‚ivist— st—li—n— di €—leontologi— e ƒtr—tigr—fi— volume IIT noF Q ppF RPIERQS xovem˜er PHIH

„ri qe‚qexy „i‚‚i ‚yƒƒi sxƒ ve‚ pe xeƒX ex y†i‚†si‡

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pihi‚sgy weƒsxs D €eyvy we‚se ‚sxevhs D he‚se €i„‚ ƒy 8 qsy†exxs ƒ ‚hs

‚e™eivedX tune PVD PHIHY —™™eptedX ƒeptem˜er PPD PHIH

uey wordsX v—te wio™eneEi—rly€lio™eneD q—rg—noD insul—r f—uE plitomeryxD r—ttomysD ƒtertomysD heinog—lerixD €—r—lutr—F xegli —nni

n—D endemismD ˜io™hronologyD ˜iogeogr—phyF 9VHD il hip—rtimento di ƒ™ienze dell— „err— di pirenzeD —™™—nto —gli —spetE

ti t—ssonomi™i @€rol—gusAD —ffront— —l™uni —spetti evolutiviD p—leogeogr—E

e˜str—™tF efter fortyye—rsfrom its dis™overyDthe historyof the fi™i e ˜io™ronologi™i ™he permettono di identifi™—re le f—une g—rg—ni™he

field surveys —nd of the rese—r™hes on the fossiliferous deposits of q—rE ™ome r—ppresent—nti di un— ˜ioprovin™i— distint— ™on forti ™—r—tteri enE

g—no is ˜rieflyreportedF demi™iF qli —nni –WH r—ppresent—no un momento di —pprofondimento su

„he q—rg—no f—un— ˜elongs to the e˜ruzzoEepuli—n €—l—eo˜ioE —spetti evolutivi di wikroti— e —ltri t—x—F el ™ontempoD un— ™—mp—gn—D in

provin™e @in™luding —lso ƒ™ontrone —nd €—len—D e˜ruzzoAF „he fossil ™ui sono ™oinvolte ™ompetenze multidis™iplin—riD port— — un— piuÁ dett—E

—ssem˜l—ge ™omprehends —ll the ™l—sses of terrestri—l verte˜r—tesX ‚epE gli—t— ™omprensione dell— str—tigr—fi— delle ™operture sediment—rie lo™—li

tili—D emphi˜i—D eves —nd w—mm—li—F „he f—un—l ™omposition is un˜—E e delle mod—lit—Á di riempimento delle fessureF „—le l—voro eÁ st—to ripreso

l—n™ed —nd presents the fe—tures of — typi™—l insul—r endemi™ f—un—F „he

re™entemente in o™™—sione delle —ttivit—Á di ri™er™— fin—lizz—te —llo studio

˜ulk of the —ssem˜l—ge is ™omposed of sm—ll m—mm—ls —nd ˜irdsF f—sed

dei depositi qu—tern—ri — ™ui ™ooper—no v—rie universit—Á it—li—neD l— ƒoE

on liter—tureD we report the most ™h—r—™teristi™ fe—turesD phylogeneti™

printendenz— e gli enti lo™—liF v9ultimo de™ennio eÁ dedi™—to —gli studi

rel—tionshipsD —nd evolution—ryrem—rks on — list of m—mm—lsD fo™using

t—ssonomi™i sui gliridiD sull9—vif—un— e sull9erpetof—un—F ƒono frequentiD

p—rti™ul—rlyon the —d—pt—tions of the most endemi™ t—x—F woreoverD —

infineD l—vori in ™ui le f—une endemi™he del q—rg—no vengono ™it—te

˜io™hronologi™—l s™heme of the m—jor f—un—l events is proposed —nd

™ome esempio p—rti™ol—rmente interess—nte per l— ™ontinuit—Á dell— doE

illustr—tedF elthough still imperfe™tD the s™heme is the first synopsis of

™ument—zione o per l9evidenz— e l— din—mi™— dei pro™essi evolutiviF

—ll the ˜io™hronologi™—l —nd t—xonomi™—l inform—tion on m—mm—ls so

v— f—un— delle „erre ‚osse ™omprende tutte le ™l—ssi di verteE

f—r —v—il—˜leF iventu—llyD the hypotheses on the origin —nd modes of

˜r—ti terrestriX ‚ettiliD enfi˜iD ™™elli e w—mmiferiF v— ™omposizione

™olonis—tion —re summ—rized —nd shortlydis™ussedF

present— tutti i ™—r—tteri tipi™i di un— f—un— endemi™— insul—reD —n™he se le

prom this overview the q—rg—no f—un— is indeed of gre—t interE

fessure piuÁ —nti™he ™ontengono t—x— non endemizz—tiF s t—x— endemi™i est for studies of evolution —nd ˜iogeogr—phyF xonethelessD there —re

vengono qui tr—tteggi—ti nelle loro linee essenzi—liD evidenzi—ndo possiE persistent un™ert—inties on its —ge —nd originD —s well —s on the —ge of

˜ili rel—zioni ™on —nten—ti ™ontinent—li e i prin™ip—li trend evolutiviF infilling of the k—rsti™ fissure system of this unique fossiliferous lo™—lityF

†iene quindi proposto per l— prim— volt— un qu—dro ˜io™ronoE

logi™oD suddiviso in f—si di popol—mentoD in ™ui si eÁ ™er™—to di riunire

‚i—ssuntoF s depositi di riempimento delle fessure ™—rsi™he nel

tutte le inform—zioni rel—tive —i m—mmiferi finor— des™ritti in q—rg—noF q—rg—no nordEo™™ident—le @„erre ‚osseA ™ontengono un— ri™™— f—un—

uesto qu—dro h— tuttor— un ™—r—ttere prelimin—re per le diffi™olt—Á endemi™—F p—une simili sono st—te ritrov—te in zone limitrofe del q—rE

in™ontr—te nell9integr—re inform—zioni t—ssonomi™he e ™ronologi™he d— g—noD m— —n™he nell— zon— di endri— @€ugli—AD — €—len— e — ƒ™ontrone

l—vori f—tti d— diversi spe™i—listi e per l— in™ompletezz— delle ™onos™enze

@e˜ruzzoAF „—li f—une endemi™he sono st—te riunite nell— p—leo˜ioproE

su —l™uni t—x— e sulle f—une di numerose fessureF vin™i— e˜ruzzoEepul—F

snfineD viene riper™ors— l— stori— delle ipotesi sull9origine delle sn questo ™ontri˜uto si present— un— ˜reve stori— delle ri™er™he

f—une dell— p—leo˜ioprovin™i— e˜ruzzoEepul—D ™on p—rti™ol—re —ttenE sul q—rg—noD ™on p—rti™ol—re riferimento —ll— ™omposizione dell— f—un—

zione —l settore g—rg—ni™oF ve ipotesi sull9et—Á e le mod—lit—Á di proveE e — ™onsider—zioni ˜io™ronologi™he e ˜iogeogr—fi™heF s primi reso™onti

nienz— sono poi ™onfront—te e dis™usseF delle ri™er™he sull— f—un— delle „erre ‚osse ris—lgono —ll9inizio degli

uest— p—nor—mi™— mostr— ™heD dopo qu—r—nt9—nni di ri™er™heD —nni –UHD qu—ndo w—tthijs preudenth—l pu˜˜li™oÁ un— not— prelimin—re

l— f—un— del q—rg—no eÁD d— un l—toD un ™—so di studio molto interess—nte sui ritrov—menti di verte˜r—ti nell9—re— g—rg—ni™—D individu—ndone il ™—E

per gli —spetti evolutivi e ˜iogeogr—fi™iD m—D d—ll9—ltroD —n™or— present— r—ttere endemi™o insul—reF

—spetti d9in™ertezz— ™he rigu—rd—no prin™ip—lmente l9et—Á e l— provenienE ƒeguironoD —d oper— di diversi —utoriD l— des™rizione delle —viE

z— dei t—x— e in se™ondo luogo —n™he l9et—Á dei depositi fossiliferi stessiF f—une e delle forme — ™—r—ttere endemi™o piuÁ m—r™—toX wikroti—D roE

I niversit—Á di €—lermoD hip—rtimento di qeologi— e qeodesi—D †i— er™hir—fi PPD sEWHIPQD €—lermoD st—lyF

iEm—ilX fm—sinidunip—FitD d—ri—peUPdunip—FitD gsurdidunip—Fit

P niversit—Á di pirenzeD hip—rtimento di ƒ™ienze dell— „err—D †i— v— €ir— RD sESHIPI pirenzeD st—lyF iEm—ilX p—olom—ri—Frin—ldidunimiFit

RPP w—sini pFD ‚in—ldi €F wFD €etruso hF 8 ƒurdi qF

sntrodu™tion ™i—ted with this f—un— r—ised ™onsider—˜le interest in the

s™ientifi™ ™ommunity@erz—rello et —lF PHHTAF

‚ed soil deposits @„erre ‚osseA fill most of the

preudenth—l @IWUIA —™tu—lly—nti™ip—ted most of

extensive p—l—eok—rst network in the wesozoi™ limeE

the results th—t were to ˜e pu˜lished on the „erre ‚osse

stone on the xorthE‡estern slopes of wount q—rg—noF

f—un—F sn the following ye—rs preudenth—l des™ri˜ed

ƒever—l fissure fillings yielded — very ri™h —mount of

some endemi™sD exhi˜iting most pe™uli—r fe—turesD su™h

fossil rem—ins of — verydiversified —ssem˜l—ge of verteE

—s heinog—lerix @preudenth—l IWUPA —nd wikroti— @preuE

˜r—tesF „hese fossils —ttest to — su™™ession of f—un—l setE

denth—l IWUTAF we—nwhileD he proposed the first ˜ioE

tlement eventsD whi™h o™™urred when the —re— w—s isoE

™hronologi™—l —rr—ngement of the fissure fillings @preuE

l—ted —nd inh—˜ited ˜yendemi™ t—x—F wost of the fossiE

denth—l IWUTAF preudenth—l9s @IWUIA influenti—l study

liferous deposits h—ve ˜een exposed ˜yintensive qu—rE triggered — flurryof studiesF

rying in — sm—ll —re—D of —˜out PH squ—re kilometresD ƒever—l monogr—phs were pu˜lished on the q—rE

in™luded ˜etween the towns of epri™en— —nd €oggio g—no f—un—X f—llm—nn @IWUQD IWUTA des™ri˜ed the ˜ird

smperi—le @pigF IAF ƒimil—r f—un—D howeverD w—s —lso rem—insD futler @IWVHA revised the genus heinog—lerixD

found out of this —re—D iFeFD —t ƒF x—zz—rio @—˜out S km ‡illemsen @IWVQA —n—lyzed the otter €—r—lutr— g—rg—E

northw—rdAD wonte qr—n—t— @PS km ƒouthEi—stAD —s well nensisD veinders @IWVRA the endemi™ rumin—nt roplitoE

—s in —n unidentified lo™—lity˜etween f—rlett— —nd enE meryxD preudenth—l @IWVSA studied the ™ri™etidsD —nd

dri— @preudenth—l IWUIAD —˜out IHH km ƒouthEi—st of h——ms 8 preudenth—l @IWVSA des™ri˜ed the gi—nt dorE

the qu—rry—re—F „he „erre ‚osse f—un—D together with mouse ƒtertomys l—ti™rest—tusF

those re™overed from ƒ™ontrone —nd €—len—Eg—po di huring the VHsD rese—r™hers of the hep—rtment of

piume @e˜ruzzoD ™entr—l st—lyAD —re ˜elieved to prove i—rth ƒ™ien™es of ploren™e @st—lyA ™ondu™ted new field

the existen™e of the soE™—lled e˜ruzzoEepuli—n €—l—eoE surveysD —nd g—ve new impulse to the studies on q—rg—E

˜ioprovin™e @™fF w—zz— 8 ‚ustioni PHHVAF „his ™ontriE no —n—lyzing the evolution of the f—un— @in p—rti™ul—r

˜ution intends to ˜e — st—teEofEtheE—rt report of the wikroti— —nd €rol—gusAD —s well —s the p—l—eogeogr—phy

knowledge on the q—rg—no f—un—F of the —re—D ˜ut they—lso proposed — new ˜io™hronoloE

gi™—l —rr—ngement of the fissure fillings @he qiuli et —lF

IWVT—D ˜D IWVU—D ™Y w—zz— IWVU—D ˜D ™Y w—zz— 8 —fonte

IWVUAF prom these studies the q—rg—no f—un— w—s —™E e ˜rief history of the rese—r™hes in q—rg—no

knowledged to ˜elong to — distin™t endemi™ ˜ioprovin™e

pield surveys were first ™ondu™ted in the epri™eE whi™h w—s ™—lled the epuloEh—lm—ti™ ‚e—lm @he qiuli

n—E€oggio smperi—le —re— from IWTW to IWUR @preuE et —lF IWVU™AF he qiuli 8 „orre @IWVRA interpreted the

denth—l persF ™ommFA ˜yw—tthijs preudenth—l @x—turE evolution—rytrends o˜served in wikroti— —nd €rol—gus

—lisD x—tion——l x—tuurhistoris™h wuseumD of veidenD —s the result of ™ompetitive interrel—tionships ˜etween

the xetherl—ndsAF sn his first reportD preudenth—l themD while „orre @IWVTA stressed the gr—du—listi™ —sE

@IWUIA identified the pe™uli—r ––gig—ntism —nd —˜err—nt pe™ts of the their evolutionF †—lleri @IWVRA g—ve — su˜E

morphologies99 of some of the t—x—D thus —™knowledging st—nti—l ™ontri˜ution proposing — €lio™ene —ge to the

the insul—r fe—tures of the „erre ‚osse f—un—F woreoverD m—rine deposits lo™—llyse—ling the fissure fillingsF „hese

preudenth—l reported the o™™urren™e of —n e—rly€leiE m—rine sediments were previouslyd—ted to the „ortoE

sto™ene @g—l—˜ri—nA f—un— ™h—r—™terized ˜y elloph—iE ni—n ˜yh9eless—ndro et —lF @IWUWAF yther studies folE

omysD whi™h h—s l—ter on ˜een ™—lled €irro xord p—un— lowedD devoted espe™i—llyto the evolution of spe™ifi™

˜yhe qiuli et —lF @IWVU—AF „he find of stone tools —ssoE t—x—D n—mely wikroti— @—fonte 8 w—sini IWWPY e˜˜—zzi

pigF I E vo™—tion of the sites of the

e˜ruzzoEepuli—n €—l—eoE

˜ioprovin™eF

ƒqu—res indi™—te lo™—lities

with „erre ‚osse f—un—D

found in fissure deposits in

the forel—nd dom—inF „ri—nE

gles indi™—te lo™—lities with

rem—ins found in ˜e—™h to

m—rine deposits in the ™h—in dom—inF

„he q—rg—no „erre ‚osse insul—r f—un—sX —n overview RPQ

et —lF IWWQY €—rr— et —lF IWWWY willienE€—rr— PHHHY willien everD the pred—tors —re f—irly™ommon —nd represented

˜y™ro™odilesD sn—kes —nd p—rti™ul—rly˜y˜irds of preyF 8 t—eger PHHIAD €rol—gus @engelone PHHSAD the inse™tiE

„he l—tter —re represented ˜ytwo endemi™ spe™ies of vores @p—nf—ni IWWWY v—n den roek ystende PHHIAD —nd

e—gle @q—rg—no—etusA —nd six spe™ies of owlD —mong roplitomeryx @v—n der qeer IWWWD PHHSD PHHVAF

whi™h three ˜—rn owl in™luding the veryl—rgeEsized we—nwhileD —t the ˜eginning of the WHsD further

„yto gig—nte—F xonEendemi™ sm—ll m—mm—ls —re —lso field surveys were ™ondu™ted jointly ˜y the universities

found in the —ssem˜l—geF „he f—un—l list of the m—mm—E of ploren™eD €is—D —nd g—merinoF „he results of these

li—n t—x—D with some t—xonomi™—l det—ils —nd evolutionE surveys were summ—rized in e˜˜—zzi et —lF @IWWTA mulE

—ryrem—rksD is reported in „—˜F IF tidis™iplin—ryreportD whi™h re™onstru™ts the evolution

of the k—rst systemD —s well —s the origin —nd str—tigr—E

indemi™ t—x—F roplitomeri™id—e —re most pe™uE phyof the whole out™ropping su™™essionD from the

li—r endemi™ rumin—ntsF „hey—re represented ˜ysever—l „erre ‚osse soils up to the €leisto™ene @g—l—˜ri—n to

spe™ies of different sizeD —t le—st two of whi™h very ‚e™entA depositsF „he studyin™ludes —lso — det—iled

sm—llF „hey—re ™h—r—™terized ˜ythe fusion of t—rs—l str—tigr—phyof the m—rine sediment—ry™over th—t loE

˜ones @n—vi™oE™u˜oid —nd met—t—rs—lAD whi™h fun™tionE ™—llyse—ls the fissuresD together with —n interpret—tion

—llylimits l—ter—l movements of the —rti™ul—tion @veinE of the p—l—eogeogr—phi™ evolution of the —re—F

ders 8 ƒond——r IWURAD suggesting lo™omotion on r—gged yver the l—st ten ye—rsD interest on the q—rg—no

terr—insF „he most rem—rk—˜le fe—tureD howeverD is the f—un— ˜ursted —g—inF helfino @PHHPA —nd helfino et —lF

o™™urren™eD —t le—st in some spe™iesD of five ™r—ni—l —pE @PHHUA studied the herpetof—un— for the first timeD while

pend—ges @one on the re—r end of the n—s—l ˜oneD —nd qohli™h 8 €—vi— @PHHVA identified — new spe™ies of ˜irdF È

two ™ouples upon e—™h or˜itA ™oupled with longD s—˜reE qliridsD twentyye—rs—fter h——ms 8 preudenth—l9s

like upper ™—nines @pigF PAF „he morphologi™—l dist—n™e @IWVSA des™ription of ƒtertomys l—ti™rest—tusD re™eived

from the other rumin—nt f—miliesD together with the o™E the —ttention of sever—l rese—r™hersD iFeFD preudenth—l 8

™urren™e of — set of di—gnosti™ fe—turesD whi™h is not w—rtõÂnEƒu—Ârez @PHHTAD ‚in—ldi @PHHTAD w—rtõÂnEƒu—Ârez 8

found in the other rumin—ntsD led veinders @IWVRA to preudenth—l @PHHUA —nd ‚in—ldi 8 w—sini @PHHWAF sn

refer these endemi™s to — new f—milyF „he phylogeneti™ these p—pers five new ƒtertomys spe™ies —re des™ri˜edF

rel—tionships of these rumin—nt —re still de˜—tedD —s well ‚in—ldi 8 w—sini @PHHWA —lso proposed — ™orrel—tion

—s the —ge of their ™ontinent—l —n™estorF e™™ording to ˜etween the fissure ˜io™hronologies proposed ˜ypreuE

w—zz— 8 ‚ustioni @PHHVAD the —n™estor might ˜e tr—™ed denth—l @IWUTA —nd he qiuli et —lF @IWVU˜AF

˜—™k in the v—te yligo™ene ˜efore the divergen™e ˜eE ƒt—rting from IWWHD the „erre ‚osse f—un— is ™onE

tween gervid—eD fovid—e —nd qir—ffid—eF yther —uthors sidered —n ex—mple of utmost interestD ˜e™—use of the

@eFgF woy—ÁEƒol—Á et —lF IWWWA suggested the possi˜ilityof ™ontinuityof its do™ument—tion —nd of the promising

—n e—rlyv—te wio™ene —n™estor @ wi™romeryxAD while potenti—ls it h—s to provide —™™ur—teD reli—˜le —nd se—mE

veinders @IWVRA suggested the i—rlywio™ene emphiE less inform—tion on evolution—rypro™essesF st is often

mos™hus —s — suit—˜le ™—ndid—te to the —n™estryof the mentioned with ƒ™ontrone in p—pers devoted to the

f—milyF iventu—llyD v—n der qeer @PHHVA —nd v—n den xeogene p—l—eogeogr—phyof the wediterr—ne—n @eFgFX

roek ystende et —lF @PHHWA —re in f—vor of — €—l—eomeE he qiuli et —lF IWWHY woy—ÁEƒol—Á et —lF IWWWY w—sini et

ry™id—e —n™estorF —lF PHHP—D ˜D PHHVY ‚ook et —lF PHHTD PHHVY de †os et —lF

heinog—lerix is —n inse™tivore ˜elonging to the PHHUY w—zz— 8 ‚ustioni PHHVAF

i™hinosori™in—eD — su˜f—milynow—d—ysrestri™ted to

ƒouthern esi—F ixt—nt i™hinosori™in—e in™lude sm—ll

to medium sized spe™ies th—t live in tropi™—l or su˜troE

„he f—un—

pi™—l forests in humid environmentsF „he su˜f—milyw—s

„he ri™h „erre ‚osse f—un— in™ludes —ll ™l—sses of widespre—d in iurope —t le—st from the yligo™ene to the

terrestri—l verte˜r—tesX ‚eptili—D emphi˜i—D eves —nd i—rly€lio™eneD p—rti™ul—rlywith represent—tives of the

w—mm—li—F st is highlyun˜—l—n™edD howeverD —nd preE tri˜e q—leri™iniF heinog—lerix is ™h—r—™terized ˜yshort

sents —ll the fe—tures of —n insul—r popul—tionF „he ˜ulk —nd strong legsD — l—rge he—d with elong—ted muzzleD —nd

of the —ssem˜l—ge is ™omposed of endemi™ mi™rom—mE highlyspe™i—lized ™hewing —pp—r—tus with strongD sh—rp

m—ls —nd ˜irdsD some of whi™h of gig—nti™ sizeF w—™roE in™isorsD stout ––third —nd fourth premol—rsD with ˜lunt

m—mm—ls —re poorlyrepresentedX theyin™lude — group ™uspsD —d—pted for ™rushing99 @futler IWVHAD —nd shortD

of rumin—nts of the endemi™ f—milyroplitomeri™id—eD strong row of mol—rsF „he re—r p—rt of the m—ndi˜le

together with sever—l spe™ies of the gi—nt inse™tivore shows — verylow ™ondyleDwhi™h suggests th—t this

heinog—lerixD whi™h is mentioned —s — l—rge m—mm—l moonr—t ™ould widelyopen —nd r—pidlysn—p its j—wsF

˜e™—use of its —nom—lous sizeF g—rnivor— —re represented sn q—rg—noD heinog—lerix is present —lre—dyin the most

only˜ythe m—rine otter €—r—lutr— g—rg—nensisF rowE —n™ient fissures with — singleD sm—llEsized —nd shortE

RPR w—sini pFD ‚in—ldi €F wFD €etruso hF 8 ƒurdi qF

„—˜F I E yver—ll t—xonomi™ ™omposition of the m—mm—li—n —ssem˜l—ges of the wikroti— f—un— @—fter he qiuli et —lF IWWH —nd w—sini et —lF

PHHVD upd—tedAF wikroti— line—ges —re num˜ered —fter he qiuli et —lF @IWVU˜AF

„he q—rg—no „erre ‚osse insul—r f—un—sX —n overview RPS

™ies with simpler dent—l morphology@ ƒF d——msiD ƒF simE

plex —nd ƒF degiuliiAF „hese groups likelyrepresent two

independent evolution—ry˜r—n™hes stemming from —

™ommonD pro˜—˜lynonEendemi™D —n™estor @w—rtõ ÂnE

ƒu—Ârez 8 preudenth—l PHHUY ‚in—ldi 8 w—sini PHHWAF

foth — sm—ll —nd — l—rgeEsized spe™ies usu—llyo™™ur

togetherD with few ex™eptionsX fissure ‚in—s™it— ID where

three sm—ll spe™ies o™™urD p—rti—llyoverl—pping in sizeD

two of whi™h of the simplerEtoothed group @ƒF d——msi

—nd ƒF simplexAD —nd fissure pQPD where onlythe sm—llE

sized ƒF degiulii o™™ursF ‚in—ldi 8 w—sini @PHHWA re™ogE

nized the o™™urren™e of five distin™t phyleti™ line—gesX

three of the ™omplexEtoothed group —nd two of the

simpler oneF ynlythe evolution of the ƒF d——msiEƒF

degiulii line—ge ™—n ˜e followed throughout the su™™esE

sion of fissuresF st ™h—nged modestlyin morphology—nd

it presents — redu™tion in size th—t is —pp—rentlyopposite

to wh—t is imposed ˜ythe soE™—lled ––isl—nd rule99F „he

—n™estryof ƒtertomys is dis™ussedX preudenth—l 8 w—rE

tõÂnEƒu—Ârez @PHHTA suggested — wessini—n wyomimus —nE pigF P E ‚e™onstru™tion of the ™r—nium of roplitomeryx m—ttheiD

™estorD where—s ‚in—ldi @PHHTA ™onsidered — most likely —n unusu—l —nterior view @from veinders IWVRAF

widdleEe—rlyv—te wio™ene —n™estor of the €eridyE

romysEwyomimus line—geF

€rol—gus is widespre—d iurope—n wio™ene o™hoE muzzled spe™ies @heinog—lerix freudenth—liAF st underE

tonid @— veryf—r rel—tive of ext—nt pik—sAD whi™h is preE went — r—di—tion whi™h produ™ed —t le—st two line—gesD

sent in q—rg—no tooF w—zz— @IWVU—D ˜D ™A shows th—t differing in size —nd in the sh—pe of the foreEmuzzleF

q—rg—no €rol—gus underwent signifi™—nt evolution—ry enother q—leri™ini spe™iesD sm—ller th—n the wioE

™h—nges in the dent—l morphologyD ™oupled with — ™ene represent—tives of the tri˜eD is very™ommon in the

m—rked in™re—se in size ™omp—red to its ™ontinent—l „erre ‚osse fissure fillingsF elso this tinygymnure

™ounterp—rtsF e lessEderived —nd sm—ller spe™ies @€rol—E shows some endemi™ fe—turesD su™h —s —n enl—rged —nd

gus —pri™eni™usA is widespre—d in —ll fissure fillingsD stout premol—r seriesF st is phylogeneti™—lly rel—ted to

where—s in the youngest fissures — l—rger —nd more deE the €—r—sorexEƒ™hizog—lerix group @p—nf—ni IWWWY w—siE

rived spe™ies @€rol—gus imperi—lisA —lso o™™ursF e™™ordE ni unpu˜lishedAF „he phylogeneti™ rel—tionships of heiE

ing to w—zz— et —lF @IWWSA q—rg—no €rol—gus derived nog—lerix —re still dis™ussedF futler @IWVHA suggested —

from €rol—gus oeningensis —nd ™olonized the ˜ioproE possi˜le deriv—tion from —n yligo™eneEi—rlywio™ene

vin™e in the widdle wio™eneF engelone @PHHUA holds — unknown esi—n —n™estorD while v—n den roek ystende

e—stern iurope—n €F oeningensisElike —n™estor for the @PHHIA proposed — widdleEv—te wio™ene €—r—sorexElike

q—rg—no endemi™D —nd suggests — v—te wio™ene @e—rly —n™estorD —ssuming — ™ommon —n™estryfor ˜oth the giE

wessini—nA ™oloniz—tionF en —rti™ul—ted skeleton tent—E g—nti™ heinog—lerix —nd the verysm—llEsized gymnureF

tivelyreferred to €rol—gus ™fF —pri™eni™us h—s ˜een reE elso the —d—pt—tion of heinog—lerix is dis™ussedF preuE

™overed in e—rlywessini—n l—goon sediments in €—len—E denth—l @IWUPA reg—rded this moonr—t —s — s™—vengerD

g—po di piume @w—zz— et —lF IWWSAF where—sD —™™ording to the el—˜or—ted —rgument—tion of

wikroti— is —n u˜iquitous murid of the „erre futler @IWVHAD it ™ould h—ve ˜een —n —™tive pred—torD

‚osse fissure fillingsD —nd perh—ps the most ™h—r—™terisE possi˜lyspe™i—lized in hunting fishes —s well —s ™rust—E

ti™ genus of the —ssem˜l—geF „he endemi™ evolution of ™e—nsF

this murid is ™h—r—™terized ˜y— verym—rked growth in ƒtertomys is —n endemi™ dormouse of the su˜f—mE

sizeD —™™omp—nied ˜y—n in™re—se in hypsodonty—nd ilywyomimin—eDwith dent—l morphology™ompli™—ted

™omplexityof the dent—l ™rownsF ƒt—rting from the ˜—si™ ˜y—n in™re—sed num˜er of ™rests on the o™™lus—l surf—™e

murid p—tternD —s ™rown height in™re—ses tu˜er™les unite of the teeth @eFgF five to fourteen ™rests in the upper

forming tr—nsvers—l ™rests in lower mol—rs —nd in the mol—rsAF „he genus in™ludes six spe™ies whi™h ™—n ˜e

upper third mol—rD while the ™ir™ul—r steph—nodont p—tE —rr—nged in two different groupsF „he first in™ludes

tern of the first —nd se™ond upper mol—rs is —™™entu—tedF l—rgeE to veryl—rgeEsized spe™ies with ™omplex to very

„he most signifi™—nt fe—ture in the dent—l evolution of ™omplex dent—l morphology@ ƒF d—uniusD ƒF lyrifer —nd ƒF

wikroti—D howeverD is the in™re—se of ™rown ™omplexityD l—ti™rest—tus E the l—tter m—y˜e the l—rgest glirid so f—r

knownAD while the se™ond in™ludes mediumEsized speE resulting from the —ddition of up to five new tr—nsvers—l

RPT w—sini pFD ‚in—ldi €F wFD €etruso hF 8 ƒurdi qF

™rests in the first lower —nd in the third upper mol—rsF th—t develop on the l—˜i—l ˜order of the mol—rsD —s well

„he —dv—n™ed spe™ies from ƒ—n qiov—nnino possess the —s the ––™rests th—t tend to ˜e™—me domin—ting on the

most highlymodified teethF rypsodontyDtr—nsvers—l ™usps99D whi™h give the teeth — ™—rvedEin —spe™tF epp—rE

Q

™rests —nd in™re—sing ™omplexityof the w —nd w —re entlyD this endemi™ evolved —long — single phyleti™ lineE I

™h—r—™ters sh—red —lso ˜ywi™rotin—e rodentsX for this —ge @rF ˜eetsiErF n—z—riiErF g—rg—ntu—AF

re—son preudenth—l @IWUTA ™oined the origin—l n—me wiE preudenth—l @IWVSA re™ognises — ™ert—in —ffinityof

™roti—F nfortun—telyD the n—me h—d to ˜e ™h—nged in r—ttomys with gri™etulodon @xeo™ri™etodon in preuE

––wikroti—99 ˜ypreudenth—l PHHTD sin™e it w—s preEo™™uE denth—l 8 w—rtõÂnEƒu—Ârez PHIHAD ˜ut the third mol—rs

pied ˜y— vepidopter— genusF „he m—™ros™opi™ morphoE th—t persist to ˜e l—rge in the endemi™ genus rules out

logi™—l ™h—nges in the teeth —re —™™omp—nied ˜ymodE — dire™t —n™estryF ‚e™entlyD preudenth—l 8 w—rtõÂnEƒu—ÂrE

ifi™—tions in the p—ttern of en—mel mi™rostru™tureX proE ez @PHIHA proposed — possi˜le —n™estryfrom gri™etus

gressive in™re—se in prism in™lin—tion ™—uses — swit™h @epo™ri™etus in preudenth—l 8 w—rtõÂnEƒu—Ârez PHIHAD

from the primitive r—di—l p—ttern to protoEt—ngenti—l whi™h is still —nother genus present in the q—rg—no —sE

˜—nds @—fonte 8 w—sini IWWPAF foth morphologi™—l sem˜l—geF

—nd mi™rostru™tur—l ™h—nges suggest th—t the dentition

xonEendemi™ t—x—F „he „erre ‚osse —ssem˜l—ge of wikroti— ˜e™ome highlyspe™i—lized for —n —˜r—sive

in™ludes —lso nonEendemi™ t—x—F ‡ith the ex™eption of dietF „his spe™i—liz—tion ™ould ˜e linked to the —™quisiE

epodemusD they—ll o™™ur onlyin the older fissuresF tion of — fossori—l h—˜it @€—rr— et —lF IWWWAF ix™—v—tingD

hryomys —pulus is s™—r™elyrepresented onlyin livingD —nd possi˜lyfeedingD in ˜urrows likelyfor™ed

three fissures @fi—n™one ID ‚in—s™it— I —nd pISAF „his these murids to grind high per™ent—ges of ™o—rse items

dormouse h—d ˜een identified —s iliomys in e—rlier reE @eFgFD soil p—rti™lesAF wikroti— underwent sever—l spe™i—E

portsY then it w—s given its ™orre™t t—xonomi™ des™ripE tion events produ™ing — r—ther ™omplex ˜r—n™hing phyE

tion ˜ypreudenth—l 8 w—rtõ ÂnEƒu—Ârez @PHHTAF „hese logenyD of whi™h —t le—st five line—ges —re ˜etter known

—uthors in™luded it in hryomys on the ˜—sis of the morE @pigF QAF €—r—llel morphologi™—l evolution o™™urred in

phologyof the lower mol—rsD though —dmittingD howE the different line—gesD —lthough the m—jor phylogeneti™

everD th—t iliomys —nd hryomys —re quite indistinguishE trend ™h—r—™terizes the soE™—lled ––resident line—ge99F sn

—˜le dent—llyF foth gener— —re known sin™e the widdle those th—t —re ™onsidered the most —n™ient fissuresD wikE

wio™ene @h——ms 8 de frujin IWWSAY iliomys w—s more roti— shows simpler morphologyD ™lose to the ˜—si™ murE

™ommon in iurope —s opposed to hryomys whi™h w—s id p—tternD —lthough —lre—dywith — ™ert—in degree of

widespre—d in xorth efri™— —nd xe—r i—stF hypsodonty —nd l—rger sizeF sn younger fissures two

xeo™ri™etodon @gri™etulodon in preudenth—l IWVSA nonEresident spe™iesD whi™h likelyevolved in some

neigh˜ouring isl—ndsD re—™hed the q—rg—no isl—nd ˜y is known from three fissures @fi—n™one ID ‚in—s™it— I

—nd „refossi IAD where—s epo™ri™etus @gri™etus in preuE dispers—lD envis—ging the soE™—lled ––—r™hipel—go effe™t99

denth—l IWVSA o™™urs onlyin ‚in—s™it— IF gri™etids of @preudenth—l IWUIY he qiuli et —lF IWVT—AF „he morphoE

logi™—l evolution of wikroti— is strikingD to — point th—tD the he qiuli ™olle™tion h—ve re™ently˜een ™he™ked ˜y

endre— ƒ—vorelli @work in progressAF e™™ording to if it h—d developed in nonEinsul—r ™ontextD it would h—ve

preudenth—l @IWVSAD ˜oth those ™ri™etids —re simil—r to justified the ™re—tion of — thoroughlydistin™t supr—E

generi™ t—xonD iFeFD — tri˜eF „he forerunner of wikroti— their ™ongeneri™s from the „uroli—n f—un— of grevillente

@de fruijn et —lF IWUSAD where —lso iliomys is presentF is somewh—t un™ert—inF preudenth—l @IWUTA suggested —

weg—™ri™etodon is yet —nother ™ri™etidD repreE possi˜le —n™estryfrom ƒteph—nomys @see —lso e˜˜—zzi

et —lF IWWQAF preudenth—l 8 w—rtõÂnEƒu—Ârez @IWWWA sugE sented ˜yonlyone spe™imen in ‚in—s™it— ID whi™h

preudenth—l @IWVSA —nd preudenth—l 8 w—rtõÂnEƒu—Ârez gested th—t epodemus ™ould ˜e ™onsidered — suit—˜le

—n™estorD tooF „his opinion is sust—ined ˜ypreudenth—l @PHIHA ™onsidered due to ™ont—min—tionF weg—™ri™etoE

8 w—rtõÂnEƒu—Ârez @PHIHA ™onsidering the o™™urren™e of don o™™urred in iurope from the middle yrle—ni—n

@wxRA to the e—rly†—llesi—n @wxWAF —n unp—ired tu˜er™le in the —nterior p—rt of the w of I

epodemus —nd wikroti— —nd th—t in the l—tter genus ––—ll v—rtetium ™fF dehmi h—s ˜een retrieved in fissures

su™™essive —dditions of ™rests p—ss through — st—ge with pIS —nd pPI—E™ @p—nf—ni IWWWAY its o™™urren™e in the

veiden ™olle™tion h—s not ˜een ™he™kedF „his shrew is su™h —n —nterior ™usp99F

r—ttomys is —n endemi™ h—mster of the „erre known in iurope from i—rlyto widdle wio™eneF

‚osse fissure fillingsD l—™king onlyin the youngerfisE „he murid epodemus is the most widespre—d

nonEendemi™ t—xon —t q—rg—noF nlike the other ™onE sures of ph—ses Q —nd R @see ˜elowAF st shows — m—rked

endemi™ tenden™ytow—rd ever l—rger sizesD —™™omp—E tinent—l t—x—D whi™h h—d gone extin™t pre™o™iouslyD epoE

nied ˜y— progressive ™ompli™—tion of the —nterior p—rt demus ˜e™—me residentD —nd underwent evolution—ry

of the first mol—rsF e™™ording to preudenth—l @IWVSAD modifi™—tions th—tD —™™ording to he qiuli et —lF

other ™h—r—™teristi™ fe—tures of this genus —re the fl—nges @IWVU˜D IWWHAD —re quite signifi™—nt in the l—st ph—se of

„he q—rg—no „erre ‚osse insul—r f—un—sX —n overview RPU

pigF Q E ƒ™heme showing the evolution of the dent—l morphology—nd en—mel mi™rostru™ture of the first lower mol—r of wikroti— @—fter

e˜˜—zzi et —lF IWWQ modifiedAF

IX dentineY PX r—di—l en—mel with —xi—llyin™lined prismsY QX r—di—l en—mel with prisms norm—l to the o™™lus—l surf—™eY RX r—di—l en—mel

with —˜E—xi—llyin™lined prismsY SX primitive t—ngenti—l en—mel st—ge eY TX primitive t—ngenti—l en—mel st—ge fY UX t—ngenti—l en—melF

possi˜le fissures on the ˜—sis of the evolution of ˜oth f—un—l settlement @fissure pQPD pigF RAF st h—s never ˜een

wikroti— —nd r—ttomysD —ssuming —n irreversi˜le inE —n—lyzed in det—ilD —lthough ˜oth preudenth—l @IWVSA

™re—se of size of the wikroti— m—in line—geF he qiuli —nd he qiuli et —lF @IWVU˜A re™ognized —dv—n™ed fe—E

et —lF @IWVU˜AD in ™ontr—stD sele™ted the ri™hest s—mplesD tures re™—lling those of the l—te „uroli—nEe—rly‚us™ini—n

pinpointing — su™™ession of f—un—l settlement pulsesD m—inl—nd ™ounterp—rtsF wore det—ils on the —n™estryof

˜—sed prim—rilyon the evolution of wikroti— —nd €roE this mouse —re reported ˜elowF

l—gusD —nd sele™ting the most p—rsimonious —mong the

—ltern—tive solutions @see ‚in—ldi 8 w—sini PHHW for —

det—iled dis™ussionAF „he ˜io™hronologi™—l sequen™es fio™hronology —nd outline of the ph—ses of f—un—l setE

o˜t—ined using these two methods —re f—irly™omp—rE tlement

—˜leD with minor in™onsisten™iesF ‚in—ldi 8 w—sini

fe™—use the deposits from the different fissures @PHHWA tried to integr—te he qiuli et —lF9s @IWVU˜A s—mples

—re not ordered str—tigr—phi™—llyD rese—r™hers h—d to seE with some of preudenth—l9s @IWUTAD o˜t—ining — sumE

quen™e them ˜io™hronologi™—llyon —n evolution—ry˜—E m—rysu™™ession whi™h in™ludes —lso d—t— on the evoluE

sisF „his w—s possi˜le ˜e™—use the f—un—l ™omposition tion of ƒtertomysF „hey—lso extensivelydis™ussed the

—nd endemi™ fe—tures of the t—x— from the different fisE diffi™ulties theyh—d ™ome up with tryingto ™om˜ine

sures v—ryin — f—irly™oherent w—yF the two ˜io™hronologi™ solutionsF yne for —llD preuE

preudenth—l @IWUTA —nd he qiuli et —lF @IWVU˜A denth—l @IWUTA h—d not ™onsidered the evolution of €roE

followed two slightlydifferent ˜io™hronologi™—l str—teE l—gusD —nd he qiuli et —lF @IWVU˜A h—d not in™luded th—t

giesF „he former tried to —rr—nge ™hronologi™—lly—ll of ™ri™etidsF sn the present ™ontri˜ution upd—ted d—t— on

RPV w—sini pFD ‚in—ldi €F wFD €etruso hF 8 ƒurdi qF

€h—se I fissures ™ont—in

v—ried f—un—l —ssem˜l—ges not

e—syto group in su˜Eph—sesD

espe™i—lly˜e™—use of the in™oE

herent f—un—l ™omposition of fisE

sure ‚in—s™it— I @see preudenth—l

IWUTD IWVS —nd ‚in—ldi 8 w—sini

PHHW for — dis™ussion on this fisE

sureAF st is ™h—r—™terized ˜ythe

s™—ttered o™™urren™e of nonEenE

demi™D ™ontinent—l t—x—D —s well

—s ˜ynot too endemi™ sm—ll

m—mm—lsD su™h —s wikroti— p—rE

v—D wikroti— evolution—rylineE

—ge ID r—ttomys ˜eetsiD et™F „his

ph—se is —lso ™h—r—™terized ˜y—

primitive heinog—lerixD where—s

inform—tion on roplitomeryx is

s™—nt or not —v—il—˜leF e l—rger

—nd more —dv—n™ed heinog—lerix

o™™urs in the l—te p—rt of this pigF R E „ent—tive distri˜ution of the m—mm—li—n —ssem˜l—ge in the „erre ‚osse depositsF

ph—seF „his ph—se of f—un—l setE „he ™hronologi™—l order derived from he qiuli et —lF @IWVU˜A integr—ted ˜ysever—l

tlement foresh—dows the o™™urE sour™es of d—t—F ƒh—ded rows indi™—te the distri˜ution of the t—x—Y ˆX o™™urren™e reported

in the liter—tureY oX o™™urren™e not reported in the liter—tureD —nd ˜—sed on unpu˜lished ren™e of f—un—l —rriv—ls from the

d—t—Y cX dou˜tful o™™urren™e reported in the liter—tureY @cAX dou˜tful identifi™—tionY no

m—inl—nd —nd possi˜lyf—un—l exE

sym˜ol me—ns inferred o™™urren™eY bX phyleti™ tr—nsition within —n evolution—ry line—geF

™h—nges with the neigh˜ouring

fseID fi—n™one IY ‚sxID ‚in—s™it— IY „pID „refossi IY gQeD g—nt—tore QeY pxhD pin—

isl—nds of the p—l—eo—r™hipel—goD hY xRD x—zz—rio RY €IfD €osti™™hi— IfY ƒqD ƒ—n qiov—nninoF

fro—d ™olumns represent groups of fissuresX pPI—E™ a pPI— to pPI™Y xREpI a x—zz—rio RD ˜ut it —lso do™uments evolutionE

fi—n™one PD ghiroÁ QD ghiroÁ UeD €izzi™oli RD ghiroÁ IIfD ghiroÁ Se —nd pIY pW a pWD wonte —rytrends in the resident t—x—F

qr—n—t— ID ghiroÁ TD ghiroÁ IReD ghiroÁ IPD ghiroÁ IHeEfD ghiroÁ PƒExD ghiroÁ WD pin— r

w—sini et —lF @PHHVA ˜elieve th—t

—nd ghiroÁ PUY €If a €osti™™hi— IfD qerv—sio ID ghiroÁ RD ghiroÁ PHeEiD ghiroÁ PR —nd

—ll ––™ontinent—l99 t—x—D ˜ut epoE

qerv—sio PY ƒq a ƒ—n qiov—nnino —nd €irro IIeF wikroti— line—ges —re num˜ered —fter

demusD ˜e™—me extin™t ˜e™—use he qiuli et —lF @IWVU˜AF

of their in—˜ilityto —d—pt to the

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the med—l of the ––—dv—nt—ge of the resident99F ™ri™etids of the he qiuli ™olle™tion —re ™onsidered @ƒ—E

he qiuli et —lF9s @IWVU˜A €h—se P ™—n ˜e ™onsidered vorelliD work in progressAF

tr—nsition—l from the oldest fissures to the more re™ent pigure R is —n —ttempt to ™om˜ine ˜oth sequen™es

ones —nd it h—s ˜een —rti™ul—ted in two su˜Eph—sesF ‚eE into — single ™hronologi™—l s™hemeD ˜—sed on the highE

sident t—x— underwent phyleti™ evolution—ry ™h—ngesD est possi˜le num˜er of m—mm—li—n t—x—F st o˜viously

su™h —s those o˜served in r—ttomysD wikroti— —nd the founds m—inlyon wikroti—D ƒtertomysD r—ttomys —nd

sm—llEsized ƒtertomysF „he —rriv—l of endemi™s from €rol—gusD ˜ut it —lso in™ludes roplitomeryx —nd heinoE

neigh˜ouring isl—ndsD su™h —s ƒtertomys l—ti™rest—tusD g—lerixF xot —ll fissures reported in the liter—ture ™—n ˜e

™—nnot ˜e ex™ludedF sn this ph—seD —pp—rentlythree difE fit into the sequen™eX we h—ve in™luded onlythe ˜est

ferent sizes of roplitomeryx @v—n der qeer PHHSD PHHVA ™ontrolled ones whi™h yielded the ri™hest —nd most

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€h—se Q is the ˜est do™umented oneF st is ™h—r—™E l—r —ssem˜l—ges ™ont—ining t—x— —t simil—r evolution—ry

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phyleti™ ™h—nges in the l—te p—rt of this ph—seD eFgF the 8 e˜˜—zzi PHHT —nd €etruso et —lF PHHV for — dis™usE

sm—llEsized ƒtertomys —nd the resident line—ge of wiE sion on this prin™ipleAF sn order to m—n—ge the highest

kroti—F et the s—me time ™ri™etids ˜e™—me extin™t —nd possi˜le —mount of ˜io™hronologi™—l inform—tionD he

€rol—gus imperi—lis —ppe—rsD either ˜ydispers—l or ˜yin qiuli et —lF9s @IWVU˜A four ph—ses of f—un—l settlement

situ evolutionF „he most re™ent p—rt of the ph—se re™ords h—ve ˜een —rti™ul—ted in su˜Eph—sesD following w—sini

et —lF @PHHVAF —lso the phyleti™ tr—nsition of heinog—lerix minor into

„he q—rg—no „erre ‚osse insul—r f—un—sX —n overview RPW

hF ˜revirostris —nd of hF intermedius into hF koenigsE ph—se —s — suit—˜le ™ondition for the dispers—ls onto

the q—rg—no isl—ndsF ‚e™entlyD following the ™l—ims of w—ldiX ˜oth ™—ses exhi˜it size in™re—seF p to four difE

v—n den roek ystende et —lF @PHHWA on the ƒ™ontrone ferent sized roplitomeryx t—x— o™™urF

f—un— @see ˜elowAD preudenth—l 8 w—rtõÂnEƒu—Ârez @PHIHA €h—se R is do™umentedD to d—teD only˜yfissure

—dmitted th—t the most suit—˜le me™h—nism to expl—in pQP @he qiuli et —lF IWVU˜AF „he sm—ll m—mm—l —ssemE

the un˜—l—n™ed f—un— of the q—rg—no is sweepst—ke disE ˜l—ge is poorlydiversifiedF v—rgerEsized wikroti— —nd

pers—l whi™hD howeverD should h—d o™™urred during the ƒtertomys spe™ies ˜e™—me extin™tD where—s sm—llerE

f—vour—˜le lowEst—nd ™onditions of the wessini—nF preuE sized onesD —long with epodemus —nd q—lerix @€—r—E

denth—l 8 w—rtõÂnEƒu—Ârez @PHHTD PHIHA refused the hyE sorexAD underwent phyleti™ modifi™—tion —nd slightly

pothesis th—t q—rg—no ™ould h—ve ˜een re—™hed ˜ysu™E in™re—sed in sizeF €rol—gus still o™™urs with two differE

™essive w—ves of migr—tionD insisting th—t m—mm—ls ent spe™iesF foth roplitomeryx —nd heinog—lerix —re

––m—y h—ve re—™hed q—rg—no simult—neously99 @preuE presentD ˜ut theyh—ve not ˜een studied in the det—ilF

denth—l 8 w—rtõÂnEƒu—Ârez PHHWAD de f—™to ™ontr—di™ting „he drop in diversityof the sm—ll m—mm—ls h—s ˜een

the sweepst—ke me™h—nismF interpreted —s — sign—l of — trophi™ ™risisD possi˜lydue

futler @IWVHA —rgued th—t the un˜—l—n™ed ™ompoE to the shrinking of the isl—nd @he qiuli et —lF IWVU˜D

sition of the q—rg—no f—un— ™ould ˜e expl—ined ˜y IWWHY w—sini et —lF PHHVAF

sweepst—kesD ˜ut he ™on™luded th—t this me™h—nism

w—s not ex™lusiveD sin™e it would not —™™ount for the

o™™urren™e of emphi˜i—D whi™h ™—nnot ™ross exp—nses „he origin —nd —ge of the f—un—

of s—lt se— w—terF „o resolve this in™onsisten™yD he interE

„heoriesD modelsD —nd rel—ted —ssumptions on the preted the q—rg—no f—un— —s ˜eing ™omposed of — ™ore

me™h—nisms —nd pro™esses th—t origin—te the isl—nd f—uE of —n ––impoverished reli™ of —n older f—un—99 @—™tu—lly—

n—sD —re p—rti™ul—rlyrelev—nt ± —nd intim—telyt—ngled ± vi™—ri—n™e pro™essAD to whi™h new f—un—l elements were

to the question of the —ge of the f—un—sF e™tu—llyD the —dded ˜yl—ter inv—sions @— dispers—list pro™essAF futler

deposition of the fossiliferous sedimentsD together with w—sD indeedD the first who ™onsidered the q—rg—no f—un—

the ™olonis—tion events ˜ythe —n™estors of the insul—r —s the result of different events s™—ttered in time —nd

t—x— might h—ve ˜een s™—ttered in timeF „he two quesE with different modesF

tions should therefore ˜e ex—mined sep—r—telyD even he qiuli et —lF @IWVU™A —lso ™onsidered the q—rg—E

though they—re often dis™ussed jointlyin the liter—E no f—un— —s the result of different dispers—l events s™—tE

tureF „he different opinions of the —uthors on the oriE tered through timeF „he events th—t ˜rought the —n™esE

gin of insul—r t—x— refle™tD in p—rtD two ™ontr—sting theE tors of €rol—gusD heinog—lerixD roplitomeryx —nd ƒterE

oreti™—l —ppro—™hesD vi™—ri—n™e —nd dispers—lismF nE tomys into the insul—r dom—inD were ˜elieved to h—ve

fortun—telyD there is — ™ert—in —mount of ™onfusion in o™™urred in notE˜etter defined moments during the

the use of these ™on™eptsD —s stressed —lso ˜yw—sini et i—rlywio™eneEe—rlywiddle wio™eneF e se™ond w—ve

—lF @PHHP˜D PHHVD PHHWAF of dispers—lsD involving the —n™estors of hryomysD wiE

preudenth—l @IWUIA suggested th—t the q—rg—no kroti— —nd epo™ri™etusD w—s —ssumed to h—ve o™™urred

w—s popul—ted vi— dispers—ls from the m—inl—nd through during the regressive ph—se of the l—te wessini—nEe—rliest

— sort of ™ontinent—l ˜ridgeD possi˜ly–– — series of isl—nds —n™le—nF „he dispers—l of epodemus is ™onsidered ˜y

whi™h merely offered — limited p—ss—ge for the f—un—99 the —uthors —s — keyto define the —ge of the „erre ‚osse

@filtering ™orridorAF „he —re— of origin of the f—un— ™ould fissuresF sn — first re™onstru™tionD epodemus is —ssumed

h—ve ˜een either the emerged epennine ™h—inD —™ross to h—ve re—™hed the isl—nds together with the other nonE

the poggi— qr—˜enD or the f—lk—nsD through — threshold endemi™ rodents @hryomysD et™FA during the wessini—nE

—™ross the edri—ti™ ƒe—F re first ˜elieved th—t migr—tions i—rly€lio™eneF en —ltern—tive possi˜ility@s hypothesis

o™™urred in the l—te †—llesi—nEe—rly„uroli—nD whi™h he in he qiuli et —lF IWVU™A is ˜—sed on the morphologi™—l

™onsidered older th—n the ƒerr—v—lli—n on the ˜—sis of —ffinityof q—rg—no epodemus with epodemus domiE

in™orre™t d—ting of the ™—l™—renites th—t se—l the „erre n—ns from gs—rnot— PD —nd envis—ges — dispers—l of this

‚osse depositsF e l—te †—llesi—nEe—rly„uroli—n —ge of single murine —t the very˜eginning of the €i—™enti—nD

™oloniz—tion w—s —lso supposed ˜ywoy— ÁEƒol—Á et —lF during — m—rine regression do™umented lo™—llyD —s well

@IWWWA ˜—sed on preudenth—l @IWUID IWUTAF v—ter onD —s elsewhere in the wediterr—ne—n ˜—sinF

preudenth—l @IWVSA —ssumed the s—me mode of ™oloniE e™™ording to the l—tter interpret—tionD the fossiliE

z—tionD ˜ut —t — different timeF re re™ognized — gener—l ferous fissures together with the whole historyof the

—ffinityof the q—rg—no f—un— with those of some wesE endemi™ f—un— would h—ve ˜een ™on™entr—ted in the

sini—n lo™—litiesD su™h —s grevillente @preudenth—l IWVSAD €i—™enti—nF pollowing preudenth—l @IWUIAD he qiuli et

—lF @IWVU™A pinpointed the m—jor se— level regressionsD il erquilloD w—r—men— @preudenth—l 8 w—rtõÂnEƒu—Ârez

—nd indi™—ted them —s the most suit—˜le o™™—sions for PHHTAD —nd he ™onsidered the wessini—n low se—Elevel

RQH w—sini pFD ‚in—ldi €F wFD €etruso hF 8 ƒurdi qF

ing isl—ndsA whi™h o™™urred in two time interv—ls when dispersion onto q—rg—noD when the soE™—lled ––wid

the ˜—rriers sep—r—ting the isl—nds from the m—inl—nd edri—ti™ ‚idge99 @— su˜m—rine stru™tur—l high —™ross

were e—sier to ˜e ™rossedF „he older period is not ™le—rly the edri—ti™ ƒe—A emergedF he qiuli et —lF @IWVU™A did

definedD ˜e™—use it would sp—n from the l—te widdle not spe™ifyexpli™itlywhether dispers—ls o™™urred vi—

wio™ene to the v—te wio™eneD involving the whole €—E l—nd ˜ridgesD filtering ˜—rriersD or sweepst—kes ™on™enE

l—eo˜ioprovin™e @˜oth the e˜ruzzo —nd epuli— se™torsAF tr—ted during the lowEst—nd interv—lsF „heywere the

„he se™ond ph—se is —ssumed to h—ve o™™urred in the firstD howeverD who ™onsidered the q—rg—no f—un—s

l—te wessini—nD involving onlythe reli™ q—rg—no —re—D —s —nd isl—nds within — wider p—l—eogeogr—phi™—l ™ontextD

suggested ˜yhe qiuli et —lF @IWVU™AF ƒweepst—kes —re —ttempting — synthesis of m—rine str—tigr—phyD te™toni™sD

preferred respe™t to full ™onne™tion ˜yl—nd ˜ridgesD p—l—eogeogr—phyof l—ndsD —nd terrestri—l m—mm—l disE

sin™e theypermit filtered —rriv—ls of sm—ll m—mm—lsF tri˜utionF

woreoverD in the opinion of the —uthorsD full ™onne™E „he dis™overyof the ƒ™ontrone fossil f—un— @‚usE

tions with the m—inl—nd —re to ˜e ex™luded sin™e they tioni et —lF IWWPA h—d — deep imp—™t on the dis™ussions

would likelyh—ve ™—used the extin™tion of the endemi™ on the origin of the q—rg—no f—un—F ƒ™ontrone is lo™—ted

f—un— @see the l—te €leisto™ene of ƒi™ilyD —s —n ex—mpleAF in the epennine ™h—in —˜out IIS km ‡ of q—rg—no @pigF

w—sini et —lF @PHHP˜A introdu™ed the ™on™ept of ––polyE IAF st yielded — f—un— ™omposed of l—rgeEsized verteE

ph—si™ popul—tion99 to indi™—te f—un—l settlements deE ˜r—tes found in ™o—st—l sediments whi™h were d—ted

rived ˜y˜ioevents s™—ttered in timeD whi™h in turn ˜ythe —uthors to the e—rly„ortoni—nF „he o™™urren™e

m—yinvolve different me™h—nisms @vi™—ri—n™e or disE of roplitomeryx —nd heinog—lerixD —lthough exhi˜iting

pers—listAF €olyph—si™ popul—tions —re very ™ommon in more —r™h—i™ fe—turesD testifies the stri™t —ffinitywith the

the wediterr—ne—n —re— @eFgFD €lioE€liesto™ene of ƒi™ily q—rg—no f—un— @w—zz— 8 ‚ustioni IWWTD PHHVAF w—zz—

—nd ƒ—rdini—AF woreoverD w—sini et —lF @PHHP˜D PHHVD et —lF @IWWSA —nd w—zz— 8 ‚ustioni @IWWTA suggested

PHHWA rem—rked the diffi™ulties in oper—ting — ™le—rE™ut th—t the ™oloniz—tion of the e˜ruzzoEepuli—n —re—

distin™tion ˜etween vi™—ri—n™e —nd dispers—list me™h—nE would h—ve o™™urred during two interv—lsD v—te yligoE

isms when — det—iled re™onstru™tion is requiredD stresE ™eneEi—rlywio™ene —nd wessini—nD in ™onjun™tion with

sing th—tD for inst—n™eD — tempor—ryl—nd ˜ridge @— pe™uE m—jor m—rine regressionsF huring the first interv—l the

li—r ™—se of vi™—ri—n™eA or — filtering ˜—rrier @dispers—listA —n™estor of roplitomeryx @—nd heinog—lerixD —™™ording

do not h—ve e—sydistinguish—˜le effe™ts on the f—un— of to w—zz— 8 ‚ustioni PHHVA would h—ve re—™hed the

the isl—ndsF insul—r dom—inD while in the se™ond interv—l most of

‚e™entlyD w—zz— 8 ‚ustioni @PHHVA re™onsidered the sm—ll m—mm—ls typi™—l of the „erre ‚osse f—un—

the question of the e˜ruzzoEepuli—n fioprovin™e in would h—ve re—™hed the q—rg—no —re— from the m—inE

the light of the dis™overyof heinog—lerix —t ƒ™ontroneD l—nd @w—zz— et —lF IWWSAF

—nd of the very™—reful onEshore —nd offEshore geologiE e˜˜—zzi et —lF @IWWTA ™onsidered th—t q—rg—no

™—l investig—tions in the e˜ruzzoEepuli—n —re— —nd ––w—s wh—t rem—ined of — ˜ioprovin™e th—t w—sD —t le—st

edri—ti™ ƒe— ˜y€—t—™™— et —lF @PHHV—D ˜AF „he i—rly„orE in the v—te wio™eneD ™onsider—˜ly l—rger99D of whi™h

toni—n —ge of the ƒ™ontrone f—un— is ™onfirmed ˜y€—E ƒ™ontrone represents —n older re™ordF fy—nd l—rgeD they

t—™™— et —lF @PHHV—A who referred the fossiliferous l—yer ™onsidered the q—rg—no f—un— —s the reli™ of —n older

to —n —ge slightlyyoungerth—n the pirst ‚egul—r y™E f—un—l settlementD resulting from multiple dispers—l

™urren™e of xeoglo˜oqu—drin— —™ost—ensisD some IHFS eventsF ‚ook et —lF @PHHTA interpreted the q—rg—no —re—

w—D even though some un™ert—intystill persistsD —s —s — r—ther st—˜le stru™tur—l highD ˜eing p—rt of the foreE

stressed ˜ypreudenth—l 8 w—rtõÂnEƒu—Ârez @PHIHAF l—nd of the epennine orogeneD while ƒ™ontrone w—s

€—l—eogeogr—phi™—l interpret—tions ˜yw—zz— 8 involved in the te™toni™ disruption linked to the edifi™—E

‚ustioni @PHHVA —nd €—t—™™— et —lF @PHHV˜A ™onfirmed tion of the ™h—inF „his interpret—tion is fully™onsistent

—lso the o™™urren™e of two m—jor regressions during with the vi™—ri—n™e modelF

the v—te yligo™eneEi—rlywio™ene —nd the wessini—nD w—sini et —lF @PHHP˜D PHHVA —ppro—™hed the quesE

—nd permitted to identify— stru™tur—l high @gentr—lE tion of the origin of the q—rg—no ™ommunities distinE

edri—ti™ fridge ac wid edri—ti™ ‚idge of he qiuli et guishing t—x— —s stronglyendemi™D intermedi—teGmoderE

—lF IWVU™A whi™h w—s likelyf—irlyemerged during the —telyendemi™ —nd nonEendemi™F

m—rine lowEst—ndsD ™onne™ting h—lm—ti— —nd the „he first ™—tegoryin™ludes roplitomeryx —nd

heinog—lerixF „hese two t—x—D following previous interE e˜ruzzoEepuli—n —re— ––either in the form of stripe of

pret—tionsD —re ™onsidered —s survivors from the older l—nd or —s — ™h—in of isl—nds99 @w—zz— et —lF PHHWAF w—zz—

ph—se of f—un—l settlement of the e˜ruzzoEepuli—n €—E 8 ‚ustioni @PHHVA —dopted the model of l—nd ˜ridgesD to

l—eo˜ioprovin™eF por intermedi—teGmoder—telyendemi™ justifythe —rriv—ls of m—mm—ls —ndD —s in their previous

—s well —s for nonEendemi™ f—un—sD w—sini et —lF sugE p—perD theysust—ined two migr—tion w—vesX the older

gested — sort of sweepst—ke me™h—nism @r—fting or flo—tE oneD —llowing the —rriv—l of the —n™estors of roplitomeE

„he q—rg—no „erre ‚osse insul—r f—un—sX —n overview RQI

ryx —nd heinog—lerixD —nd the se™ond one ˜ringing sm—ll q—rg—no f—un— —nd not on its originD we should —dd

some ™ommentsF m—mm—ls into the q—rg—no —re—F fy—nd l—rgeD w—zz— 8

„he l—te i—rly€lio™ene —ge of the ˜—se of the ‚ustioni @PHHVA interpreted the f—un— of q—rg—no —s the

m—rine deposits th—t lo™—llyse—l the „erre ‚osse k—rst result of ˜oth — pure vi™—ri—n™e me™h—nism @the reli™ of

fissures @†—lleri IWVRY e˜˜—zzi et —lF IWWTY €—vi— et —lF — wider —nd older €—l—eo˜ioprovin™eAD —nd — dispers—list

PHIHA is —n upper ™hronologi™—l ™onstr—int for the enE me™h—nism vi— filtering ˜ridge in the wessini—nF

demi™ f—un—F „he lower ˜ound—ryD inste—dD is more proE engelone @PHHUA ™onsidered €rol—gus un—˜le to

˜lem—ti™D —lso ˜e™—use the su™™ession of ph—ses of f—un—l ™ross m—rine ˜—rriersD —nd she sust—ined th—t this o™hoE

settlement likelyinvolved — su˜st—nti—l time interv—lF sn tonid dispersed on isl—nds vi— l—nd ˜ridges in the wesE

this ™ontext the nonEendemi™ t—x— from the older fisE sini—nF

sures —re most suit—˜le for inferring the —ge of the ˜eE †—n den roek ystende @PHHIA suggested th—t

ginning of the fossil re™ordF e™™ording to he qiuli et —lF ˜oth q—rg—no irin—™eid—e @the gi—nt heinog—lerix —nd

@IWVU˜A —nd preudenth—l 8 w—rtõÂnEƒu—Ârez @PHIHAD the the sm—ll gymnureA h—ve — €—r—sorex —n™estorD whose

o™™urren™e of epodemus would implyth—t the oldest immigr—tion should ˜e younger th—n wxW @e—rly †—lleE

––wikroti— f—un—99 ™ould not ˜e older th—n the „uroli—nD si—nAF pin—llyD v—n den roek ystende et —lF @PHHWAD ™riE

˜e™—use the first o™™urren™e of epodemus in iurope is ti™ising w—zz— 8 ‚ustioni @PHHVAD fo™used on the tempo

in wxIPF roweverD the —˜ove mentioned —ffinities with

—nd mode of ™oloniz—tion of the e˜ruzzoEepuli—n —re—F

l—te „uroli—nEe—rly‚us™ini—n ™ontinent—l spe™ies @preuE

„hey—ssumed — sweepst—ke me™h—nismD ™onsidering

denth—l IWVSY he qiuli et —lF IWVU˜A —lso suggested the th—t this model should ˜e the most suit—˜le to expl—in

possi˜ilityof — younger—ge @wxIQEwxIRAF por th—t the un˜—l—n™ed fe—tures of the q—rg—no f—un—D whi™h in

re—son there is — gener—l —greement on the v—te wioE their opinion is equiv—lent to the f—un—s th—t —re ™omE

™eneEi—rly€lio™ene —ge of the –– wikroti— f—un—99F ynly monlyfound on the o™e—ni™ isl—ndsF „he —rriv—ls were

v—n den roek ystende et —lF @PHHWA h—ve sust—ined —n supposed to ˜e ™on™entr—ted in — definite time interv—l

e—rly„ortoni—n —ge for ˜oth ƒ™ontrone —nd q—rg—noF when —ll the —n™estors of the q—rg—noEƒ™ontrone f—un—D

nfortun—telyD they did not ™onsider th—t the oldest in™luding rodents —nd other sm—ll m—mm—ls @not found

re™ord of epodemus in iurope is some P w— l—ter —nd

—t ƒ™ontroneAD re—™hed the e˜ruzzoEepuli—n —re—F „hey

—pp—rentlyignore the p—pers ™ont—ining su˜st—nti—l disE

d—ted this interv—l to the e—rly„ortoni—nD —dopting the

™ussion on the —ge of the q—rg—no f—un—F „he —ttri˜uE

ƒ™ontrone ™hronology˜yw—zz— 8 ‚ustioni @PHHVAD —nd

tion —t —n e—rly„ortoni—n —ge to the q—rg—no f—un— is

strongly™l—iming —g—inst the possi˜ilityth—t the e˜ruzE

therefore unten—˜leD in our opinionF

zoEepuli—n f—un— m—y˜e derived from dispers—ls —ndG

purther ™omments ™—n ˜e —dded on the mode of

or vi™—ri—n™e s™—ttered throughout time @polyph—si™ poE

origin of the f—un—sF he qiuli et —lF @IWVT˜D IWVU˜A —nd

pul—tionsAF

w—zz— 8 ‚ustioni @IWWTD PHHVA tried to identifythe

preudenth—l 8 w—rtõÂnEƒu—Ârez @PHIHA refuse the

most f—vour—˜le p—l—eogeogr—phi™ ™onditions for verteE

e—rly„ortoni—n d—ting of ƒ™ontrone ˜one ˜eds m—king

˜r—te dispers—ls to the insul—r dom—inD on the ˜—sis of

— verydet—iled ™riti™ism of €—t—™™— et —lF @PHHV—AF fy

geologi™—l d—t—F „his method h—s ˜een exh—ustivelydisE

—nd l—rgeD they™on™lude th—t the fossiliferous l—yers

™ussed ˜yezz—roli @IWWTA —nd —pplied to sever—l insul—r

should ˜e ™onsistentlyyoungerof IHFS w— —nd older

f—un—sF iven though the —forementioned —uthors use

th—n TFR w— @p‚y of „ur˜orot—li— multilo˜—AF sn this

typi™—l dispers—list ™on™eptsD their —ppro—™hes —re —t

time interv—lD preudenth—l 8 w—rtõÂnEƒu—Ârez @PHIHA

the ˜ound—rywith vi™—ri—n™e ˜iogeogr—phyD—s stressed

™onsider the o™™urren™e of suit—˜le —n™estors of the

˜yw—sini et —lF @PHHP˜AF yn the other h—ndD v—n den

e˜ruzziEepuli—n f—un—D —nd ™on™lude th—t the wxIP roek ystende et —lF @PHHWA —dopted the typi™—l dispersE

zone @l—te „ortoni—nEe—rlywessini—nA ™ould ˜e — suiE —list model of sweepst—kesD —d—pted to the isl—nd theory

t—˜le interv—l for the immigr—tion of the f—un— onto the ˜yhermitz—kis 8 ƒond——r @IWUVA —s dis™ussed ˜yde

insul—r dom—inF preudenth—l 8 w—rtõÂnEƒu—Ârez @PHIHAD †os et —lF @PHHUAF preudenth—l first —dopted the ƒimpE

—s yet reported ˜eforeD sh—re the interpret—tion of v—n son9s @IWRHA model of l—nd ˜ridges @preudenth—l IWUID

den roek ystende et —lF @PHHWA of — single migr—tion IWUTD IWVSA ˜utD re™entlyD he —lso em˜r—™es sweepst—kes

possi˜lyo™™urred ˜ysweepst—kesD even though they @preudenth—l 8 w—rtõÂnEƒu—Ârez PHIHAF roweverD —ll the

—dmit th—t the p—l—eontologi™—l d—t— prevent to st—te —uthorsD with the ex™eption of v—n den roek ystende et

if the migr—tion w—ve origin—tes from e—stern or western —lF @PHHWAD —greed th—tD when vi™—ri—n™e is ruled outD

iuropeF f—un—s disperse onlyunder veryf—vour—˜le p—l—eogeoE

„his overview evidentlyshows th—t the question gr—phi™ ™onditionsD iFeFD when the ™o—sts of sour™eEl—nds

of the —ge of the fossiliferous deposits of q—rg—no —nd —nd t—rgetEisl—nds —re rel—tively™lose e—™h other @see

of the d—ting of the f—un—l migr—tions —re still not ™omE —lso w—sini et —lF PHHP˜AF prom this point of viewD if

™oloniz—tion is vi— l—nd ˜ridgesD filtering ˜—rriers or pletelysettledF po™ussingD —t firstD onlyon the —ge of the

RQP w—sini pFD ‚in—ldi €F wFD €etruso hF 8 ƒurdi qF

gon™lusion sweepst—kesD is not p—rti™ul—rlyrelev—ntF ispe™i—llyin

the ™—se of f—un—s like those of q—rg—noD for whi™h the

„his overview shows th—t —fter fortyye—rsof knowledge of ––surrounding99 p—l—eogeogr—phi™—l setting

rese—r™hes the q—rg—no f—un— isD on one h—ndD — most

—nd ™oev—l f—un—s is verys™—nt —nd impre™iseF

interesting study™—se for evolution —nd ˜iogeogr—phyD

„he —ssumption of o™e—ni™ isl—nd ˜yv—n den

˜utD on the otherD it still presents — ™onsider—˜le roek ystende et —lF @PHHWA —lso requires some ™ritiE

—mount of un™ert—intyD espe™i—lly reg—rding —ge —nd

™ismsF ƒweepst—kes onto o™e—ni™ isl—nds —re expe™ted

proven—n™e of the t—x—D —s well —s the —ge of the fossiE to ˜e s™—ttered in timeD —nd not ™on™entr—ted in p—rtiE

liferous depositsF fy—nd l—rgeD this ™ompil—tion shows

™ul—rlyf—vour—˜le interv—lsF iven if we —ssume th—t

th—t over the l—st ye—rs interest on this —n™ient insul—r

—rriv—ls —re ™on™entr—tedD too m—nyt—x—D iFeFD some

system is rising —g—inF ƒever—l new pu˜li™—tions —nd ten gener— of m—mm—lsD would h—ve dispersed on q—rE

works in progress —re supplying new tesser—e to the g—no following — sto™h—sti™ me™h—nismD su™h —s — tyE

mos—i™ of the q—rg—no historyD even though the det—ils pi™—l sweepst—keF preudenth—l 8 w—rtõÂnEƒu—Ârez @PHIHA

—re f—r to ˜e exh—ustivelyre™onstru™tedF et the st—te of h—ve —n —pp—rentlysimil—r positionD ˜ut theylook for

the —rtD q—rg—noD like m—nyother p—l—eontologi™—l o˜E f—vour—˜le ™onditions in whi™h sweepst—kes o™™urredD

je™tsD re™—lls —n smpressionist p—intingD whi™h is ™le—r

—nd they—ssume — lesser num˜er of in™omersD ™onsidE

enough if seen from — dist—n™eD ˜ut ˜lur when the ering m—nyof the endemi™ t—x— —s derived from less

™—nv—s is o˜served in the det—ilsF endemi™ ™ounterp—rt found in the s—me „erre ‚osse

—ssem˜l—geD eFgF wikroti— origin—ted from epodemusD

e™knowledgmentsF ‡e w—nt to th—nks w—ssimo helfino @ niE

r—ttomys from epo™ri™etus —nd heinog—lerix from the versityof ploren™eD st—lyA—nd w—tthijs preudenth—l @ niversityof

qr—n—d—D ƒp—inY x—tur—lisD the xetherl—ndA for — ™riti™—l ™he™k of the verysm—ll q—lerixF sn our opinionD howeverD ™onsiderE

list of referen™es on q—rg—noD endre— ƒ—vorelli @ niversityof ploren™eD ing how widelydifferent is endemism in q—rg—noD

st—lyA for the upd—ted d—t— on ™ri™etidsD —nd €—ul w—zz— @ niversity of

r—nging from highlyendemi™ to nonEendemi™D ––™ontiE

ploren™eD st—lyA for — ™riti™—l re—ding of the m—nus™riptF „he work is

nent—l99 spe™iesD it seems veryh—rd to suppose — single

supported ˜ypondi d9eteneo @ex TH7A of the niversityof €—lermoD

migr—tion w—veF ye—r PHHV —nd ˜y €‚sx PHHV @godeX PHHV‚„gtr•HHRAD niversity of

perr—r—F

‚ipi‚ixgiƒ

f—llm—nn €F @IWUQA E possil †ogel —us dem xeogen der r—lE e˜˜—zzi vFD fenvenuti wFD fos™hi—n qFD homini™i ƒFD w—sini

˜inseln q—rg—no @st—lienAF ƒ™ript— qeologi™—D IUX IEUSF pFD wezz—˜ott— gFD €i™™ini vFD ‚ook vFD †—lleri qF 8

f—llm—nn €F @IWUTA E possil †ogel —us dem xeogen der r—lE „orre hF @IWWTA E ‚evision of the xeogene —nd €leiE

˜inseln q—rg—no @st—lienAF weiter „eilF ƒ™ript— qeoE sto™ene of the q—rg—no region @epuli—D st—lyAF „he

logi™—D QVX IESWF m—rine —nd ™ontinent—l su™™ession —nd the m—mm—l

futler €FwF @IWVHA E „he gi—nt erin—™eid inse™tivore heinoE f—un—l —ssem˜l—ges in —n —re— ˜etween epri™en— —nd

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q—rg—noD st—lyF ƒ™ript— qeologi™—D SUX IEUPF QVQERHPF

h——ms ‚F 8 de fruijn rF @IWWSA E e ™l—ssifi™—tion of the e˜˜—zzi vFD w—sini pF 8 „orre hF @IWWQA E ivolution—ry

qlirid—e @‚odenti—A on the ˜—sis of dent—l morpholE p—tterns in the first lower mol—r of the endemi™ murid

ogyF rystryxD T@IEPA@IWWRAX QESHF wi™roti—F u—ternF snterFD IWX TQEUHF

h——ms ‚F 8 preudenth—l wF @IWVSA E ƒtertomys l—ti™rest—tusD engelone gF @PHHSA E ivolution—rytrends in dent—l morE

— new glirid @dormi™eD ‚odenti—A from the insul—r f—uE phologyof the genus €rol—gus @y™hotonid—eD v—goE

n— of q—rg—no @€rovF poggi—D st—lyAF ƒ™ript— qeologi™—D morph—A in the wediterr—ne—n isl—ndsF wonF ƒo™F ristF

UUX PIEPUF x—tF f—le—rsD IPX IUEPTF

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@IWUWA E sl xeogene del wonte q—rg—noF follF ƒo™F gomorph—A of st—lyF qeo˜iosD RHX RHUERPIF

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preudenth—l wF 8 w—rtõÂnEƒu—Ârez iF @IWWWA E p—milywuriE he qiuli gFD w—sini pF 8 „orre hF @IWVU—A E „he l—test

d—eF snX ‚ossner qFiF 8 reissig uF @idA E „he wioE È †ill—fr—n™hi—n f—un—s of st—lyF „he €irro xord p—un—

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preudenth—l wF 8 w—rtõÂnEƒu—Ârez iF @PHHTA E qlirid—e @‚oE he qiuli gFD w—sini pF 8 „orre hF @IWWHA E ssl—nd endemism

denti—D w—mm—li—A from the l—te wio™ene fissure fillE in the e—stern wediterr—ne—n m—mm—li—n p—leof—un—sX

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preudenth—l wF 8 w—rtõÂnEƒu—Ârez iF @PHHWA E „he —ge of he qiuli gFD w—sini pFD „orre hF 8 foddi †F @IWVU˜A E inE

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f—un—sF snX p—nti pF 8 ƒp—llett— gF @idsA E sntern—tion—l q—rg—no ™—se historyF follF ƒo™F €—leontolF st—lFD

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logn— PVEPW ƒeptem˜er PHHWF e˜str—™t fookX RPERSF he qiuli gFD w—sini pFD „orre hF 8 †—lleri qF @IWVT˜A E

preudenth—l wF 8 w—rtõÂnEƒu—Ârez iF @PHIHA E „he —ge of

w—mm—l migr—tion events in emerged —re—s of the

immigr—tion of the verte˜r—te f—un—s found —t q—rg—E

epuli—n pl—tform during the xeogeneF qiornF qeolF

no @epuli—D st—lyA —nd ƒ™ontrone @l9equil—D st—lyAF

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gF‚F €—levolD WX WSEIHHF he qiuli gFD w—sini pF 8 †—lleri qF @IWVU™A E €—leogeoE

qohli™h FfF 8 €—vi— wF @PHHVA E e new spe™ies of €—l—eorE È

gr—phi™ evolution of the edri—ti™ —re— sin™e yligo™ene

tyx @evesX q—lliformesX €h—si—nid—eA from the xeoE

to €leisto™eneF ‚ivF stF €—leontolF ƒtr—tFD WQ@IAX IHWE

gene of q—rg—noD st—lyF yry™tosD UX WSEIHVF

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veinders tF @IWVRA E roplitomeri™id—e f—mF novF @‚umin—nE he qiuli gF 8 „orre hF @IWVRA E ƒpe™ies interrel—tionships

ti—D w—mm—li—A from xeogene fissure fillings in q—rE

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w—rtõÂnEƒu—Ârez iF 8 preudenth—l wF @PHHUA E qlirid—e @‚oE

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wusF qeolF f—r™elon—D IR@PHHTAX QUESWF

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w—sini pF 8 e˜˜—zzi vF @PHHTA E e prelimin—ryqu—ntit—tive

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