™—use it is l—rger —nd it h—s —

˜E qly™ymeris insu˜ri™—D right v—lve @egqWUEIHAF we—ker r—di—l orn—ment—tionF

qly™ymeris infl—t— together with qly™ymeris glyE

€ve„i P

™ymerisD represents the l—rger spe™ies @pigF QA with — rGv

ell spe™imens —re xIY —A extern—l viewD ˜A intern—l viewF

r—tio higher th—n I @pigF RAD whi™h —llow to distinguish it

I—E˜E qly™ymeris gly™ymerisD right v—lve @egqPU˜isEUAY P—E˜E qly™yE

from the other qly™ymeris spe™iesF

meris insu˜ri™—D left v—lveF xote the well preserved ™olor p—ttern

ƒtr—tigr—phi™ —nd geogr—phi™ o™™urren™eF qlyE @egqIWUESAY Q—E˜E qly™ymeris gly™ymerisD right v—lve @egqPRQEIAY

R—E˜E qly™ymeris infl—t—D right v—lve @egqPW˜isEUAY S—E˜E qly™ymeris ™ymeris infl—t— is — spe™ies known from the wio™ene to

infl—t—D right v—lve @egqPW˜isEPVAF the €leisto™ene @w—r—sti 8 ‚—ffi IWVHA of the wediterE

r—ne—n ƒe—F sts re™ent o™™urren™e @elt—˜— et —lF PHHTA is

€ve„i Q still de˜—tedF

ell spe™imens —re xIY —A extern—l viewD ˜A intern—l viewF

I—E˜E qly™ymeris infl—t—D left v—lve @egqPW˜isEQPAY P—E˜E qly™ymeris

qly™ymeris insu˜ri™— @fro™™hiD IVIRA insu˜ri™—D left v—lve @egqPHREQAY Q—E˜E qly™ymeris insu˜ri™—D right

v—lve @egqPTRERAY R—E˜E qly™ymeris insu˜ri™—D left v—lveF xote the €lF ID figF RY €lF PD figF PY €lF QD figs PEU

well preserved extern—l lig—ment @egqPSWEIAY S—E˜E qly™ymeris inE

su˜ri™—D left v—lve @egqWRETAY T—E˜E qly™ymeris insu˜ri™—D left v—lve IUSV er™— numm—ri— vinn—eusD pF TWS

@egqPHRERAY U—E˜E qly™ymeris insu˜ri™—D right v—lve @egqIWUERAF IVHS €e™tun™ulus ™or v—m—r™kD pF PIU

IVIR er™— insu˜ri™— fro™™hiD pF RWPD plF ˆsD figsF IH—D ˜

€ve„i R IVIW €e™tun™ulus viol—™es™ens v—m—r™kD pF SP

IVPS €e™tun™ulus insu˜ri™us @fro™™hiA ± forsonD pF IPR

ell spe™imens —re xIY —A extern—l viewD ˜A intern—l viewF

IVPT €e™tun™ulus insu˜ri™us @fro™™hiA ± ‚issoD pF QIV

I—E˜ E eequipe™ten oper™ul—risD right v—lve @egqIWUETAY P—E˜ E eeE

IVQI €e™tun™ulus insu˜ri™us @‚issoA ± fronnD pF IHV

quipe™ten oper™ul—risD right v—lve @egqPPPEIAY Q—E˜ E eequipe™ten

IVQU €e™tun™ulus insu˜ri™us @fro™™hiA ± qoldfussD pF ITI

oper™ul—risD right v—lve @egqWUEVAY R—E˜ E er™ti™— isl—ndi™—D right

IVTV €e™tun™ulus insu˜ri™us @fro™™hiA ± w—yerD pF RRERT

v—lve @egqPISEVAY S—E˜ E eequipe™ten oper™ul—risD right v—lve

IVUQ €e™tun™ulus viol—™es™ens @v—m—r™kA ± go™™oniD pF IPT

@egqIWVEPAY T—E˜ E eequipe™ten oper™ul—risD left v—lve @egqPQWEPAY

IVUV €e™tun™ulus insu˜ri™us @fro™™hiA ± €—ron—D pF WI

U—E˜ E eequipe™ten oper™ul—risD right v—lve @egqIHREPAF

IVUW €e™tun™ulus numm—rius @vinn—eusA ± teffreysD pF QP

IVVT €e™tun™ulus insu˜ri™us @fro™™hiA ± †erriD pF RRP

€ve„i S

IVWS €e™tun™ulus viol—™es™ens @v—m—r™kA ± erduiniD pF SH

IVWV €e™tun™ulus insu˜ri™us @fro™™hiA ± elmer— 8 fofillD pF IPT ell spe™imens —re xIY —A extern—l viewD ˜A intern—l viewF

IVWV exine— insu˜ri™— —nd v—rF @fro™™hiA ± ƒ—™™o in fell—rdi 8 I—E˜ E eequipe™ten oper™ul—risD left v—lve @egqIWRESAY P—E˜ E eequiE

pe™ten s™—˜rell—D left v—lve @egqREPAY Q—E˜ E eequipe™ten s™—˜rell—D ƒ—™™oD pF QQD plF VD figsF IIEPI

right v—lve @egqREWAY R—E˜ E eequipe™ten s™—˜rell—D right v—lve IWHU €e™tun™ulus @exine—A insu˜ri™us @fro™™hiA ± gerulli srelliD

@egqRESAY S—E˜ E er™ti™— isl—ndi™—D left v—lve @egqPSRETAY T—E˜ E pF IPID plF ˆsD figF l˜D PF

er™ti™— isl—ndi™—D left v—lve @egqPSQEPAF IWIS €e™tun™ulus insu˜ri™us @fro™™hiA ± h—lloniD pF RSQ

wollusk f—un— of the vower €leisto™ene erd— ‚iver se™tion @xorthern st—lyA UI €ve„i I

UP gripp— qF 8 ‚—ineri qF €ve„i P

wollusk f—un— of the vower €leisto™ene erd— ‚iver se™tion @xorthern st—lyA UQ €ve„i Q

UR gripp— qF 8 ‚—ineri qF €ve„i R

wollusk f—un— of the vower €leisto™ene erd— ‚iver se™tion @xorthern st—lyA US €ve„i S

UT gripp— qF 8 ‚—ineri qF

egqPHHD egqPHID egqPHPD egqPHQD egqPHRD egqPHSD IWRW— qly™ymeris insu˜ri™us @fro™™hiA ± ‚uggieriD pF PU

egqPHTD egqPIHD egqPIID egqPISD egqPSPD egqPSSD IWRW˜ qly™ymeris insu˜ri™us @fro™™hiA ± ‚uggieriD pF TW

egqPSTD egqPST˜isD egqPSUD egqPSVD egqPSWD egqPTHD IWSH €e™tun™ulus @exine—A insu˜ri™us @fro™™hiA ± pest—D pF WT

egqPTID egqPTI˜isD egqPTPD egqPTQD egqPTRD egqPTTD IWSI qly™ymeris @qly™ymerisA insu˜ri™us @fro™™hiA ± ‚ossi

egqPTUD erd— river se™tionD g—stell9erqu—toD st—lyF ‚on™hettiD pF PPD figsF T—Ef

IWSI qly™ymeris viol—™es™ens @v—m—r™kA ± sm˜esiD pF IPW

IWSP €e™tun™ulus viol—™es™ens @v—m—r™kA ± ve™ointreD pF TP

hes™riptionF wedium sizedD ˜i™onvex shellD

IWSU— qly™ymeris insu˜ri™us @fro™™hiA ± ‚uggieriD pF QI

ne—rly equiv—lveD slightly inequil—ter—l with tr—nsverseD

IWSU˜ €e™tun™ulus insu˜ri™us @fro™™hiA ± ‚uggieriD pF T

su˜qu—dr—te to su˜re™t—ngul—r outlineY shell su˜st—n™e IWTP qly™ymeris insu˜ri™us @fro™™hiA ± €—p—ni 8 €elosioD pF IT

not so thi™kY some spe™imens h—ve — rounded —nterior IWTR qly™ymeris @qly™ymerisA ™or @v—m—r™kA ± irun—l irentoz

8 yztemurD pF PTP m—rgin —nd — str—ight posterior oneY v—lves infl—ted with

IWTU qly™ymeris @qly™ymerisA ™or @v—m—r™kA ± gon—to et —lFD pF

orthogyr—te to slightly opisthogyr—te um˜oF

PTI

yrn—ment—tion of ˜oth v—lves ™onsisting of fineD

IWUI qly™ymeris @qly™ymerisA ™or @v—m—r™kA ± €es™e 8 ‚—pettiD

regul—r —nd dense r—di—l ™ostell—eD num˜ering IEQ in S pF IPPD plF ID figF P

mm —t the ventr—l m—rgin in —dult spe™imensY r—di—l IWUR qly™ymeris @qly™ymerisA viol—™es™ens @v—m—r™kA ± w—l—E

test—D pF QID plF PD figsF I—E™ orn—ment—tion more m—rked th—n ™on™entri™ oneY ™onE

IWUR qly™ymeris viol—™es™ens @v—m—r™kA ± €—renz—nD pF SHD plF TD

™entri™ —nd very thin growth linesD whi™h in some spe™iE

figF RQ

mens ˜e™ome more pronoun™ed in the ventr—l p—rt

IWVH qly™ymeris @qly™ymerisA insu˜ri™— @fro™™hiA ± w—r—sti 8

where they form ™o—rse l—mell—eY ventr—l interior m—rgin ‚—ffiD pF V

regul—rly flutedF sn some juvenile spe™imens —nd in the IWVP qly™ymeris @qly™ymerisA insu˜ri™— @fro™™hiA ± fenigni 8

gorselliD pF TUS juvenile region of the —dult spe™imens the violet ™olor

IWVP qly™ymeris @qly™ymerisA insu˜ri™us @fro™™hiA ± endreÂsD pF

orn—ment—tion is preserved with ™on™entri™ v—ri—tions

IIVD plF PD figsF IER

of tone —nd reddish mottlesF

IWVP qly™ymeris @sFsFA —ffF viol—™es™ens @v—m—r™kA ± w—rtinell 8

snterior of ˜oth v—lves with prionodont dentition homeÁne™hD pF QVP

with teeth rel—tively shortD l—ter—lly † sh—pedD ˜e™oming IWVQ qly™ymeris @qly™ymerisA —ffF insu˜ri™— @fro™™hiA ± homeÁE

ne™hD pF WQD plF QD figsF IEQD UD V o˜soles™ent or verti™—lly dire™ted medi—llyY lig—ment

IWVT qly™ymeris @qly™ymerisA —ffF qF @qFA insu˜ri™— @fro™™hiA ±

—re— l—ter—lly exp—nded ˜ut lowD tri—ngul—rD fl—t or

homeÁne™hD pF IQUD figsF RuEv

slightly ™on™—veY extern—l lig—ment —mphideti™ duplivinE

IWVV qly™ymeris insu˜ri™— @fro™™hiA ± qomezEel˜—D pF IUHD plF

™ul—rF VRD figF R

himy—ri—n —nisomy—ri—n —nd integrip—lli—te shellY IWVW qly™ymeris @qly™ymerisA insu˜ri™— @fro™™hiA ± endreÂsD pF

QQTD plF PD figsF IER @™um synFA ™ir™ul—r p—lli—l lineY —nterior —ddu™tor mus™le s™—r with

IWWP qly™ymeris insu˜ri™— @fro™™hiA ± gossign—ni et —lFD pF QID

tri—ngul—r sh—peD slightly l—rger th—n the posterior su˜E

figF PTV

™ir™ul—r to su˜ov—l mus™le s™—rY ˜oth —ddu™tor mus™le

IWWQ qly™ymeris @qly™ymerisA insu˜ri™— @fro™™hiA ± voz—no

s™—rs —re well impressed on — myophori™ pl—tformY s™—rs pr—n™is™o et —lFD plF ID figsF WEIH

of r—di—lly dire™ted fi˜ers of mus™le m—ntle —long the

w—teri—lX porty —rti™ul—ted spe™imenX egqPWEPD egqRTEIEPE p—lli—l lineF ƒu˜rhom˜oid—l posterior ped—l retr—™tor

QD egqRWEPEQD egqWIEQEIHD egqWREIQY egqWU˜isEID egqIWUEIEPD

s™—rF

egqIWVESETD egqPHHETEUD egqPHPEID egqPHREIEPD egqPIHEID

himensionsF ƒee pigs Q —nd RF egqPSPEPD egqPSTEIEPEQERD egqPST˜isEIEPD egqPSVEIEPD

his™ussionF qly™ymeris insu˜ri™— h—s — gre—t inE egqPSWERD egqPTIEIEPD egqPTI˜isEIEPEQD egqPTPEIEPEQD

tr—spe™ifi™ v—ri—˜ilityY —™™ording to endreÂs @IWVWA two egqPTQEIERY sixtyEthree right v—lvesX egqPW˜isEREVEIHEIQD egqQHE

SD egqQIEID egqRPEPD egqRP˜isEPD egqRQEID egqRSEPD egqTHE forms ™—n ˜e distinguishX the firstD ™—lled ––insu˜ri™—99D

ID egqTTEIEPESD egqVWEIEPEQD egqWHEQD egqWIEPESD egqWPEIEPD

™orresponds to th—t des™ri˜ed ˜y fro™™hi @IVIRAD

egqWQEQERESETEUD egqWREIERD egqWSEIEPD egqWTEQERD egqWUEPE

where—s the se™ondD ™—lled ––viol—™es™ens99D ™orresponds

QESEIHD egqWVEIEPD egqWWEID egqIWUERD egqIWVEQD egqPHHEISY

to th—t des™ri˜ed ˜y v—m—r™k @IVIWAY these two forms egqPHIERD egqPHREUY egqPISERD egqPSPEIETD egqPSSEID

differs m—inly for the outlineD more qu—dr—ngul—rD regE egqPSUEID egqPSVEQERD egqPSWEPD egqPTHEPD egqPTPERESD

egqPTQEPD egqPTREIEPERESD egqPTUEIEPY sixtyEfour left v—lvesX ul—r —nd equil—ter—l in the ––viol—™es™ens99 formF qly™yE

egqWEPD egqIQEPD egqPW˜isEIPEIREIUEIWD egqRSEID egqRWEID meris viol—™es™ens —nd qly™ymeris insu˜ri™— —re now

egqSQEITD egqSWEID egqTTEQERETD egqTVEID egqVQEID

™onsidered synonymY qF viol—™es™ens is usu—lly used to

egqWHEIEPEREIPD egqWIEIERD egqWPEQD egqWQEIEPEIQD egqWRE

identify the ‚e™ent spe™imensD where—s qF insu˜ri™— is

PEQESETD egqWTEIEPD egqWUEIEREWEIIEIPD egqWVEQD egqIWUESEIID

for fossil onesF egqIWWEIEPD egqPHQEIEPEQD egqPHREQERESETD egqPHSEIEPEQD

egqPHTEID egqPIIEID egqPSPEQERD egqPSUEPD egqPSWEIEQESD e™™ording to sever—l —uthors @from ƒ—™™o in felE

egqPTHEID egqPTQESD egqPTREQD egqPTTEID egqPTUEQF l—rdi 8 ƒ—™™o IVWV to ƒirn— IWUVAD in —ddition to qlyE

y™™urren™eX egqWD egqIQD egqPWD egqPW˜isD egqQHD

™ymeris viol—™es™ens @v—m—r™kD IVIWAD other synonyms

egqQID egqRPD egqRP˜isD egqRQD egqRSD egqRTD egqRWD

of qF insu˜ri™— —re qly™ymeris ™or @v—m—r™kD IVHSA —nd egqSQD egqSWD egqTHD egqTTD egqTVD egqVQD egqVWD

qly™ymeris numm—ri— @vinn—eusD IUSVAF qly™ymeris egqWHD egqWID egqWPD egqWQD egqWRD egqWSD egqWTD

numm—ri— is ™onsidered — nomen o˜litum ˜y ƒirn— egqWUD egqWU˜isD egqWVD egqWWD egqIWUD egqIWVD egqIWWD

wollusk f—un— of the vower €leisto™ene erd— ‚iver se™tion @xorthern st—lyA UU

@IWUVAD —lthough ‡o‚wƒ ™onsiders this spe™ifi™ n—me —s himensionsF ƒee pigs Q —nd RF

his™ussionF „hese spe™imens h—ve intermedi—te the v—lid oneF qly™ymeris ™or h—s ˜een inv—lid—ted ˜eE

™h—r—™ters —mong the —˜ove des™ri˜ed spe™iesX they h—ve ™—use of the in—dequ—te des™ription given for it ˜y v—E

— rGv r—tio higher th—n I —s qly™ymeris infl—t— @pigF RAD m—r™k @IVHSAF por these re—sonsD we h—ve determined

— su˜™ir™ul—r outline —s qly™ymeris gly™ymeris —nd — the spe™imens under ex—m —s qly™ymeris insu˜ri™—

thi™k shell with strong teeth —s qly™ymeris ˜im—™ul—t— @fro™™hiD IVIRAF

@€oliD IUWSAF et the s—me time they differ from spe™iE sn gener—lD qly™ymeris insu˜ri™— h—s — medium

mens of qly™ymeris infl—t— ˜e™—use they l—™k the ™h—rE sized shellD — su˜qu—dr—te to su˜re™t—ngul—r outlineD —

—™teristi™ —ngle in the —nterior v—lve outline —nd from pronoun™ed r—di—l orn—ment—tion —nd — low lig—ment

qly™ymeris gly™ymeris ˜e™—use they —re more infl—ted —re—Y for these ™h—r—™ters it differs from qly™ymeris inE

—nd h—ve — more prominent um˜oF qly™ymeris ˜im—™uE fl—t— @fro™™hiD IVIRAD from qly™ymeris ˜im—™ul—t— @€oliD

l—t— is the most simil—r spe™ies even if the erd— spe™iE IUWSA —nd from qly™ymeris gly™ymeris @vinn—eusD IUSVAF

mens do not —tt—in the l—rge size typi™—l of the repreE vooking —t the gr—ph in figF Q qly™ymeris insu˜riE

sent—tives of this spe™iesY furthermoreD they —re not well ™— is the spe™ies with the sm—llest size ˜ut it —lso repreE

preserved so — spe™ifi™ —ssignment is diffi™ultF sents the spe™ies with the higher num˜er of spe™imensY

qly™ymeris spF indF is simil—r in size to qF infl—t— —mong these spe™imens juvenile forms m—y o™™urD

—nd qF gly™ymeris @see pigF QAD presuming the —ffinity whi™h thus lower the me—surementsY despite th—tD the

with one of these two spe™iesY howeverD —s o˜served in m—jority of the spe™imens rem—ins of sm—ller size th—n

the system—ti™ des™riptionD the extern—l —nd intern—l other qly™ymeris spe™iesF qF insu˜ri™— h—s — rGv r—tio

™h—r—™ters of qly™ymeris spF indF —re different from qF sm—ller th—n ID whi™h —llow to distinguish it from other

infl—t— —nd qF gly™ymerisD promoting the f—™t th—t it is — qly™ymeris spe™ies @pigF RAF

distin™t spe™iesD —lthough the preserv—tion ™onditions do

ƒtr—tigr—phi™ —nd geogr—phi™ o™™urren™eF qlyE

not —llow — more —™™ur—te —ssignmentF

™ymeris insu˜ri™— is known from the i—rly wio™ene to

the ‚e™ent from the etl—nti™ y™e—n to the wediterr—E

ne—n ƒe— @w—r—sti 8 ‚—ffi IWVHAF

yrder €e™tinoid— ed—ms 8 ed—msD IVSU

ƒuperf—mily €e™tinoide— ‚—finesqueD IVIS

qly™ymeris spF indF

p—mily €e™tinid—e ‚—finesqueD IVIS

w—teri—lX „wo right v—lvesX egqIREUD egqPRERY one left ƒu˜f—mily eequipe™tinin—e xordsie™kD IWTW

v—lveX egqUTEPSF

y™™urren™eX egqIRD egqPRD egqUTD erd— river se™tionD

qenus eequipe™ten pis™herD IVVT g—stell9erqu—toD st—ly

„ypeEspe™iesX ystre— oper™ul—ris vinn—eusD IUSV from the ‚e™ent

of pr—n™eF

hes™riptionF v—rge sizedD ˜i™onvex shellD equivE

—lveD equil—ter—l with su˜™ir™ul—r to su˜qu—dr—te outlineY

‚em—rksF ƒimil—r to ghl—mys ‚odingD IUWVD ˜ut shell su˜st—n™e very thi™kY v—lves infl—ted with orthogyE

it differs ˜y its more rounded outlineD ne—rly equ—l —uriE r—te to slightly opisthogyr—te um˜oF

™lesD sh—llower ˜yss—l not™hD —nd fewerD usu—lly not ˜iE

yrn—ment—tion ™onsisting of very fine —nd reguE

fur™—ting r—di—l ri˜sF l—rly —rr—nged r—di—l ™ostell—e —nd ™on™entri™ growth

linesY the l—tter in the ventr—l p—rt of the v—lve ˜e™ome

eequipe™ten oper™ul—ris @vinn—eusD IUSVA dense —nd ™o—rse growth l—mell—eY this produ™es — ™le—r

€lF RD figs IEQD SEUY €lF SD figF I division of the v—lve in two p—rtsX IA the juvenileE—dult

dorsoEmedi—n p—rt ™h—r—™terized ˜y thin growth lines

IUSV ystre— oper™ul—ris vinn—eusD pF TWV

—nd PA the oldest ventr—l p—rt with ™o—rse growth l—melE

IVUQ €e™ten oper™ul—ris @vinn—eusA ± go™™oniD pF QVS

l—eY ventr—l interior m—rgin regul—rly flutedF IVVR €e™ten oper™ul—ris @vinn—eusA ± weliD pFIP

IVWV €e™ten oper™ul—ris @vinn—eusA ± elmer— 8 fofillD pF IIP snterior of v—lve with prionodont dentition with

IWHP €e™ten oper™ul—ris @vinn—eusA ± he ƒtef—niD pF S ™o—rse teethD ™urvedD l—™king medi—llyY lig—ment —re—

IWHT ghl—mys @eequipe™tenA oper™ul—ris @vinn—eusA ± goliniD

highD l—ter—lly exp—ndedD tri—ngul—r —nd fl—tY extern—l

pF ITQ

lig—ment —mphideti™ duplivin™ul—rF

IWHU ghl—mys @eequipe™tenA s™—˜rell— @v—m—r™kA ± gerulli srE

himy—ri—n —nisomy—ri—n —nd integrip—lli—te shellY elliD pF WPD plF SD figsF IUD IV

IWIH eequipe™ten oper™ul—ris @vinn—eusA ± ƒ™h—fferD pF QTD plF ITD ™ir™ul—r p—lli—l lineY —nterior —ddu™tor mus™le s™—r with

figsF IIEIS tri—ngul—r sh—pe —nd l—rger th—n the posterior su˜™ir™uE

IWTP ghl—mys oper™ul—ris @vinn—eusA ± foni 8 ƒ—™™hi †i—lliD pF

l—r oneY ˜oth —ddu™tor mus™le s™—rs —re well impressed

IHWD plF IPD figsF RD TY plF IQD figF IUY plF IRD figF IH

on — myophori™ pl—tformY s™—rs of r—di—lly dire™ted fiE IWTP ghl—mys @eequipe™tenA oper™ul—ris @vinn—eusA ± €—p—ni 8

€elosioD pF IS ˜ers of mus™le m—ntle —long the p—lli—l lineF

UV gripp— qF 8 ‚—ineri qF

IWTS vyrope™ten @eequipe™tenA oper™ul—ris @vinn—eusA ± qli˜ert y™™urren™eX egqPD egqRD egqTD egqWD egqIID egqIPD

egqIQD egqIRD egqPRD egqPSD egqPTD egqPUD egqPU˜isD 8 †—n de €oelD pF PT

egqPW˜isD egqQHD egqQID egqQPD egqQQD egqQRD egqQUD IWTT ghl—mys @eequipe™tenA oper™ul—ris @vinn—eusA ± „e˜˜leD pF

egqRID egqRI˜isD egqRPD egqRP˜isD egqRQD egqRRD egqSHD THD plF SD figF ˜Dd

egqSID egqSI˜isD egqSPD egqSQD egqSRD egqSSD egqSTD IWTV eequipe™ten @eequipe™tenA oper™ul—ris @vinn—eusA ± fon—E

egqSUD egqSWD egqTHD egqTID egqTTD egqTVD egqUHD donn—D pF PVP

egqUTD egqUUD egqVHD egqVID egqVPD egqVQD egqVRD IWTW eequipe™ten oper™ul—ris @vinn—eusA ± ‡—llerD plF ID figsF SD

egqVSD egqVWD egqWHD egqWID egqWPD egqWQD egqWRD UEIH

egqWUY egqIHHD egqIHID egqIHPD egqIHQD egqIHRD IWTW eequipe™ten oper™ul—ris @vinn—eusA ± xordsie™kD pF RWD plF

egqIHSD egqIITD egqIQPD egqIQQD egqIWRD egqIWSD †ssD figF QPFHH

egqIWTD egqIWUD egqIWVD egqIWWD egqPHHD egqPHH˜isD IWUH vyrope™ten @eequipe™tenA oper™ul—ris @vinn—eusA ± fu™E

egqPHID egqPHPD egqPHUD egqPHVD egqPHWD egqPIPD ™heriD pF PST

egqPIQD egqPIQ˜isD egqPISD egqPIUD egqPPHD egqPPPD

IWUH ghl—mys @eequipe™tenA oper™ul—ris @vinn—eusA ± ‚—ffiD pF

egqPPQD egqPQSD egqPQTD egqPQUD egqPQVD egqPQWD

IIH

egqPRHD egqPRQD egqPSPD egqPSQD egqPSSD erd— river se™tionD

IWUI eequipe™ten @eequipe™tenA oper™ul—ris @vinn—eusA ± €es™e

g—stell9erqu—toD st—ly

8 ‚—pettiD pF IPTD plF PD figF P @™um synFA

IWUP ghl—mys @eequipe™tenA oper™ul—ris @vinn—eusA ± g—prottiD

pF SVD plF ID figF S hes™riptionF wedium sized ˜i™onvex shellD inE

IWUR ghl—mys @eequipe™tenA oper™ul—ris @vinn—eusA ± w—l—test—D equiv—lve with orthogyr—teD ™onvex um˜oY ™ir™ul—r outE

pF RSD plF QD figsF Q—D˜

line with heightGlength r—tio ne—rly equ—l to IF ‚ight

IWUT ghl—mys oper™ul—ris @vinn—eusA ± g—prottiD plF RD figF S

v—lve equil—ter—l with slightly ™urved ™—rdin—l m—rginY

IWVT ghl—mys @eequipe™tenA oper™ul—ris @vinn—eusA ± p—r—oneD

—nterior —nd posterior —uri™les very different in size —nd pF RPD figsF UEIR

sh—peX the —nterior one is l—rgeD su˜re™t—ngul—r with — PHHW eequipe™ten oper™ul—ris @vinn—eusA ± timenez et —lFD pF UD

figsF Q™Ee deep˜yss—l not™hY the posteriorone is sm—ller —nd with

— tri—ngul—r outlineF veft v—lve slightly inequil—ter—lD

w—teri—lX „wo —rti™ul—ted spe™imensX egqIQPERD egqIQQEIY

more ™onvex th—n the right v—lveD with the dors—l region

PTI right v—lvesX egqPEPEQERD egqREIHEIQD egqTESD egqWETD

representing the gre—t ™onvexityY posterior p—rt more egqIIEID egqIPESD egqIQEQEVD egqIREISD egqPREUEIRD

developed th—n the —nterior oneY ™—rdin—l m—rgin egqPSEPEQEUEVEIREIVD egqPTEIEPEQETEVD egqPUETD egqPU˜isEQE

str—ightY —nterior —nd posterior —uri™les ne—rly simil—r SETD egqQHEIEUEVD egqQPEIEPD egqQREQD egqQUEVD egqRIEIEQE

RESETEUEWEIHEIPEIQEISEITEPREPSEPTEPUEPWD egqRI˜isEPEQERESETD in sizeD the —nterior is slightly l—rger th—n the posterior

egqRPEIEQERESETEUEVEWEIHEIQEIREIVEIWEPHEPIEPPEPQD egqRP˜isEIESE

one —nd h—s — sh—llow ˜yss—l not™hF

TEUEVEIHEIIEIREITEIUEPHEPQD egqRQEQD egqRREIEPD egqSI˜isEIEPE

yrn—ment—tion of ˜oth v—lves ™onsisting of ITEPI

QERETEUEVEIHD egqSPEIfD egqSQEIEQETEUEVEWEIHEIIEIPEIRD egqSRE

r—di—l ™ost—eD ™o—rser in some spe™imensD fl—tter in

IERESD egqSSEID egqSTESD egqSUEPEQERD egqSWEPEQESETEVD

othersY ™ost—e su˜qu—dr—ngul—r to su˜rounded in se™E egqTHEPEQESD egqTTEIPEIQD egqTVEPD egqUHEIEQD egqUTERESE

TEUEVEWEIHEIIEIPEIQEIVEPHEPID egqUUEID egqVHEIEPEQD egqVIEIE tionD ne—rly of the s—me width of the intersp—™esY some

PEQD egqVPEPD egqVQEPEQD egqVREIEPEQERESETEUD egqVSEID spe™imens show — se™ond—ry orn—ment—tion on the ™ostE

egqVWETEVD egqWHETEIHD egqWIEIEPEQEVDD egqWPEREVD egqWQEVE

—eD ™onsisting in IEP thinner ™ost—eY growth lines thin

IHEIID egqWREIHD egqWUEUEVY egqIHHEIEPERD egqIHIEIEPD

—nd denseD more evident in the ™ost—l intersp—™esY in egqIHPEQD egqIHQEIEPEQD egqIHREIEPEQD egqIHSEID egqIITEID

some spe™imens growth lines —re ™o—rser —nd imp—rt — egqIQPEIEQD egqIWREPERETEUD egqIWSEIEQESEUD egqIWUETEUEWEIHD

l—mellose p—tternF euri™les orn—mented ˜y thin ™ost—e egqIWVEPEQERD egqIWWEQD egqPHHEIEQERESD egqPHH˜isEIEPD

egqPHIEIEPEQD egqPHUEIEQERD egqPHVEPEQERD egqPISEIEPEQD —nd growth linesY —nterior —uri™le of the right v—lve orE

egqPIUEIEPETD egqPPHERESD egqPPPEIEPEQETD egqPPQEPEQERESD n—mented ˜y ™o—rser ™ost—e —nd ˜y very dense growth

egqPQSEPEQESD egqPQTEIEQERETD egqPQUEIERD egqPQVEIEPD

lines in — tri—ngul—r zone in ™orresponden™e of the ˜yE egqPQWEIEQERD egqPRHEIEPEQD egqPRQERD egqPSPESD egqPSQEID

ss—l not™hY ™tenolium presentF †—lve interior refle™ts the egqPSSEPY ISH left v—lvesX egqTETD egqIPEID egqIQEREIHD

extern—l orn—ment—tionF egqPSERESETEIQEPHD egqPTESEUD egqPUEUD egqPU˜isEIEPERD

snterior of ˜oth v—lves with — tri—ngul—r resilium egqPW˜isEITEIVD egqQHEPD egqQIEPD egqQQEIEPD egqQREIEPD

egqQUEVD egqRIEPEVEIIEIREPPEPQEQID egqRI˜isEID egqRPEIIEISE pitF wonomy—ri—n shellD well m—rked mus™le s™—rY the

ITEIUD egqRP˜isEQEREWEIPEIQEISD egqRREQD egqSHEID egqSIEIEPE

rounded —nd l—rger mus™le s™—r represents the —tt—™hE

QERD egqSI˜isESEWEIIEIPD egqSPEIeD egqSQESEIQEISD egqSSEPEQD

ment of the stri—ted mus™leD —nd the ˜e—nlike mus™le egqSTEIEPEREIHD egqSUEID egqSWEREUD egqTHERETD egqTIEID

s™—r represents the —tt—™hment of the smooth mus™leF egqTTEUEVEWEIHD egqUHEPD egqUTEIREISEITEIUEIVD egqVPEIESD

himensionsF ƒee pigF SF egqVWERESEUEWEIHD egqWHEUEVEIIEIREPHD egqWIETEUEWD egqWPESE

his™ussionF eequipe™ten oper™ul—ris is — very TEUD egqWQEWD egqWREUEVEWD egqWUETD egqIHIEQERD egqIHPEIEPD

egqIHQERESETD egqIHRERESD egqIQPEPD egqIQQEPEQERD egqIWRE ™ommon spe™ies in the erd— ‚iver su™™essionF e™™ordE

IEQESD egqIWSEPERETD egqIWTEID egqIWUEVEIID egqIWVEIEUD

ing to timenez et —lF @PHHWA this spe™ies is simil—r to eF

egqIWWERESD egqPHHEPD egqPHPEPD egqPHUEPD egqPHVEID

s™—˜rell— @v—m—r™kD IVIWAY they differ —s eF oper™ul—ris egqPHWEIEPD egqPIPEID egqPIQEID egqPIQ˜isEID egqPIUEQD

h—s — less ™urved um˜o —nd — ™ir™ul—r —nd more regul—r egqPPPERD egqPPQEID egqPQSEIERD egqPQTEPESETD egqPQUEPEQD

outline th—n eF s™—˜rell—D whi™h is very —symmetri™ @the egqPQVEQERESD egqPQWEPD egqPRQEQY yne undetermined v—lveX

—nteroEposterior di—meter is l—rger th—n the dorsoEvenE egqIWTEPF

wollusk f—un— of the vower €leisto™ene erd— ‚iver se™tion @xorthern st—lyA UW

pigF S E reight —nd length v—lues of

spe™ies of eequipe™tenF

IWSH ghl—mys @eequipe™tenA s™—˜rell— @v—m—r™kA ± pest—D pF WR tr—l oneAY furthermore eF s™—˜rell— h—s ™o—rser —nd fewer

IWSP ghl—mys s™—˜rell— @v—m—r™kA ± ve™ointreD pF SS

™ost—e —nd more ™onvex v—lvesF €ro˜—˜ly these two speE

IWSS ghl—mys @eequipe™tenA s™—˜rell— @v—m—r™kA ± woroniD pF

™ies h—d — ™ommon origin in the wio™ene —s in this time

IQH

interv—l they —re diffi™ult to distinguish one from the

IWSV ghl—mys s™—˜rell— @v—m—r™kA ± irun—l irentozD pF IRWD plF

other @w—l—test—D IWURAF PRD figsF QD R

eequipe™ten —ngelonii @he ƒtef—ni 8 €—nt—nelliD IWSW ghl—mys @eequipe™tenA seniensis @v—m—r™kA ± endersonD pF

WVD plF ˆs†D figsF V—D ˜ IVUVA is —lso simil—r to eF oper™ul—risD m—inly in the

IWTQ ghl—mys s™—˜rell— @v—m—r™kA ± †enzo 8 €elosioD pF IRVD juvenile st—gesF sn —dulthoodD represent—tives of the

plF RUD figF P

two spe™ies differ sin™e the shell of eF —ngelonii is thi™kE

IWTT eequipe™ten @eequipe™tenA seniensis @v—m—r™kA ± gomp—gE

erD the —uri™les —re less developedD the r—di—l ri˜s h—ve —

noniD pF ITVD plF ID figsF P—D˜

very ™h—r—™teristi™ tri—ngul—r se™tionD —nd it h—s spinesF

IWUH ghl—mys @eequipe™tenA s™—˜rell— @v—m—r™kA ± ‚—ffiD pF IIRD

fetween the two spe™ies of eequipe™ten @pigF SA plF PUD figsF IDPY plF PWD figsF P—D˜ @™um synFA

IWUP ghl—mys @ergope™tenA seniensis @v—m—r™kA ± g—prottiD pF we do not o˜serve p—rti™ul—r trend —s in qly™ymeris

SWD plF ID figF Q

@pigF QAF sn f—™t eF s™—˜rell— —nd eF oper™ul—ris show

IWUR ghl—mys @eequipe™tenA seniensis @v—m—r™kA ± w—l—test—D pF

simil—r size —nd in ˜oth ™—ses height —nd length —re

RUD plF QD figsF T—D˜

ne—rly equ—l @rGv r—tio equ—l IA indi™—ting — su˜™ir™ul—r

IWUT ghl—mys @eequipe™tenA seniensis @v—m—r™kA ± g—prottiD plF

outlineF RD figF Q

ƒtr—tigr—phi™ —nd geogr—phi™ o™™urren™eF eeE IWVT ghl—mys @eequipe™tenA s™—˜rell— @v—m—r™kA ± p—r—oneD pF

IRD figF IS quipe™ten oper™ul—ris is known from the wio™ene to

PHHW eequipe™ten s™—˜rell— @v—m—r™kA ± timenez et —lFD pF UD figsF the ‚e™ent of the etl—nti™ y™e—n —nd the wediterr—ne—n

QfEiD R—D˜

ƒe— @w—r—sti 8 ‚—ffi IWVHAF

w—teri—lX „wentyEtwo right v—lvesX egqIEID egqPESD egqQE

RESETEUD egqREQESETEUEVEWEIIEISD egqSEQERD egqTEPD egqVEPD

eequipe™ten s™—˜rell— @v—m—r™kD IVIWA

egqIPERD egqIQEIID egqQQERDc egqTTEIIY thirteen left v—lvesX

€lF S figs PER

egqIEPD egqPETD egqQEQD egqREPEREIPD egqSEPD egqTEREVD

egqIPEPEQETD egqIQESF

IVIW €e™ten seniensis v—m—r™kD pFIVP

y™™urren™eX egqID egqPD egqQD egqRD egqSD egqTD

IVIW €e™ten s™—˜rellus v—m—r™kD pF IVQ

egqVD egqIPD egqIQD egqQQDc egqTTD erd— ‚iver se™tionD g—sE

IVUW €e™ten s™—˜rellus @v—m—r™kA ± weliD pF SD T

tell9erqu—toD st—lyF

IVUWEIVVP €e™ten s™—˜rellus @v—m—r™kA ± pont—nnesD pF IVUD plF

IPD figsF PD Q

IVWV €e™ten s™—˜rellus @v—m—r™kA ± elmer— 8 fofillD pF III hes™riptionF wedium sized ˜i™onvex shellD inE

IWHP €e™ten s™—˜rellus @v—m—r™kA ± he ƒtef—niD pF S

equiv—lve —nd inequil—ter—lY slightly tr—nsverse su˜ov—l

IWHT eequipe™ten s™—˜rellus @v—m—r™kA ± goliniD pF ITU

outline —nd orthogyr—teD ™onvex um˜oY ™—rdin—l m—rgin

IWIQ ghl—mys s™—˜rellus @v—m—r™kA ± qignouxD pF QTW

str—ightY dorsoEventr—l di—meter sm—ller th—n —nteroE IWIR ghl—mys @eequipe™tenA s™—˜rellus @v—m—r™kA ± fongoD pF

posterior oneY posterior region more developed th—n RUP

IWQQ ghl—mys @eequipe™tenA seniensis @v—m—r™kA ± †enzoD pF VID the —nterior oneF ‚ight —nd left v—lves —uri™les simil—r

plF UD figsF IVD PIY pF VP

in sh—pe @tri—ngul—rA —nd sizeD with the —nterior one

IWQT €e™ten @eequipe™tenA seniensis @v—m—r™kA ± pried˜ergD plF

l—rger th—n the posterior oneY the —nterior —uri™le h—s — ssD figsF QUEQ

sm—ll ˜yss—l not™h in ˜oth v—lvesF IWRH ghl—mys s™—˜rell— @v—m—r™kA ± ‚ogerD pF QQPD QRP

VH gripp— qF 8 ‚—ineri qF

yrn—ment—tion of ˜oth v—lves ™onsisting of IQEIU ‚em—rksF er™ti™— differs from €ele™yor— h—llD

IWHPD ˜e™—use the former h—s — thi™ker shell —nd l—™ks ™o—rse r—di—l ™ost—eD thinner th—n the ™ost—l intersp—™esD

the p—lli—l sinusY it differs from g—llist— €oliD IUWI —s the —nd of thin —nd dense ™on™entri™ growth linesY ™ost—e

l—tter is glossyD usu—lly without s™ulpture —nd with the —re qu—dr—ngul—r in se™tion ne—r the um˜oD ˜e™oming

p—lli—l sinusF rounded in the ventr—l regionY the ™h—nge in the outline

of the se™tion usu—lly o™™urs in ™orresponden™e of —

er™ti™— isl—ndi™— @vinn—eusD IUTUA growth h—lt where the shell forms — sh—rpstepY—fter

€lF R figF RY €lF SD figs SET this tr—nsition r—di—l ™ost—e —nd intersp—™es ne—r the

ventr—l m—rgin ˜e™ome orn—mented ˜y thinner ri˜sX QE

IUTU †enus isl—ndi™— vinn—eusD pF IIQI

T on the ™ost—eD PEQ in the ™ost—l intersp—™esY in some

IUUV †enus ˜u™™—rdium fornD pF TQD plF RD figF II

spe™imensD m—inly —dult onesD there —re IEQ growth h—lts

IUUV €e™tun™ulus ™r—ssus h— gost—D pF IVQD plF IRD figF S

forming IEQ sh—rp stepsD ˜ut only in ™orresponden™e of IVIU er™ti™— vulg—ris ƒ™hum—™herD pF IRSD plF IQD figF Q

IVTR gyprin— isl—ndi™— v—rF ™r—ssior teffreysD pF QHS the first h—lt the ™h—nge in se™tion —nd in orn—ment—tion

IVWV gyprin— isl—ndi™— E ƒ—™™o in fell—rdi 8 ƒ—™™oD †F PVD pF IHD of the ™ost—e t—kes pl—™eF euri™les orn—mented ˜y thin

plF ssD figsF IEP @™um synFA

™ost—eD more numerous in the posterior oneF †—lve interE

PHHI er™ti™— isl—ndi™— qof—s et —lF in gostello et —lFD pF PHIH

ior refle™ts the extern—l orn—ment—tionF

snterior of ˜oth v—lves with — tri—ngul—r resilium w—teri—lX pour —rti™ul—ted spe™imensX egqPSQEPD egqPSREIE

RETY thirtyEone ventr—l v—lvesX egqUUEID egqVTERESD egqPHHEWEIHE pitF wonomy—ri—n shell with not well impressedD

IPEIQD egqPHPEQD egqPIQ˜isEQD egqPIREIEPD egqPISESETEVEIHD rounded mus™le s™—rF

egqPITEIEPEQD egqPIUESETD egqPPHEIEPEQD egqPPREID egqPPTE

himensionsF ƒee pigF SF

QD egqPPVEIEPD egqPRPEIEPD egqPRQESD egqPSQEQERD egqPSREQY

his™ussionF e™™ording to sever—l —uthors @eFgF sixteen dors—l v—lvesX egqUVEIEPETD egqPHHEIIEIRD egqPIQ˜isEPD

‚—ffi IWUHY p—r—one IWVTY timenez et —lF PHHWA eequiE egqPISEUEWD egqPITERD egqPIUERD egqPPPESD egqPPSEID

egqPPTEIEPD egqPPWEID egqPSREPY fr—gmentsX egqUVEQD pe™ten s™—˜rell— shows — wide morphologi™—l intr—sE

egqPRIERF

pe™ifi™ v—ri—˜ility —ndD thereforeD some spe™ies should

y™™urren™eX egqUUD egqUVD egqVTD egqPHHD egqPHPD

˜e ™onsidered synonymsF sn p—rti™ul—rD ˜—sed on ™omE

egqPIQ˜isD egqPIRD egqPISD egqPITD egqPIUD egqPPHD

p—rison —nd revision of the —v—il—˜le liter—ture @eFgF egqPPPD egqPPRD egqPPSD egqPPTD egqPPVD egqPPWD

egqPRID egqPRPD egqPRQD egqPSQD egqPSRD erd— ‚iver se™tionD ‚—ffi IWUHD w—l—test— IWURAD eequipe™ten r—di—ns

g—stell9erqu—toD st—lyF @xyst 8 ‡estendropD IVQWAD eequipe™ten seniensis

@v—m—r™kD IVIWA —nd eequipe™ten ˜ollenensis @w—yerD

hes™riptionF v—rge sizedD ˜i™onvex —nd strong

IVUTAD here ™onsidered junior synonyms of eequipe™E

shellD equiv—lve —nd inequil—ter—lY height r—nging ˜eE

ten s™—˜rell—F

tween RI —nd WH mmD lenght r—nging ˜etween RR —nd

eequipe™ten oper™ul—ris @vinn—eusD IUSVA differs

WPFR mmY su˜ov—l outline with rounded —nterior m—rgin

from eF s™—˜rell— ˜y its more symmetri™—l v—lvesD its

—nd str—ight posterior oneY shell su˜st—n™e very thi™kY

more regul—r outlineD its lower ™onvexity —nd ˜y its

˜oth v—lves show — low ridge in the posterior p—rtD exE

thinner —nd more numerous ™ost—eF

tending from the um˜o to the ventr—l m—rginY um˜o

es o˜served in the eF oper™ul—ris dis™ussionD eF

™onvexD prosogyr—te —nd prominentY ™—rdin—l m—rgin

s™—˜rell— —nd eF oper™ul—risD show simil—r size @pigF SA

strongly ™urvedY n—rrow nymphs o™™upied ˜y thi™kD

—nd in ˜oth ™—ses height —nd length —re ne—rly equ—l @rG

—r™hed —nd ˜l—™k extern—l lig—mentY dorsoEventr—l di—E

v r—tio equ—l IA indi™—ting — su˜™ir™ul—r outlineF

meter is ne—rly equ—l to —nteroEposterior oneF

ƒtr—tigr—phi™ —nd geogr—phi™ o™™urren™eF eeE

yrn—ment—tion of ˜oth v—lves ™onsisting ofX IA

quipe™ten s™—˜rell— is known from the wio™ene to the irregul—r —nd numerous ™on™entri™ growth lines in the

qel—si—n of the wediterr—ne—n ƒe— @vower €leisto™eneA —nterior —nd middle p—rt of the v—lve —nd PA growth

@w—r—sti 8 ‚—ffi IWVHAF l—mell—e in the posterior p—rtY these two different orn—E

ment—tion p—ttern —re ˜ounded ˜y — low ridgeF

snterior of ˜oth v—lves with heterodont dentitionY

right v—lve with three prominent ™—rdin—l teeth —nd — ƒu˜™l—ss reterodont— xeum—yrD IVVR

single posterior l—ter—l toothY left v—lve with three ™—rE

yrder †eneroid— ed—ms 8 ed—msD IVST

din—ls teeth —nd one posterior l—ter—l toothD with the

ƒuperf—mily er™ti™oide— xewtonD IVWI

—nterior ™—rdin—l ™ontinuous with — series of sm—ll ridges

p—mily er™ti™id—e xewtonD IVWI —nd denti™ul—tionsF

himy—ri—n —nisomy—ri—nD integrip—lli—te shell

with su˜ov—l mus™le s™—r —nd ™ir™ul—r p—lli—l lineY —nteE

rior —ddu™tor mus™le s™—r slightly l—rger th—n the posterE qenus er™ti™— ƒ™hum—™herD IVIU

ior oneY s™—rs of r—di—lly dire™ted fi˜ers of mus™le m—nE „ypeEspe™iesX er™ti™— vulg—ris @vinn—eusD IUTUA from the €lio™ene

tle —long the p—lli—l lineF of ingl—ndF

wollusk f—un— of the vower €leisto™ene erd— ‚iver se™tion @xorthern st—lyA VI

his™ussionF er™ti™— isl—ndi™— is the only living i™—n ™o—st —nd in iurope —long the ™o—sts of s™el—ndD

spe™ies of its f—milyF „he t—xonomy —nd nomen™l—ture qre—t frit—in —nd the ƒ™—ndin—vi—n €eninsul— @h—hlgE

is well defined —lthough the old n—me gyprin— isl—ndi™— ren et —lF PHHHAF

is sometimes still used in re™ent liter—tureF „his spe™ies

xote of the —uthorsF ‡hile the p—per w—s in pro˜—˜ly represents the most popul—r ˜ore—l guestD

pressD — fr—gment of the dors—l —re— of eF isl—ndi™— w—s whi™h migr—tes into the wediterr—ne—n ƒe— during the

re™overed from — ˜ed —t —˜out S metres from the ˜—se of ™lim—ti™ deterior—tion of the i—rly €leisto™eneF „he

the se™tion of pigF PF „his suggests — first —rriv—l of the wediterr—ne—n popul—tions ˜e™—me extin™t —˜out WFV

spe™ies in the ˜—sin possi˜ly in the l—te qel—si—nF rowE k— —go —s — ™onsequen™e of the ™lim—te shift to w—rm

w—ter ™onditions @h—hlgren et —lF PHHHAF everD so e—rly it w—s not —˜le to re™ruit su™™essfully —nd

er™ti™— isl—ndi™— is the spe™ies with the widest size est—˜lish l—rge popul—tions —s it did str—tigr—phi™—lly

r—nge @minF RI mmD m—x WPDR mmA —nd usu—lly h—ve — —˜ove from IHQ m onw—rdF

rGv minor th—n ID pointing out — slightly tr—nsverse

e™knowledgmentsF ‡e —re gr—teful to –‚iserv— x—tur—le qeoloE outlineF

gi™— del €i—™enzi—no9 —nd –wuseo qeologi™o qF gortesi9 di g—stell9erE

ƒtr—tigr—phi™ —nd geogr—phi™ o™™urren™eF eF isE

qu—to for the permission to enter the ™onserv—tion —re—sF vu™i— engioE

l—ndi™— is known from IFUUEPFHH w— to WFV k— @g—l—E

lini is th—nked for her ™riti™—l re—ding —nd suggestions on the p—perF „his

˜ri—nEv—te €leisto™eneA of the wediterr—ne—n ƒe— m—nus™ript ˜enefits of the useful —nd ™onstru™tive ™omments of ‚os—

@h—hlgren et —lF PHHHAF xow—d—ys is found —t high l—E homeÁne™hD ‚—f—el v— €ern— —nd ƒergio ‚—ffiF gl—udio gor˜ett— is

deeply th—nked for the photos of the spe™imens here presentedF titudes in the ‡estern etl—nti™ y™e—n —long the emerE

left v—lveD xP @egqRIEQPAY W—E˜ E vu™inell— div—ri™—t—D left v—lveD xS

@egqPSWEUAY IH—E˜ E gh—m— gryphoidesD left v—lve @egqQUEUAY II—E˜

E plexope™ten gl—˜erD left v—lve @egqVHERAY IP—E˜ E gh—m— pl—™entin—D

€ve„i T

right v—lve @egqPREIPAF

ell spe™imens —re xID ex™ept when indi™—tedY —A extern—l viewD ˜A

intern—l viewD ex™ept when indi™—tedF

€ve„i W

I—E˜ E xu™ul— pl—™entin—D right v—lve @egqIHEPAY P—E˜ E xu™ul— sulE

™—t—D left v—lve @egqWHEIQAY Q—E˜ E xu™ul—n— pell—D right v—lveD xP ell spe™imens —re xID ex™ept when indi™—tedY —A extern—l viewD ˜A

@egqIWUEIQAY R—E˜ E ƒ—™™ell— ™ommut—t—D left v—lveD xP @egqIHRETAY intern—l viewF

S—E˜ Ec f—r˜—ti— mytiloidesD rigth v—lve of — juvenile spe™imenD xQ IE€e™ten j—™o˜—eusD right v—lve @egqPREIAY P—E˜ E †eneri™—rdi— spFD

@egqPQEIAY T—E˜ E ƒtri—r™— l—™te—D right v—lveD xP @egqIWUEIRAY U—E left v—lve @egqIREVAY Q—E˜ E g—rdites —ntiqu—tusD right v—lve

˜Eer™— no—eD left v—lve @egqPQSEUAY V—E˜ E er™— spFD left v—lveD xP @egqPWEIAY R—E˜ E est—rte fus™—D right v—lve @egqPWETAY S—E˜ E esE

@egqPSPEUAY W—E˜ E en—d—r— ™fF eFdiluviiD left v—lve @egqIPEUAY IH—E t—rte fus™—D left v—lve @egqPSEIPAY T—E˜ E est—rte spFD left v—lveD xP

˜ E erti™ul—ted spe™imen of f—r˜—ti— mytiloidesD right —A —nd left ˜A @egqIQEWAY U—E˜ E e™—ntho™—rdi— —™ule—t—D left v—lve @egqPIUEUAY

v—lves @egqTPEIAY II—E˜ E f—thy—r™— spFD left v—lveD xP @egqIQEIPAY V—E˜ E e™—ntho™—rdi— p—u™i™ost—t—D right v—lve @egqPPQETAY W—E˜ E

IP—E˜ E emusium ™rist—tumD left v—lve @egqTEIAY IQ E pr—gment of e™—ntho™—rdi— e™hin—t—D right v—lve @egqPQPEIAY IH—E˜ E e™—ntho™—rE

€inn— spF @egqPHHEITAY IR E pr—gment of wytilus edulis @egqIIUE di— —™ule—t—D right v—lve @egqPHUESAF

IAY IS—E˜ E wim—™hl—mys v—ri—D right v—lve @egqIIWEIAY IT—E˜ E

„—lo™hl—mys ™fF „F multistri—t—D left v—lveD xP @egqIREIVAY IU—E˜ E

€ve„i IH €seud—mussium peslutr—eD left v—lve @egqPIWEIAY IV E nidentified

v—lve of emusium ™rist—tum @egqPEIAF

ell spe™imens —re xID ex™ept when indi™—tedY —A extern—l viewD ˜A

intern—l viewD ex™ept when indi™—tedF

I—E˜ E e™—ntho™—rdi— tu˜er™ol—t—D left v—lve @egqPSQESAY P—E˜ E erE

€ve„i U ti™ul—ted spe™imen of e™—ntho™—rdi— tu˜er™ol—t—D left —A v—lve —nd

—nterior view ˜A @egqPSIEIAY Q—E˜ E v—evi™—rdium ™r—ssumD left v—lve ell spe™imens —re xIY —A extern—l viewD ˜A intern—l viewF

@egqIWUEITAY R—E˜ E v—evi™—rdium o˜longumD right v—lve @egqIWUE

I—E˜ E €e™ten fl—˜elliformisD left v—lve @egqREIAY P—E˜ E plexope™ten

IUAY S—E˜ E €—pilli™—rdium p—pillosumD right v—lveD xP @egqIWUEIVAY gl—˜erD right v—lve @egqPPUEIAY Q—E˜ E €e™ten j—™o˜—eusD left v—lve

T—E˜ E €—pilli™—rdium p—pillosumD left v—lveD xP @egqIWVEVAY U—E˜ E

@egqRIEPVAY R—E˜ E plexope™ten flexuosusD left v—lve @egqVHESAF

w—™tr— stultorumD left v—lve @egqPHSERAY V—E˜ E €—rvi™—rdium exiE

guumD right v—lveD xP @egqPST˜isEQAY W—E˜ E v—evi™—rdium spFD left

v—lve @egqIWVEWAY IH—E˜ E ƒpisul— su˜trun™—t—D right v—lve @egqPQTE

€ve„i V

UAY II—E˜ E ƒpisul— su˜trun™—t—D left v—lve @egqPQTEVAY IP—E˜ E „ellin—

—l˜i™—nsD right v—lve @egqSQERAY IQ—E˜ E „ellin— in™—rn—t—D right v—lve ell spe™imens —re xID ex™ept when indi™—tedY @—A extern—l viewD @˜A

@egqPPQEUAY IR—E˜ E „ellin— in™—rn—t—D right v—lve @egqPSWEVAY IS—E intern—l viewD ex™ept when indi™—tedF

˜Evutr—ri— —ngustiorD right v—lve @egqPQUESAF I—E˜ E enomi— spFD left v—lve @egqIWUEISAY P—E˜ E €e™ten j—™o˜—eusD

left v—lve @egqPW˜isEQVAY Q—E˜ E enomi— ephippiumD left v—lve

@egqRPEPTAY R—E˜ E enomi— ephippiumD left v—lve @egqRIEQHAY S—E

€ve„i II

˜Ewoni— p—telliformisD left v—lve @egqIIVEIAY T—E˜ E voripes l—™teusD

left v—lve @egqPSWETAY U—E˜ E erti™ul—ted spe™imen of voripinus fr—E ell spe™imens —re xID ex™ept when indi™—tedY —A extern—l viewD ˜A

intern—l viewD ex™ept when indi™—tedF gilisD right —A —nd left ˜A v—lve @egqPVEQAY V—E˜ E vu™inom— ˜ore—lisD

VP gripp— qF 8 ‚—ineri qF

I—E˜ E vutr—ri— o˜long—D left v—lve @egqIPEIAY P—E˜ E woerell— disE @egqIWREWAY IP—E˜ E tuju˜inus spFD —˜—pertur—l —A —nd —pertur—l ˜A

tort—D right v—lveD xP @egqPSWEWAY Q—E˜ E „ellin— pul™hell—D left v—lve viewsD xP @egqIWREIHAY IQ—E˜ E „urritell— tri™—rin—t— pliore™ensD —˜—E

@egqPHUESAY R—E˜ E „ellin— pul™hell—D right v—lve @egqPTSESAY S—E˜ E pertur—l —A —nd —pertur—l ˜A views @IHWEIAY IR—E™ E hilom— p—tulumD

„ellin— serr—t—D right v—lve @egqRI˜isEUAY T—E˜ E hon—x ™fF hF trunE —pi™—l —AD um˜ili™—l ˜A —nd —pertur—l ™A views @egqSQEPQAF

™ulusD left v—lve @egqPIWEPAY U—E˜ E hon—x ™fF hF venustusD left v—lve

@egqPIQ˜isERAY V—E˜ E erti™ul—ted spe™imen of ƒole™urtus s™opul—D

right —A —nd left ˜A v—lves @egqIQIEIAY W—E˜ E ezorinus ™h—m—solenD

€ve„i IR right v—lve @egqIHEIAY IH—E˜ E „ellin— tenuisD right —A —nd left ˜A

v—lveD xP @respe™tively egqIUET —nd egqIUEUAY II E qlossus hum—E ell spe™imens —re xID ex™ept when indi™—tedY —A —˜—pertur—lD ˜A —perE

nusD right v—lve @egqPHHEISAY IP E †enus nuxDdors—l view @egqIQHE tur—l viewD ex™ept when indi™—tedF

IAY IQ—E˜ E †enus nuxD left v—lve @egqIQHEPAY IR—E˜ E €it—r rudisD left IEystre— edulisD —rti™ul—ted spe™imen @egqIQQESAY P E g—pulus unE

v—lve @egqVPEVAY IS—E˜ E €it—r rudisD right v—lve @egqVHEUAY IT—E˜ E g—ri™usD l—ter—l viewD xQ @egqIWREIIAY Q—E˜ E eporrh—is pespele™—ni

g—llist— ™hioneD right v—lve @egqIWUEIWAF @egqPQTEIHAY R—E˜ E g—lyptr—e— ™hinensisD —pi™—l @—A —nd l—ter—l views

@˜A @egqRI˜isEIHAY S—E˜ E eporrh—is uttingeri—n— @egqIWUEPHAY T—E™

E g—lyptr—e— ™hinensisD —pi™—l —AD um˜ili™—l ˜A —nd l—ter—l ™A viewsD xP

@egqPSQEUAY U—E™ E g—lyptr—e— spFD —pi™—l —AD um˜ili™—l ˜A —nd l—ter—l €ve„i IP

™A views @egqSWEWAY V—E™ E ˆenophor— ™risp—D —pi™—l —AD um˜ili™—l ˜A

ell spe™imens —re xID ex™ept when indi™—tedY —A extern—l viewD ˜A

—nd l—ter—l ™A views @egqIWWETAY W—E™ E ˆenophor— ™risp—D —pi™—l —AD

intern—l viewF

um˜ili™—l ˜A —nd l—ter—l ™A views @egqIIESAY IH—E˜ E x—ti™—rius sterE

IE€ele™yor— ˜ro™™hiD left v—lve @egqQUERAY P—E˜ E gh—mele— g—llin—D

™usmus™—rum @egqPW˜isEPHAY II—E˜ E iuspir— spFD xP @egqPTIEQAY

left v—lve @egqPHUETAY Q—E˜ E hosini— lupinusD left v—lve @egqPRVEIAY

IP—E™ E xeverit— josephini—D um˜ili™—l view ™A @egqPQSEIHAY IQ—E™ E

R—E˜ E gh—mele— g—llin—D left v—lve @egqWHEISAY S—E˜ E gl—usinell—

x—ti™—rius spFD um˜ili™—l view ™A @egqPST˜isERAF f—s™i—t—D right v—lveD xP @egqPSEIHAY T—E˜ E gl—usinell— spFD right

v—lveD xP @egqPQEVAY U—E˜ E hosini— lupinusD right v—lve @egqPQSE

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litit—pes rhom˜oidesD left v—lve @egqPHHEITAY IH—E˜ E €olitit—pes ™fF €F €ve„i IS

rhom˜oidesD left v—lve @egqPQUETAY II—E˜ E „imo™le— ov—t—D right

ell spe™imens —re xID ex™ept when indi™—tedY —A —˜—pertur—lD ˜A —perE v—lve @egqPHHEIUAY IP—E˜ E gor˜ul— gi˜˜—D right v—lveD xQ

tur—l viewD ex™ept when indi™—tedF

@egqIWREVAY IQ—E˜ E gor˜ul— gi˜˜—D right v—lveD xP @egqPSQETAY

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ius ™fF xF gi˜˜osulus @egqPSVEIAY S E folinus spF @egqIQQETAY T—E™ E

xeverit— josephini—D um˜ili™—l view ™AD xP @egqPSIEPAY U—E˜ E x—sE

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™e—D xQ @egqPTQETAY PH—E˜ E gonus ventri™osus @egqPVESAY PI—E˜ E view —AD intern—l view ˜AD xPD @egqPSWEIIAY T E „u˜ul—r stru™ture of

gonus spFD xP @egqPW˜isERQAY PP E €yr—midell— spF @egqIHPERAY PQ—E gl—v—gell— spF @egqIWSEVAY U—E˜ E ystre— edulisD right v—lveD extern—l

˜ E i™hinoids indetFD —pi™—l —A —nd or—l ˜A viewsD xQ @egqPTEIHAY PR E view —AD intern—l view ˜A @egqPQSEWAY V—E˜ E ystre— spFD right v—lveD

gl—do™or— spF @egqPHHEPHAY PS E hent—lium spF @egqPWERAY PT E extern—l view —AD intern—l view ˜A @egqVHEWAY W—E˜ E hiodor— gr—e™—D

ƒerpulor˜is spF @egqPW˜isERIAY PU E hitrup— spF @egqWSERAY PV E —pi™—l —A —nd l—ter—l ˜A viewsD xP @egqQPEQAY IH—E˜ E g—lliostom— ™fF

f—rn—™le indetFD top—A —nd l—ter—l ˜A views @egqPSEIUAY PW E fryozo— gF ™onulusD —˜—pertur—l —A —nd —pertur—l ˜A viewsD xP @egqSTEIQ—AY

II—E˜ E tuju˜inus stri—tusD —˜—pertur—l —A —nd —pertur—l ˜A viewsD xP indetF @egqIWWEUAF

wollusk f—un— of the vower €leisto™ene erd— ‚iver se™tion @xorthern st—lyA VQ €ve„i T

VR gripp— qF 8 ‚—ineri qF €ve„i U

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VT gripp— qF 8 ‚—ineri qF €ve„i W

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VV gripp— qF 8 ‚—ineri qF €ve„i II

wollusk f—un— of the vower €leisto™ene erd— ‚iver se™tion @xorthern st—lyA VW €ve„i IP

WH gripp— qF 8 ‚—ineri qF €ve„i IQ

wollusk f—un— of the vower €leisto™ene erd— ‚iver se™tion @xorthern st—lyA WI €ve„i IR

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ennF qeolF €—ys rellenFD „F hors seÁrieX TWWEUHWF pest— eF @IWSHA E lteriori notizie sull— f—un— plio™eni™— del  Â

v—m—r™k tFfF @IVHSA E ƒuite des meÂmoires ƒur les fossiles des qenoves—toF ennF wusF givF ƒtF x—tF qenov—D vˆs†X

environs de €—risF ennF wusF €—risD TX PIREPPID QQUE VSEIHHF

QRSF pont—nnes pF @IVUWEIVVPA E ves mollusques plio™enes de l—

Âe du ‚hone et du ‚oussillonF P volX PUVEQPPF pF v—m—r™k tFf @IVIWA E ristoire n—turelle des —nim—ux s—ns v—lle

verteÁ˜resF ferlinD €—risD QRQ ppF ƒ—vyD €—risF

wollusk f—un— of the vower €leisto™ene erd— ‚iver se™tion @xorthern st—lyA WS

v— €ern— ‚F 8 h9e˜r—mo wF @PHHWA E worphometri™ —nd woneg—tti €F 8 ‚—ffi ƒF @PHHIA E „—xonomi™ diversity —nd

str—tigr—phi™ distri˜ution of wediterr—ne—n €lio™ene system—ti™ study on three e™—ntho™—rdi— spe™ies from

˜iv—lvesF €—l—eogeogrFD €—l—eo™lim—tolFD €—l—eoe™olFD the wediterr—ne—n €leisto™ene @wollus™—D fiv—lvi—D

ITS@QAX IUIEIWQF g—rdiid—eAF qeodiversit—sD QI@QAX TTWETVPF

woneg—tti €FD ‚—ffi ƒFD ‚overi wF 8 „—vi—ni wF @PHHIA E yne ve™ointre qF @IWSPAF ‚e™her™hes sur le xeÂogeÁne et le u—E

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to™ene f—sinX from the f—dl—nd of wonte qiogo to te™tF ‚epF prF w—ro™F ƒervF qeolFD xotes wemF WWD IUR

the ƒtirone ‚iverF €—leo˜iogeogr—phy —nd €—leoe™olE ppF

ogy sntern—tion—l gonferen™eD ix™ursion quide˜ookD vinn—eus gF @IUSVA E ƒystem— x—tur—e per ‚egn— tri— n—turE

niversit—Á di €—rm—D €—rm—D PT ppF —eD se™undum ™l—ssesD ordinesD gener—D spe™ies ™um

woroni wFeF @IWSSA E v— m—™rof—un— s—heli—n— del wessiE ™h—r—™teri˜usD differentiisD synonymisD lo™isF iditio

ni—no inferiore dell— ‚epu˜˜li™— di ƒF w—rinoF qiornF de™im—D reform—t—F „omus IF v—urentii ƒ—lviiD rolE

qeolFD serie PD PSX VIEITPF mi—eD VPR ppF

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wediterr—neoF P @fiv—lviAF fios „—r—sD „—r—ntoD SRT ppF

w—l—test— eF @IWTHEIWTQA E w—l—™of—un— pleisto™eni™— di

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qr—mmi™hele @ƒi™ili—AF wemori— per servire —ll— deE

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trion—lF fullF ƒo™F qeolF pr—n™eD PPX PIHEPUUF

w—l—test— eF @IWURA E w—l—™of—un— plio™eni™— um˜r—F weE

€—ron— gFiF @IVUVA E sl €lio™ene dell9yltrepoÁ €—veseX osserE

mori— per servire —ll— des™rizione dell— g—rt— qeoloE

v—zioni str—tigr—fi™he e p—leontologi™heD etti ƒo™F itF

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w—l—test— eF 8 xi™osi— wFvF @IWSSA E s fossili del €lio™ene e

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ƒervF qeolF stFD UUX IUQEIVHF

degr—d—tion of shell ™—r˜on—teF pourth sntern—tion—l

w—l—test— eF 8 —rleng— pF @IWVTA E xorthern guests in the

€—l—eontologi™—l gongressD wendoz—D ergentin—F PV

€leisto™ene wediterr—ne—n ƒe—F qeolF ‚om—n—D PSX WIE

ƒeptem˜erEQ y™to˜er PHIRF

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€elosio qF 8 ‚—ffi ƒF @IWURA E ysserv—zioni su er™ti™— isl—nE

w—r—sti ‚F @IWUQA E v— f—un— tortoni—n— del „F ƒtirone @limite

di™— ed —ltri l—melli˜r—n™hi del g—l—˜ri—no dell9imili—

€—rmenseE€i—™entinoAF follF ƒo™F €—leontF stFD IP@IAX

o™™ident—leF v9eteneo €—rmenseEe™t— x—tur—li—D IHX

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w—r—sti ‚F 8 ‚—ffi ƒF @IWVHA E v— diversit—Á t—ssonomi™— dei €elosio qF 8 ‚—ffi ƒF @IWUUA E €relimin—ry rem—rks on molE

˜iv—lvi del €lio™ene mediterr—neoF gentro qr—fi™o lus™ —ssem˜l—ges of the ƒtirone river €leisto™ene series

niversit—Á di €—rm—X IEQHF @€—rm— €rovin™eD xorthern st—lyAF ˆ sx e gonE

w—rtinell tF 8 homeÁne™h ‚F @IWVPA E il €lio™eno m—rino de gressD firmingh—mX IEIWF

l— m—rgen dere™h— del rio pluvi— @elt impord—D qiE €ervesler €FD ™hm—n eFD rohenegger tF 8 homini™i ƒF

ron—AF istudF qeolFD QVX QUWEQVRF @PHIIA E s™hnologi™—l re™ord of environment—l ™h—nges

w—yer gF @IVTVA E g—t—logue systeÂm—tique et des™riptif des in e—rly u—tern—ry @qel—si—n±g—l—˜ri—nA m—rine deE

fossiles des terr—ins „erti—ires qui se trouvent —u wuE posits of the ƒtirone ƒe™tionD northern st—lyF €—l—iosD

seÂe peÂdeÂr—l de uri™hF Qeme g—hierF wollusquesD p—E PT@WAX SUVESWQF

mille des er™ideÂsF ƒ™h—˜elitzD uri™hD IPR ppF €es™e qFvF 8 ‚—petti gFeF @IWUIA E w—l—™of—un— plio™eni™—

weli ‚F @IVUWA E xote geologi™he sui dintorni di givit—ve™E d9e˜ruzzoF fiv—lvi del ™ir™ond—rio di ƒF ius—nio del

™hi—F etti ‚F e™™F vin™eiD serF QD †X IEIQF ƒ—ngro @ghietiAF ennF nivF ƒtudi equil—D —nno †F

weli ‚F @IVVRA E genni geologi™i sulle ™oste d9enzio e di ‚—ffi ƒF @IWUHA E s €ettinidi del €lio™ene e g—l—˜ri—no

xettuno ed elen™o dei mollus™hi plio™eni™i ivi r—™™olE dell9imili— o™™ident—le @€i—™entino e €—rmenseAF follF

ƒo™F €—leontolF stFD W@PAX WUEIQSF tiF ennF sstF „e™nF ‚om—D sˆD QI ppF

WT gripp— qF 8 ‚—ineri qF

‚—ffi ƒF @IWVTA E „he signifi™—n™e of m—rine ˜ore—l mollus™s ƒeed ‚F @IWWPA E ƒystem—ti™s evolution —nd distri˜ution of

mussels ˜elonging to the genus wytilusX —n overviewF in the i—rly €leisto™ene f—un—s of the wediterr—ne—n

emF w—l—™olF fullFD WX IPQEIQUF —re—F €—l—eogeogrFD €—l—eo™lim—tolFD €—l—eoe™olFD SPX

ƒeed ‚F @IWWSA E ƒpe™i—tion —nd geogr—phi™—l distri˜ution PTUEPVWF

within the genus wytilusF fullF w—l—™olF ƒo™F vondonD ‚—ffi ƒFD ƒt—nley ƒFwF 8 w—r—sti ‚F @IWVSA E fiogeogr—phi™

PR@RAF p—ttern —nd €lioE€leisto™ene extin™tion of fiv—lvi— in

ƒeguenz— qF @IVUQEIVUUA E ƒtudi str—tigr—fi™i sull— form—E the wediterr—ne—n —nd southern xorth ƒe—F €—leoE

zione plio™eni™— dell9st—li— meridion—leF follF ‚F ˜iologyD II@RAX QTVEQVVF

gomF qeolF stFX PWW ppF ‚—ffi ƒFD ‚io hFD ƒprovieri ‚FD †—lleri qFD woneg—tti €FD ‚—ffi

ƒignor €F ‡F 8 vipps tF rF @IWVPA E ƒ—mpling ˜i—sD gr—du—l sF 8 f—rrier €F @IWVWA E xew str—tigr—phi™ d—t— on the

extin™tion p—tterns —nd ™—t—strophes in the fossil reE €i—™enzi—n str—totypeF follF ƒo™F qeolF stFD IHVX IVQE

™ordF qeolF ƒo™F emeri™— ƒpF €—pFD IWHX PWIEPWTF IWTF

ƒirn— qF @IWUVA E €ro˜lemi di nomen™l—tur—X l— priorit—Ádi ‚iedl ‚F @IWVTA E p—un— y plor— del wediterr—ÂneoF ymeg—D

qly™ymeris insu˜ri™us @fro™™hiAF gon™higlieD IR@WEIHAX f—r™elon—D VSV ppF

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‚io hFD ƒprovieri ‚FD ‚—ffi sF 8 †—lleri qF @IWVVA E fiostr—tiE

ƒki˜inski hFyFpFD ehm—d wF 8 fe—rdmore tFeF @IWUVA E

gr—fi— e p—leoe™ologi— dell— sezione str—totipi™— del

qeneti™ eviden™e for n—tur—lly o™™urring hy˜rids ˜eE

€i—™enzi—noF follF ƒo™F €—leontF stFD PU@PAX PIQEPQVF

tween wytilus edulis —nd wytilus g—lloprovin™i—lisF

‚isso eF @IVPTA E ristoire n—turelle des prin™ip—les produ™E

ivolutionD QP@PAX QSREQTRF

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ƒquires ‚FvF @PHIHA E xorthe—st €—™ifi™ pper gret—™eous

™elles des environs de xi™e et des elpes w—ritimesF

—nd €—leo™ene qly™ymeridid ˜iv—lvesF tF €—leontFD

€—risD pFqF vevr—ultD RQW ppF

VR@SAX VWSEWIU

‚oger tF @IWRHA E €e™tinides mio™enes plio™enes er qu—tern—rE

„—v—ni qF @IWQVEIWQWA E possili del wio™ene dell— giren—i™—F

ie de ƒyrieF r—ut gommF ‚epF prF ƒyrie vi˜—nF xotes

€—l—eontogr—phi— st—li™—D QVX IPUEIVUY QWX IUEUTF

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„—vi—ni wFD ‚overi wFD smpi™™iniD ‚F 8 †igliotti vF @IWWUA E

‚ossi ‚on™hetti gF @IWSIA E s „ipi dell— «gon™hiologi— fosE

ƒegn—l—zione di u—tern—rio m—rino nell— †—l ghero

sile su˜—pennin—» di qF fro™™hiF ‚ivF stF €—leontolF

@eppennino pi—™entinoAF follF ƒo™F €—leontF stFD QTX

ƒtr—tFD SUX PETQF

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‚overi wF 8 „—vi—ni wF @PHHQA E g—l™—renite —nd s—propel

„e˜˜le xF @IWTTA E fritish fiv—lve se—shellsF ‚oy—l ƒ™ottish

deposition in the wediterr—ne—n €lio™eneX sh—llowE

wuseum ˜y rer w—jesty9s ƒt—tionery yffi™eD idinE

—nd deepEw—ter re™ord of —stronomi™—lly driven ™liE

˜urghD PIP ppF

m—ti™ eventsF „err— xov—D IS@RAX PUWEPVTF

golini ‚F @IWHTA E wonogr—fi— dei €ettinidi neogeni™i dell—

‚uggieri qF @IWRW—A E sl €lio™ene superiore di g—po™olle

ƒ—rdegn—F €—rte sX qeneri ghl—mysD rinnitesD sn—equiE

@porlõÁAF qiornF qeolFD serie PD PHX IWEQVF

pe™tenF €—l—eontogr—phi— st—li™—D IPX ISSEPHTF

‚uggieri qF @IWRW˜A E v— m—l—™of—un— del g—l—˜ri—no ‚oE

†—n gouvering tF eF @IWWUA E „he €leisto™ene ˜ound—ry —nd

m—gnoloF qiornF qeolFD serie PD PHX TQEIIHF

the ˜eginning of the u—tern—ryF g—m˜ridge niverE

‚uggieri qF @IWSU—A E qeologi— e str—tigr—fi— dell— sommit—Á

sity €ressD PWT ppF

del terzi—rio — g—stro™—roF qiornF qeolFD serie PD PTX IE †enzo ƒF @IWQQEIWQSA E possili del xeogene „rentinoD †eroE

SPF nese e fres™i—noF €—l—eontogr—phi— st—li™—D QRX QIEVRY

‚uggieri qF @IWSU˜AF wollus™hi plio™eni™i sopr—vissuti nel QSX PHIEPSSF

g—l—˜ri—noF etti ƒo™F „os™—n— ƒ™iF x—tFD serie eD TRX †enzo ƒF 8 €elosio qF @IWTQA E v— m—l—™of—un— tortoni—n—

VHEVUF del golle di †igoleno @€re—ppennino pi—™entinoA E

ƒ™h—ffer pFˆF @IWIHEIWIPA E h—s wio™—n von iggen˜urgF hie È €—l—eontF stFD SVX RQEPIQF

p—un— der ersten wediterr—nstufe des ‡iener fe™kens †erri eF @IVVTA E ezione delle forze nell9—ssetto delle †—lli

und die geologis™hen †erh—ltnisse der mge˜ung des È ™on eppendi™e sull— distri˜uzione dei fossili nell— †—lE

w—nh—rts˜erges in xiederosterrei™hF hie q—stropoE È di™hi—n— e nell9 m˜ri— intern— settentrion—leF follF

den der wio™—n˜ildungen von iggen˜urgF e˜hF uFuF È ƒo™F qeolF stFD SX RITERSRF

qeolF ‚ei™hFD PP@IAX QEIPTF ‡—ller „F‚F @IWTWA E „he evolution of the ergope™ten gi˜˜us

ƒ™hum—™her gF pF @IVIUA E iss—i d9un nouve—u systeÁme des sto™k @wollus™—X fiv—lvi—AD with imph—sis on the „erE

h—˜it—tions des vers test—™eÂsF †F IEQF @ƒ™hultzA gopenE ti—ry —nd u—tern—ry spe™ies of i—stern xord emerE

i™—nF tF €—leontolFD RQD IPS ppF h—gueD PVU ppF

wollusk f—un— of the vower €leisto™ene erd— ‚iver se™tion @xorthern st—lyA WU eppendix N. cf. sp., Venus nux Venus sp., illicardium papillosum, papillosum, illicardium subrica, Loripinus fragilis, Dosinia lupinus, Glycymeris sp., S. forskaeli S. humanus, Lutraria oblonga, Nassarius Nassarius oblonga, Lutraria humanus, cf. , Ostrea edulis, Pap edulis, Ostrea , T. multistriata, Venericardia cf. cf. Dentalium Saccostrea sp. Spisula subtruncata, Tellina pulchella pulchella Tellina subtruncata, Spisula Timoclea ovata, Xenophora crispa Xenophora ovata, Timoclea sp., sp., Glycymeris inflata, Glycymeris in sp. Corals indet., Corals Talochlamys Turritella sp., sp. pliorecens, Venus nux is Glycymeris Glycymeris , Glycymeris insubrica, Glossus Glossus insubrica, Glycymeris , Terebratula Terebratula Ostrea edulis, Pinna Pinna edulis, Ostrea sp., Echinoids indet., indet., Echinoids sp., Dosinia lupinus, insubrica, Glycymeris Lucinoma borealis, prysmaticus, Nassarius Glycymeris insubrica, Pecten flabelliformis la subtruncata, Tellina sp., sp., Tellina la subtruncata, sp., sp., sp., sp., Trochidae indet., indet., Trochidae sp., Glycymeris insubrica, Nassarius semistriatus Nassarius insubrica, Glycymeris Echinoids indet., indet., Echinoids sp., Gastropods indet., Bathyarca Bathyarca sp., la auricolata, Spisu la auricolata, Corbula gibba, Dentalium Dentalium gibba, Corbula Diodora graeca, Diodora sp., Echinoids indet., indet., Echinoids sp., Flabellum bertii, Glycymeris inflata Glycymeris Flabellum bertii, Cardium indicum, Clausinella fasciata, Pecten jacobaeus, Spisula subtruncata Spisula jacobaeus, Pecten fasciata, Clausinella indicum, Cardium sp., sp., Saccella commutata, Tellina pulchella, Tellina Glycymeris insubrica, Glycymeris Nassarius sp., Corbula gibba, Dosinia Dosinia gibba, Corbula sp., Chama gryphoides, Chama placentina, Chamelea gallina, Chamelea placentina, Chama gryphoides, Chama sp., sp., Corals indet., sp. Pitar rudis, Pitar rudis, Tellina pulchella, Timoclea ovata ovata Timoclea pulchella, Tellina rudis, Pitar rudis, Pitar ? rum, Ostrea edulis, Pelecyora brocchi, Turritella tricarinata Turritella brocchi, Pelecyora edulis, Ostrea rum, sp., Barnacles indet., Barnacles sp., Clausinella raea chinensis, Conus Conus chinensis, raea rbula gibba, Glycymeris glycymeris, Glycymeris inflata, Glycymer inflata, Glycymeris glycymeris, Glycymeris gibba, rbula Carditidae indet., Glycymeris glycymeris Glycymeris Ostrea edulis, Pecten jacobaeus, Ringicu Pecten jacobaeus, edulis, Ostrea sp. sp., Pecten jacobaeus, glycymeris, Loripinus fragilis, Ostrea edulis Ostrea fragilis, Loripinus glycymeris, Nassarius semistriatus, Pinna Pinna semistriatus, Nassarius Echinoids indet. Echinoids sp. Nucula placentina, Tellina tenuis Tellina placentina, Nucula Barbatia mytiloides, ? Turritella Nassarius Capulus ungaricus, Clavagella Clavagella ungaricus, Capulus Callista chione, Calypt chione, Callista Echinoids indet., indet., Echinoids Callista chione, Calyptraea chinensis, Dentalium Calyptraea chinensis, A.diluvii, Astarte fusca, Corbula gibba, Dosinia Dosinia gibba, Corbula fusca, Astarte A.diluvii, sp., sp., sp., Glycymeris insubrica,Glycymeris edulis Ostrea cf. sp., sp., ardium papillosum, papillosum, ardium Corbula gibba, gibba, Corbula ? Thracia sp., Echinoids indet., indet., Echinoids sp., Bryozoa indet., Bryozoa indet., Bryozoa sp., Corals indet. Corals sp. , Papillic , sp., Carditidaesp., indet., sp., Echinoids indet. Echinoids sp., Aequipecten opercularis, Aequipecten scabrella, Amusium cristatum, Astarte Astarte cristatum, Amusium scabrella, Aequipecten opercularis, Aequipecten Anomia Pelecyora brocchi, Pinna Pinna brocchi, Pelecyora Murex sp., sp., Naticarius stercusmuscarum, Ostrea edulis, Pecten jacobaeus, Ringicula auricolata, Serpulorbis Serpulorbis auricolata, Ringicula jacobaeus, Pecten edulis, Ostrea stercusmuscarum, Naticarius Glycymeris insubrica, Pelecyora Glycymeris brocchi Pelecyora brocchi Anomia Anomia Tellina sp., sp., sp., sp., sp., Acteon Acteon Biota Anadara opercularis, Aequipecten Terebratula clathtratus, Acanthocardia Co antiquatus, Cardites fusca, Astarte opercularis, Aequipecten gibba Corbula Dosinia ? Ostrea edulis Acanthocardia fragments paucicostata, Acanthocardia fragments tetragona, Arca opercularis, Aequipecten Anomia ephippium, Anomia fusca, Astarte opercularis, Aequipecten fusca, Astarte opercularis, Aequipecten Venericardia edulis, Ostrea lupinus, Dosinia opercularis, Aequipecten Aequipecten glycymeris opercularis, Glycymeris Glycymeris ventricosus, Conus ungaricus, Capulus opercularis, Aequipecten Nassarius Dentalium antiquatus, Cardites ovata Timoclea insubrica, Glycymeris lupinus, Dosinia noae, Arca opercularis, Aequipecten Aequipecten opercularis, Calyptraea insubrica chinensis, Glycymeris opercularis, Aequipecten edulis Ostrea scabrella, Aequipecten opercularis, Aequipecten opercularis Aequipecten glycymeris Glycymeris pespelecani, Aporrhais stercusmusca Naticarius gryphoides, Chama opercularis, Aequipecten Mimachlamys varia, edulis Ostrea lupinus Dosinia indet.,Bivalves Aporrhais opercularis, Aequipecten Tellina serrata Acanthocardia aculeata, Aequipecten opercularis, Callista chione, Clavagella opercularis Aequipecten insubricaGlycymeris Dentalium Lutraria indet., Echinoids insubricaGlycymeris opercularis Aequipecten tubercolata, Acanthocardia opercularis Aequipecten opercularis Aequipecten Aequipecten opercularis, Anomia Anomia opercularis, Aequipecten Ditrupa sp., Dentalium sp., Aporrhais ephippium, Anomia opercularis, Aequipecten paucicostata, Acanthocardia Nassarius semistriatus Nassarius Laevicardium oblongum, Naticarius Naticarius oblongum, Laevicardium ID Number ACG13 ACG14 ACG15 ACG16 ACG17 ACG18 ACG19 ACG20 ACG21 ACG22 ACG23 ACG25 ACG26 ACG27 ACG27bis ACG28 ACG29 ACG30 ACG31 ACG32 ACG33 ACG34 ACG35 ACG36 ACG37 ACG38 ACG39 ACG40 42bis ACG43 ACG44 ACG45 ACG46 ACG47 ACG48 ACG49 ACG50 ACG51 ACG51bis Meters 0 m m 0.15 m 1.20 ACG12 m 2.90 m 3.50 m 4.50 m 7.90 m 8.70 m 12.92 m 15.60 m 16.50 m 18.70 m37.05 ACG24 m 37.50 m 37.90 m 39.88 m 40.60 m 41.35 42 m m 42.25 ACG29bis m 43.25 m 43.60 m 44.90 m 45.65 m 46.05 m 47.35 m 47.50 49 m 51 m m 52.50 m 53.95 54 m ACG41-42 m 54.20 55 m m 55.40 m 56.35 m 57.35 m 58.35 59 m m 64.20 m 65.80 m 66.50 54.10 m54.10 ACG41bis-

WV gripp— qF 8 ‚—ineri qF r rudis, Spisula subtruncata Spisula rudis, r itella tricarinata pliorecens Tellina incarnata Tellina Spisula subtruncata Spisula sp., sp., um papillosum, Pita um papillosum, Venus nux Venus Jujubinus striatus, Naticarius stercusmuscarum, Nassarius Nassarius stercusmuscarum, Naticarius striatus, Jujubinus sp., inna inna utabilis, Ostrea edulis, Turr uscarium, Ostrea Ostrea uscarium, dulis, Papillicardi dulis, cusmuscarum, Neverita josephinia, Ostrea edulis, Tellina albicans, Tellina papillosum Flexopecten glaber, Flexopecten glaber, ardium ardium Papillic Ensis ensis, Glycymeris insubrica, Glycymeris Laevicardium ensis, Ensis sp., na, Diloma patulum, Diloma patulum, na, Pitar rudis, Polititapes senescens sp. sp. ungaricus, Chamelea gallina, Ostrea e Ostrea gallina, Chamelea ungaricus, sp., Ditrupa Ditrupa Nassarius prysmaticus, prysmaticus, Nassarius sp., sp., sp. T. multistriata, Venus nux cf. Panopea glycymeris, Venus nux Venus Panopea glycymeris, C. conulus, Chamelea galli Chamelea conulus, C. sp. cf. cf. Laevicardium oblongum, Nassarius obliquatus, Nucula sulcata, P sulcata, Nucula obliquatus, Nassarius oblongum, Laevicardium sp., Ostrea Ostrea ides, Calyptraea chinensis, Capulus Capulus chinensis, Calyptraea ides, Aequipecten scabrella, Callista chione, Glycymeris insubrica, Naticarius stercusmuscarum ? sp., ? Venus nux Venus ycymeris insubrica, Ostrea edulis, Turritella ularis, Diloma patulum, Glycymeris insubrica, Nassarius obliquatus, Naticarius ster Naticarius obliquatus, Nassarius insubrica, Glycymeris patulum, Diloma ularis, sp., Glycymeris insubrica Glycymeris Naticarius Barbatia mytilo Barbatia sp., sp., . sp Calyptraea ? Bryozoa indet., Bryozoa Venus nux Venus Pecten jacobaeus, Turritella Turritella jacobaeus, Pecten Glycymeris insubrica Glycymeris sp. sp., sp. sp. Dosinia Dosinia Aequipecten opercularis, Chamelea gallina, Diloma patulum, Ostrea Ostrea patulum, Diloma gallina, Chamelea opercularis, Aequipecten operc Aequipecten tubercolata, Acanthocardia corbis subtruncata Spisula edulis, Ostrea gallina, Chamelea opercularis, Aequipecten stercusm Naticarius obliquatus, Nassarius patulum, Diloma gallina, Chamelea opercularis, Aequipecten tubercolata, Acanthocardia Acanthocardia tubercolata, Aequipecten opercularis, Calliostoma josephinia,prysmaticus, Neverita Spisula subtruncata Pinna Pinna opercularis Aequipecten lupinus Dosinia opercularis, Aequipecten rudis Pitar oblongum, Laevicardium insubrica, Glycymeris flexuosus, Flexopecten opercularis, Aequipecten opercularis Aequipecten Barbatia mytiloides nux Venus ? nux Venus opercularis, Aequipecten tubercolata, Acanthocardia indet., Echinoids Aequipecten insubrica opercularis, Glycymeris Pitar rudis Aequipecten opercularis, Spisula subtruncata, Turritella tricarinata pliorecens, Venus nux aculeata, Acanthocardia Aequipecten opercularis, Glycymeris Ostrea islandica, Arctica opercularis, Aequipecten tubercolata, Acanthocardia pliorecens tricarinata Turritella islandica, Arctica Ostrea edulis Ostrea edulis, Ostrea glaber, Flexopecten flexuosus, Flexopecten opercularis, Aequipecten Pinna opercularis, Aequipecten Aequipecten insubrica opercularis, Glycymeris Talochlamys edulis, Ostrea flexuosus, Flexopecten opercularis, Aequipecten nux Venus opercularis, Aequipecten echinata, Acanthocardia gallina Chamelea islandica, Arctica gallina Chamelea Aequipecten opercularis, Barbatia mytiloides, Gl insubrica, Glycymeris gallina, Chamelea opercularis, Aequipecten Pinna insubrica, Glycymeris ungaricus, Capulus gallina, Chamelea opercularis, Aequipecten Dentalium gallina, Chamelea opercularis, Aequipecten tubercolata, Acanthocardia Aequipecten opercularis, Chamelea gallina, insubrica, Glycymeris Laevicardium Acanthocardia aculeata, Acanthocardia tubercolata, Aequipecten insubrica, opercularis, Glycymeris Mactra m stultorum, Nassarius Ditrupa insubricaGlycymeris insubricaGlycymeris insubricaAequipecten Glycymeris ensis, opercularis, Ensis insubricaGlycymeris insubricaGlycymeris rudis Pitar opercularis, Aequipecten Aequipecten opercularis, Aequipecten ACG52 ACG54 ACG55 ACG56bis ACG56bis2 ACG57 ACG58 ACG59 ACG60 ACG61 ACG62 ACG63 ACG64 ACG65 ACG66 ACG67 ACG68 ACG69 ACG70 ACG75 ACG76 ACG77 ACG78 ACG79 ACG80 ACG81 ACG83 ACG84 ACG85 ACG86 ACG88 ACG89 ACG90 ACG91 ACG92 ACG93 ACG94 ACG95 ACG96 ACG97bis ACG97 ACG98 ACG99 ACG100 67.70 m 67.70 m68.40 ACG53 m 68.53 m 69.85 m70.02 ACG56 m 80.30 81 m m 81.60 m 84.30 m 85.35 86 m m 86.90 89 m m 89.60 91 m m 91.40 m 92.50 m 94.20 m 94.60 m 96.20 m 98.30 101 m m 102.20 m 103.70 m 104.10 m 104.80 m 106.50 108 m m111.60 m 114.30 ACG82 m 122.90 m 125.50 m 127.95 130 m m 132.25 m 133.25 m 133.90 m 134.35 m 134.70 m 135.70 140 m m 142.50 m 142.95 m 143.40 m 143.60 144 m m 145.35

wollusk f—un— of the vower €leisto™ene erd— ‚iver se™tion @xorthern st—lyA WW

, sp. ., Ditrupa Ditrupa ., sp Glycymeris insubrica,Glycymeris rudis, Striarca lactea, lactea, Striarca rudis, sp., Nassarius prysmaticus, prysmaticus, Nassarius sp., . sp bula gibba, Dentalium bula gibba, illicardium papillosum, Pitar Pitar papillosum, illicardium , Nucula sulcata, Ostrea edulis, Papillicardium papillosum, sp. Lucinoma borealis, Nassarius prysmaticus, Ringicula Ringicula prysmaticus, Nassarius borealis, Lucinoma sp., itella tricarinata pliorecens, Venus nux Venus pliorecens, tricarinata itella Xenophora crispa Xenophora hamelea gallina, Cor hamelea gallina, Nassarius sp., Arctica islandica, Calyptraea chinensis, Cladocora Cladocora chinensis, Calyptraea islandica, Arctica Nuculana pella, Pap pella, Nuculana Spisula subtruncata, Tellina Tellina subtruncata, Spisula Laevicardium sp., sp., sp., ata, Timoclea ovata, Turr ovata, Timoclea ata, Spisula subtruncata Spisula Nucula placentina, Xenophora crispa Xenophora placentina, Nucula sp., sp., Spisula subtrunc Spisula oblongum, Mytilus galloprovincialis, Mytilus oblongum, Aporrhais uttingeriana, Callista chione, C chione, Callista uttingeriana, Aporrhais Jujubinus striatus, Jujubinus Jujubinus striatus, Jujubinus llina pulchella Epitonium turtonis, Glycymeris insubrica, Laevicardium oblongum, Laevicardium Laevicardium oblongum, Laevicardium insubrica, Glycymeris turtonis, Epitonium sp., sp., Venus nux Venus ilis, Nassarius prysmaticus, Naticarius prysmaticus, Nassarius ilis, Pitar rudis, Polititapes sp., sp., sp., ? subtruncata, Te subtruncata, udamussium peslutrae, peslutrae, udamussium Terebratula Terebratula Glycymeris insubrica,Glycymeris Laevicardium Jujubinus striatus, Nassarius prysmaticus, Ringicula Ringicula prysmaticus, Nassarius striatus, Jujubinus ? sp. Capulus ungaricus, Flexopecten flexuosus, Ostrea edulis, Pinna Pinna edulis, Ostrea flexuosus, Flexopecten ungaricus, Capulus Venus nux Venus sp., Hiatella rugosa, Laevicardium Laevicardium Hiatella rugosa, sp., sp., Ditrupa Ditrupa Ostrea edulis, Ostrea Pyramidella Bryozoa indet., indet., Bryozoa auricolata, Spisula auricolata, sp., sp., lititapes senescens, Pse senescens, lititapes Venus nux Venus Ditrupa Ditrupa Dentalium Ostrea edulis, Saccella commutata m, Laevicardium oblongum, Nassarius mutab Nassarius oblongum, Laevicardium m, sp., Dentalium gibba, Corbula ungaricus, Capulus Mytilus edulis, edulis, Mytilus sp., sp., sp., Venus nux Venus sp., homboides, Po homboides, sp. is insubrica, Glossus humanus, humanus, Glossus insubrica, is sp., Venus nux Venus sp., Venus nux Venus Aporrhais Aporrhais Anomia ? ? sp., Tellina pulchella, Venus nux nux Venus pulchella, Tellina illicardium papillosum, Ringicula papillosum, illicardium tilus edulis, Pinna edulis, tilus sp., Panopea Panopea Venus nux Venus A. paucicostata, Ostrea edulis, Serpulorbis Serpulorbis edulis, Ostrea paucicostata, A. sp., sp., cf. Pitar rudis, Polititapes r Polititapes rudis, Pitar sp., Timoclea ovata, Venus nux, Venus Venus nux, Venus ovata, Timoclea Pinna Pinna Aequipecten opercularis Aequipecten opercularis, Aequipecten pespelecani Aporrhais opercularis, Aequipecten tubercolata, Acanthocardia Naticarius opercularis, Aequipecten opercularis Aequipecten Pitar rudis, Spisula subtruncata, Turritella tricarinata pliorecens, Venus nux Anomia Turritella sp., tricarinata pliorecens, Venus nux Pinna lupinus, Dosinia nux Venus pliorecens, tricarinata Turritella sp., Pinna paucicostata, Acanthocardia Panopea glycymeris My lupinus, Dosinia Acanthocardia nux Venus Acanthocardia nux Venus nux Venus edulis, Ostrea opercularis, Aequipecten Acanthocardia echinata, Mytilus edulis, Venus nux Mimachlamys varia, Monia patelliformis, Ostrea edulis varia, Mimachlamys humanus, Glossus Acanthocardia Teredo echinata, Acanthocardia edulis,Ostrea Turritella tricarinata pliorecens, Venus nux nux Venus nux Venus edulis, Ostrea edulis,Ostrea Turritella tricarinata pliorecens, Venus nux nux Venus nux Venus chamasolen, Azorinus nux Venus nux Venus nux Venus scopula Solecurtus opercularis Aequipecten Bolinus opercularis, Aequipecten paucicostata, Acanthocardia opercularis, Aequipecten Spisula auricolata, Clavagella opercularis, Aequipecten opercularis Aequipecten Anomia opercularis, Aequipecten tubercolata, Acanthocardia echinata, Acanthocardia crassu Laevicardium striatus, Jujubinus Ditrupa chinensis, Calyptraea opercularis, Aequipecten tubercolata, Acanthocardia Pap pella, Nuculana opercularis, Aequipecten tubercolata, Acanthocardia opercularis Aequipecten Aequipecten opercularis, Dosinia lupinus, insubrica, Glycymeris Laevicardium insubrica Glycymeris islandica, Arctica opercularis, Aequipecten Galeodea echinophora, Glycymer Galeodea echinophora, Acanthocardia echinata, Acanthocardia paucicostata, Acanthocardia tubercolata, Aequipecten opercularis, Aporrhais pespelecani, pespelecani, Aporrhais opercularis, Aequipecten tubercolata, Acanthocardia paucicostata, Acanthocardia echinata, Acanthocardia ACG101 ACG102 ACG103 ACG104 ACG105 ACG106 ACG107 ACG108 ACG109 ACG111 ACG110 ACG112 ACG113 ACG114 ACG115 ACG116 ACG117 ACG118 ACG119 ACG120 ACG121 ACG122 ACG123 ACG124 ACG125 ACG126 ACG127 ACG128 ACG129 ACG130 ACG131 ACG132 ACG133 ACG195 ACG196 ACG199 ACG200bis ACG201 ACG202 173.30 m173.30 ACG198 145.70 m 145.70 m 146.20 m 146.45 m 146.55 m 146.80 m 147.10 150 m m 150.50 151 m m 151.10 m 152.40 m 152.80 154 m m 154.50 m 155.80 156 m m 156.10 m 157.90 m 158.10 m 158.70 m 159.10 160 m m 160.15 161 m m 161.40 164 m m 164.80 m 166.10 m 166.70 m 167.90 170 m 171 m m 171.50 172 m m 172.20 ACG194 m 172.30 m172.70 ACG197 m 173.60 174 m ACG200 m 174.30 m 174.70 175 m

IHH gripp— qF 8 ‚—ineri qF sp. Polititapes utrae, Spisula Spisula utrae, sp., Dosinia lupinus, lupinus, Dosinia Pitar sp., ? illicardium papillosum, papillosum, illicardium Donax Donax eudamussium pesl eudamussium sp., Tellina pulchella, Tellina sp., Ditrupa Ditrupa Nucula sulcata, Pap sulcata, Nucula sp., Pandora inaequivalvis, inaequivalvis, Pandora Pinna Pinna sp. sp., sp., Ensis ensis, Laevicardium oblongum, Spisula Spisula oblongum, Laevicardium ensis, Ensis sp., Nucula sp., Nucula placentina, placentina, Nucula ardium papillosum, Pitar rudis, Ps rudis, Pitar papillosum, ardium sp., Glycymeris insubrica, Laevicardium oblongum, Ostrea edulis, edulis, Ostrea oblongum, Laevicardium insubrica, Glycymeris ncarnata sp., p., Arca noae, Dentalium Dentalium noae, Arca p., Papillic Naticarius dulis, Pitar rudis, Polititapes rhomboides, Spisula subtruncata, Tellina la subtruncata, Turritella Turritella subtruncata, la sp., sp., Nassarius Donax Donax sp., P. rhomboides, Spisula subtruncata, Tellina pulchella Tellina subtruncata, Spisula rhomboides, P. sp., Ensis ensis, Neverita josephinia, Neverita ensis, Ensis Spisula Pitar rudis, subtruncata Mitra cf. cf. , Arctica islandica, Ditrupa Ditrupa islandica, Arctica , sp., sp., sp. sp. Pitar rudis, Spisu rudis, Pitar Ditrupa Laevicardium oblongum, Ostrea edulis, Ostrea Ostrea edulis, Ostrea oblongum, Laevicardium uncata, Venus nux Venus uncata, sp., sp., nus nux, Venus Venus nux, nus na, Laevicardium oblongum, Tellina i Tellina oblongum, Laevicardium na, is insubrica, Laevicardium Laevicardium insubrica, is um tiberii, Spisula subtruncata Spisula um tiberii, , Chamelea gallina, gallina, Chamelea , s, Spisula subtr Spisula s, Polititapes senescens sp., ubtruncata, Ve ubtruncata, sp. Ostrea edulis, Pitar rudis, Polititapes Chamelea gallina, Corbula gibba, Ditrupa Ditrupa gibba, Corbula gallina, Chamelea sp., Xenophora crispa crispa Xenophora sp., sp., ani, Arctica islandica Arctica ani, Calyptraea chinensis, Dentalium elliformis, Nassarius mutabilis, Neverita josephinia, Ostrea e subtruncata, Turritella tricarinata pliorecens, Venus nux Venus Venus Azorinus chamasolen, Chamelea gallina, Ditrupa Ditrupa gallina, Chamelea chamasolen, Azorinus Spisula subtruncata Spisula D. trunculus, Dosinia lupinus, Flexopecten glaber, Naticarius Naticarius glaber, Flexopecten lupinus, Dosinia trunculus, D. sis, Chamelea gallina, Glycymer gallina, Chamelea sis, ecten opercularis, Chamelea galli Chamelea ecten opercularis, sp., sp., sp., cf. ensis, Chamelea gallina, Epitoni gallina, Chamelea ensis, lititapes senescens, Spisula s Spisula senescens, lititapes D. venustus, Naticarius stercusmuscarum, Spisula subtruncata Spisula stercusmuscarum, Naticarius venustus, D. Bryozoa indet., indet., Bryozoa Anomia Anomia Donax sp. ? , Glycymeris glycymeris, Naticarius glycymeris, Glycymeris , sp., lititapes rhomboides, Polititapes senescen Polititapes rhomboides, lititapes Lutraria angustior, Mytilus Mytilus angustior, Lutraria indet. , Calyptraea chinensis, Corbula gibba, Flexopecten glaber glaber Flexopecten gibba, Corbula chinensis, Calyptraea , Spisula subtruncata Spisula sp. opercularis, Aporrhais pespelec Aporrhais opercularis, dia paucicostata, Aequip dia paucicostata, sp. Carditidae a islandica, Calyptraea chinen Calyptraea a islandica, Anomia Anomia chinensis, Pitar rudis, Po rudis, Pitar chinensis, M. galloprovincialis,M. Neverita Polititapes senescens ? ctica islandica, Calyptraea chin Calyptraea ctica islandica, sp., ? cf. cf. Bryozoa indet., indet., Bryozoa Calyptraea chinensis, chinensis, Calyptraea sp., elea gallina, Cladocora Cladocora elea gallina, sp. nata, Aequipecten nata, Solecurtidae sp. Solecurtidae aculeata, Acanthocar aculeata, Spisula subtruncata, Tellina pulchella Tellina subtruncata, Spisula Arctica islandica, Pitar rudis, Po rudis, Pitar islandica, Arctica Turridae indet. Turridae Venus nux Venus sp., sp., sp. sp., Panopea Panopea Acanthocardia tubercolata, Chamelea gallina, insubrica, Glycymeris Naticarius Acanthocardia tubercolata, Acanthocardia paucicostata, Acanthocardia ? Chamelea gallina, insubrica, Glycymeris Mactra stultorum, Spisula subtruncata insubricaGlycymeris opercularis, Aequipecten aculeata, Acanthocardia lupinus Dosinia opercularis, Aequipecten Aequipecten opercularis, Dosinia lupinus, Nassarius mutabilis, Spisula subtruncata insubricaGlycymeris insubrica,Glycymeris edulis Naticarius Ostrea stercusmuscarium, nux Venus opercularis, Aequipecten opercularis Aequipecten Donax islandica, Arctica opercularis, Aequipecten islandica Arctica Arctic Aequipecten opercularis, Pitar rudis, Polititapes rhomboides, Polititapes senescens, Spisula Anomia opercularis, Aequipecten paucicostata, Acanthocardia echinata, Acanthocardia aculeata, Acanthocardia Tellina subtruncata, subtruncata Spisula aculeata, Acanthocardia Cham islandica, Arctica subtruncata, Ar Aequipecten opercularis, rudis Pitar paucicostata, Acanthocardia echi Acanthocardia Acanthocardia islandica Arctica Aporrhais Calyptraea islandica, Arctica Calyptraea Flexopecten chinensis, glaber Mytilus islandica, Arctica paucicostata, Acanthocardia paucicostata Acanthocardia echinata, Acanthocardia Lutraria nux Venus echinata, Acanthocardia Ostrea nux Venus s Aporrhais ephippium, Anomia opercularis, Aequipecten semistriatus, Acteon paucicostata, Acanthocardia echinata, Acanthocardia Ensis ensis, Flexopecten glaber, Laevicardium oblongum, Monia pat Dentalium chinensis, Calyptraea pespelecani, Aporrhais ephippium, Anomia opercularis, Aequipecten Turritella pulchella, Tellina subtruncata, Spisula senescens, Aequipecten opercularis, opercularis, Aequipecten opercularis, Aequipecten tubercolata, Acanthocardia Ditrupa opercularis, Aequipecten Anomia islandica, Arctica islandica, Arctica Dosinia lupinus, Mytilus pulchella, Turritella tricarinata pliorecens ACG203 ACG205 ACG206 ACG207 ACG208 ACG209 ACG211 ACG210 ACG212 ACG213 ACG213bis ACG214 ACG218 ACG220 ACG221 ACG222 ACG223 ACG224 ACG225 ACG226 ACG227 ACG228 ACG229 ACG230 ACG231 ACG232 ACG233 ACG234 ACG237 ACG238 ACG239 ACG240 ACG241 ACG242 ACG242bis 186.40 m186.40 ACG217 175.70 m 175.70 176 m m 176.10 ACG204 m 176.60 m 177.40 m 177.60 m 177.80 m 180.40 m 181.20 m 181.80 m 182.10 m 182.80 m 185.60 m185.70 ACG215-216 m 186.50 187 m m 188.80 ACG219 m 189.30 m 189.70 m 189.90 m 193.20 m 193.40 m 193.60 m 193.90 194 m m 194.10 196 m m 197.80 m 202.40 m 205.40 207 m m208.40 ACG235 209 m m 209.80 m 210.40 m 210.60 m 217.20 m 217.90 218 m 208.60 m208.60 ACG236

wollusk f—un— of the vower €leisto™ene erd— ‚iver se™tion @xorthern st—lyA IHI sp. Turritella sp., subtruncata, Tellina Tellina subtruncata, Nassarius scens sp., Spisula subtruncata, Tellina Tellina subtruncata, Spisula rdium papillosum, Pitar rudis, Mytilus sp., sp., josephinia, Spisula Spisula josephinia, Loripes lacteus, Lucinella divaricata, , Ditrupa Ditrupa , Glycymeris insubrica, Glycymeris Spisula subtruncata, sp. llina pulchella, Thracia pube Thracia llina pulchella, sp., ysmaticus, Papillica ysmaticus, Helicidae indet,Helicidae N. gibbosulus, Neverita Neverita gibbosulus, N. cf. cf. ssarius prysmaticus, Tellina pulchella Tellina prysmaticus, ssarius rum, Nassarius pr Nassarius rum, inia, Ostrea edulis, Spisula subtruncata, Tellina pulchella, Turritella subtruncata, Tellina albicans, Tellina pulchella Tellina albicans, Tellina subtruncata, ans, Tellina incarnata, Te incarnata, Tellina ans, na, Corbula gibba, Dentalium gibba, Corbula na, na, Corbula gibba, Dentalium gibba, Corbula na, exopecten glaber, Na exopecten glaber, ata, Tellina albic Tellina ata, Tellina pulchella Tellina Tellina incarnata Tellina sp., sp., . sp a lupinus, Fl a lupinus, chinensis, Chamelea galli Chamelea chinensis, Nucula , Glycymeris glycymeris, Mactra stulto glycymeris, Glycymeris , sp., chamasolen, Chamelea galli Chamelea chamasolen, Dosinia lupinus, Flexopecten glaber, Nassarius Nassarius glaber, Flexopecten lupinus, Dosinia tylus, Spisula subtrunc Spisula tylus, sp., D. venustus, insubrica,D. Dosinia Glycymeris lupinus, ensis, Ensis bula gibba, Dosini bula gibba, Glycymeris insubrica,Glycymeris edulis Ostrea cf. cf. Donax Donax sp., sp., na, Dosinia lupinus Dosinia na, N. gibbosulus, Nassarius musivus Nassarius gibbosulus, N. Glycimeris insubrica, Lucinella divaricata, Moerella distorta, Nassarius mutabilis, Nassarius cf. cf. sp., nella divaricata, Moerella distorta, Pholas dactylus, Spisula dactylus, Pholas distorta, Moerella divaricata, nella Arctica islandica, Calyptraea islandica, Arctica Arctica islandica, Azorinus islandica, Arctica Dosinia lupinus, Nassarius Nassarius lupinus, Dosinia sp., sp., sp., D. trunculus, Donax Donax trunculus, D. a, Chamelea gallina, Cor gallina, Chamelea a, cf. cf. N. prysmaticus, Spisula subtruncata, Tellina Tellina subtruncata, Spisula prysmaticus, N. Neverita josephinia, Pholas dac Pholas josephinia, Neverita sp. cf. cf. sp., Dentalium Chamelea gallina, Euspira Euspira gallina, Chamelea Corbula gibba, Dentalium sp., Dosinia lupinus, Flexopecten glaber, Neverita joseph Neverita glaber, Flexopecten lupinus, Dosinia sp., Dentalium gibba, Corbula sp. sp., sp., a tubercolat Arctica islandica, Chamelea galli Chamelea islandica, Arctica Parvicardium exiguum Parvicardium sp., sp., Chamelea gallina, insubrica Glycymeris Chamelea gallina, Cylichna cylindracea, Donax Donax cylindracea, Cylichna gallina, Chamelea sp., sp., paucicostata, Acanthocardi paucicostata, Aequipecten opercularis, Aporrhais Aporrhais opercularis, Aequipecten Venus pulchella, Tellina subtruncata, Spisula Chamelea gallina gallina, Chamelea tubercolata, Acanthocardia tricarinata pliorecens glaber Flexopecten lupinus, Dosinia tubercolata, Acanthocardia Acanthocardia tubercolata, Acanthocardia Acanthocardia Acanthocardia tubercolata, Chamelea gallina, Dosinia lupinus, Flexopecten glaber, Neverita josephinia, Tellina pulchella glaber Flexopecten lupinus, Dosinia gallina, Chamelea tubercolata, Acanthocardia Dentalium gibba, Corbula gallina, Chamelea tubercolata, Acanthocardia pulchella, Turritella tricarinata pliorecens, Turritella Arca opercularis, Aequipecten tubercolata, Acanthocardia Turritella tricarinata pliorecens, Venus nux Anomia opercularis, Aequipecten tubercolata, Acanthocardia tricarinata pliorecens, Venus nux Nassarius chinensis, Calyptraea islandica, Arctica Aequipecten insubrica, opercularis, Glycymeris Venus nux Dosinia lupinus, Flexopecten insubrica glaber, Glycymeris insubrica,Glycymeris Naticarius Acanthocardia aculeata, Acanthocardia tubercolata, Chamelea gallina, Dosinia lupinus, insubrica, Glycymeris Mactra stultorum Chamelea gallina, Dosinia lupinus, insubrica, Glycymeris Nassarius Chamelea gallina Acanthocardia Luci insubrica, Glycymeris cylindracea, Cylichna gallina, Chamelea Acanthocardia tubercolata, Calyptraea chinensis, Chamelea gallina, insubrica, Glycymeris Nassarius Acanthocardia tubercolata, insubrica Glycymeris insubrica,Glycymeris Spisula subtruncata Acanthocardia tubercolata, Acanthocardia Acanthocardia tubercolata, Acanthocardia Acanthocardia tubercolata, Chamelea gallina, Donax Donax gallina, Chamelea tubercolata, Acanthocardia Acanthocardia Moerella distorta, Nassarius mutabilis, Nassarius ACG244 ACG246 ACG247 ACG248 ACG249 ACG250 ACG254 ACG255 ACG256 ACG256bis ACG257 ACG258 ACG258bis ACG260 ACG261- 261bis ACG262 ACG264 ACG266 ACG267 223.80 m223.80 ACG252 224.20 m224.20 ACG253 218.60 m218.60 ACG243 m 218.80 m220.40 ACG245 m 220.50 m 221.20 m 221.40 m 221.90 223 m m223.20 ACG251 m 224.30 m 224.50 m 230.80 m 231.10 m 231.80 m 231.90 234 m m234.40 ACG259 m 234.80 m 235.70 236 m m236.40 237 m ACG263 m 239.20 241 m