BIOCHRONOLOGY OF SELECTED MAMMALS, MOLLUSCS AND OSTRACODS FROM THE MIDDLE PLIOCENE TO THE LATE PLEISTOCENE IN ITALY. THE STATE OF THE ART

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1 Rivist tlin di Plentli e Strtirfi vlume luj numer 3 prne Dtcembre 7997 BOCHRONOLOGY O SELECTED MAMMALS, MOLLUSCS AND OSTRACODS ROM THE MDDLE PLOCENE TO THE LATE PLESTOCENE N TALY. THE STATE O THE ART E. GLTOZZ, L. ABBAZZ2,P. ARGENT3, A. AZZAROL2,L. CALO4, L. CAPASSO BARBATO4, G. D STEANO4, D. ESU4, G. CCARELL2, O. GROTT4, T. KOTSAKSS,. MASN6, P. MAZZA2, c. MEZZABOTTA2, M. R. PALOMBO4, C. PETRONO4, L. ROOK2, B. SALA/, R. SARDELLA4, E. ZANALDAO & D. TORRE' Key-urds: Cntinentl bichrnly, Pli-Pleistcene, lre nd micr mmmls, mlluscs, strcds. Rissunt, Gli Autri hnn elbrt quttr tvle di distri buine crnlic delle fune mmmiferi (mcr e micrq, mlluschi e d strcdi di cqu dlce e slmstr, prendend in cnsiderine i più sinifictivi icimenti del Pli-Pleistcene dell penisl itlin. n quest lvr viene istituit un nuv Età Mmmiferi (Aurelin) crrelbile cn l pne superire del Pleistcene Medi e cn il Pieistcene Superire: di quest età sn stte definite due Unità unistiche (Trre in Pietr e Vitini) per l'aurelin inferire e medi, mentre nessun iciment è stt cnsidert rppresenttiv per 'Aurelin superire. k unità bicrnliche sn stte clibrte cn l mnetstrtirfi, cn le scle istpiche dell' ssi en e ttrvers étim rdimetriche. Abstrct. The Authrs hve elbrted fur rne chns f mmmlin (lre nd smll), mlluscs nd fresh-wter nd brckish strcdes funs, fr the selected Pli-Pleistcene fssiliferus lclities f the tly. A new Mmnl Ae (Aurelin) crreltble t lte Middle nd Lte Pleistcene hs been defined. nside this e tw unl Units (Trre in Pietr nd Vitini) hve been defined s chrcteristic fr erly nd middle Aurelin, while n isements hve been chsen fr the lte Aurelin. Bichrnlicl units re clibrted n mnetstrtirphic nd istpic scles nd by rdimetric dtins. lntrductin. Present dy knwlede but the terrestril mmml funs, nn mrine mlluscs nd freshwter brckish strcds f the tlin Pli-Pleistcene enbles us t ttempt t cnstruct n interted bichrnly. This hs been chieved by mens f cnstructin fur distributin chrts f selected tx frm the mst sinificnt tlin lclities. t is wrthwhile underlinin tht the mmml es used fr the bichrnlicl scle f the lre mmmls re is used (minly fr histricl resns) fr the bichrnly f the mlluscs nd strcds. The species distributin chrts refer t the unl Units (.U.) successin (Arli, 1977; Arli et l., 7982, 1988). This successin cnstitutes the bichrnlicl scle fr the cntinentl successin in the tlin peninsul, nd is clibrted with the echrnlicl scle by mens f rdimetric nd plemnetic dtin. Successive funl units re nt seprted by bundry becuse f the impssibility f definin them with precisin. The unreiibility f the bundries derives frm tw cuses f different rder: cntinentl successins re lrely discnti- ") nuus nd mmml remins fe usuliy cncentrted in hrins with punctul distributin in strtirphic successins. Therefre nly exceptinlly cn the trnsitin t the next sscitin be recnised in strtirphic successin. b) the definitin f.u., which is bsed n ll the species frm lcl funs selected s typicl sscitins. This ltter pint merits brief discussin. unl Units in bichrnly crrespnd t wht in bistrtirphy re cennes nd, in the sme wy, their limits re nt definble with precisin. The distinctin between successive cennes is nt bsed n sinle fx ^ppernce,/disppernce bievents, but n different sscitins f chrcteristic tx. n fct bievents chrcterisin successive.lj. ccur durin the trnsitin between the tw.u., nd their clibrtin is cnse- 1l C..r' St".li^ ^.r il Quternri e 'Evluine mbientle CNR c/ Dipniment di Sciene dell Terr, Università deli Studi "L Spien" di Rm p.le A. Mr 5, Rm, tly. 2) Dipniment di Sciene dell Terr, Università deli Studi di irene. 3) Dipniment di Sciene dell Terr, Università deli Studi di Perui. 4) Dipniment di Sciene dell Terr, Università deli Studi "L Spien" di Rm. 5) Dipniment di Sciene Geliche, Università deli Studi di Rm Tre. 6) Dipniment di Geli e Gedesi, Università deli Studi di Plerm. 7) Dipniment di Geli e Pientli, Università deli Studi di errr. 8) Dipniment di Sciene dell Terr, Università deli Studi di Miin.

2 374 E. Glii et l. quently ffected by sme inccurcy (fr wider discussin n this subject we refer the reder t De Giuli et. L, 1986; Lindsy, 1990). Clerly this fcrr enertes certin "bckrund nise" in the cntents f the distributin chrts. n dditin n even mre imprtnt bis is cused by the fct tht different unl Jnits re nr hmeneus frm the pint f view f the diversity f ccurrin species. An exmple f this is iven by the funl renewl tht ccurred t the beinnin f the Middle Pleistcene, between the Clle Curti nd the Slivi unl Units. n fct the few species ccurrin t Clle Curti - which in ny cse re indictive f renewl - des nt mke it pssible t estblish whether this funi chne ws lredy under wy previus t this funl unit. Despite these limittins, the picture which results frm this wrk ives resnbly cceptble infrmtin but the timin f the funl renewls durin the Pli-Pleistcene. The distributin chrts. The unrelibility f bichrnlies bsed n mmml ssembles mentined bve frced us t use sme rphic cnventins nd symbls in the distributin chrts which need sme explntin. On the left in i. 1 nd 2 re the cmpsire curve f xyen istpes mdified frm Shckletn (1995), the chrnliclly clibrted mnetstrtirphic scle (Bksi, 1993) nd the echrnlicl scle. The bundry between the middle nd the lte Plicene is plced t istpic ste 103, just befre the beinnin f the mrked clin event which tk plce t but 2.5 M (R.i et l., 1994). The bundry between the edy nd the Middle Pleistcene is plced clse t istpic ste 25, fllwin the prpsl mde by Cit 6c Csrrdri (1994). The bundry between the Middle nd the Lte Pleistcene is plced t istpic ste 5e (Auirre & Psini, 1985). unl units clibrted by rdimetric nd/r plemnetic nlyses re reprted in the clumn "Clibrted lcl funs". The rphic cnventin indictin the trnsitin between Mmmi Aes is verticl br which cvers the spce between the lst nd the first funl unit f tw successive mmml es. The sme rphic cnventin is used rc indicte the bse f selected events, which is plced in n intermedite psitin between tw successive funi units. The distributin f the tx is represenred by cntinuus verticl line when the presence f tx is dcumented. \When the presence f txn is inferred r is dcumented by scrce r frmentry mteril, it is represented by dmed line. The precise successin f the ppernce,/disppernce f ech txn chrcterisin successive funl units is nt knwn. Only in few cses is it pssible t recnstruct the reltive chrnlicl psitin f ech lcl fun within funl unit. r this resn lcl funs in funl unit re cnventinlly reprted in lphbeticl rder nd first/lst ccurrences f tx re cnventinlly plced t the bse f ech funl unit, represented by dtted line. The synchrny f events bservble n the distri butin chrt is ttributble t this rphic cnvenrins. r the resns discussed bve, the temprl p between ppernces is smetimes emphsised by the scrcity f lcl funs, especilly in the lder prt. urthermre, the scle f the funl renewl (mesured by the number f new ppernces) is bised by the îefer r lesser richness f the lcl funs which mke up ech funl unit. Lre mmmls. Villfrnchin Mmml Ae. The ldest fun we re delin with is the Trivers.J., which dtes bck t the Middle Plicene. Erly Plicene mmmls in tly re exrremely prly represented. A few fssils were cllected in the Vl di Pun (Sien), ssined Stepbnrhinus merhinus, Sus minr, Alephis liryx nd elsintherium ertisi (Arli et l., 1988). A further find f this e is frmentry metpdil f Alephis liryx {rm the fluvilcustrine bsin f Mntecrl ner Lucc (Dlln Nrdi, 1988). Erly Plicene mmmis re nt sufficiently well represented in tly t be included in the distributin chrt. The fun f the Trivers.U. (the first funl unit f the erly Villfrnchin) shw mrked renewl with respect t previus sscitins which re well chrcterised in lclities f Spin nd suthern rnce (see Arli et l., 1988, with references) nd which re cmpsed f tx which re typicl f humid nd dense frested envirnments. The Trivers.U. is chrcterised by the ssemble f fssils frm vrius sites in the Villfrnc d'asti re (S. Pl, Slbrit, Dusin, rnce R.D.B., Cs Crtin, Cs Cssintt, Arbschi, Arndelli) nd tkes its nme frm smll river in the re. The vertebrte berin fluvi-lcustrine successin lies n Erly Plicene (MPL 4) mrine clys. The e f this.u. shuld be rund 3.2};,4, s suested by plemnetic nlyses t rnce R.D.B. (Lindsy et l., 1980), lthuh new nlyses by Lindsy et l. (1995, nd rl cmmunictin) suest tht the rnce R.D.B. sectin shuld be ttributed t the upper prr f the Gilbert mnetchrnnîe. The first dte hs hwever been cnfirmed by the plemnetic survey recently crried ut t Snt Brbr ner Csrelnuv dei Sbbini in the upper Vldrn (the first fluvilcusrrine cycle

3 E. Glii et l, à 18 curue Í (Shckletn, l9s5) : is._-- =--=: à = -):- r = -_j.-- ìslpic Sle lslpic Ste M?0736M\ Gril del Brin, Gf1l del Cvll, P iccl lft T ien1e, S Buclne, C rvelerl, l,,l spin Sedi del Divl lre n Piet (u l.), Vilni Csle di Guid, Mlqrll, R n, lr in Piet (l.) Cv Ner lú infi, nln Rnlcc. Ces, lserni, Pnle G eri (vrius sltes), Vldemin Sl v. Mnle T nd!!r À"1 3 sb -È '-* t "i _ ==-> =t :> <-: =lt- -= t -, ^- L,,*, C le Cudi. Mfie Pel É 'Éî -=: :r= -= 6'.-.=-= : -< =---:-, _j Upper Vldfn (Cs Mlssin. Oliv ill ttt >- -= -:",m--l,* == --.1 '* -.:-- -? [*- ì --;- ]-'- ': -= ì 1 Arcf e Snl Brbf, rvers (vfius slles) Bichrnly f selected lre mmmls f tlin Peninsul frm Middle Plicene t Lte Pleistcene p.2 C.{ f î;:4 4 4 È e i 4 4 * 1,.q.9! e r - Ò * / t EÉ:fÍEÌ;ÈÈ:eiri;l'i2 :È*' e+; )ì;!eéi3l q i ú ú 1È E;::: É r: t ;: i: È; i Èi É:; ; ; t iiéi! i be!+ìs;: *dsièiè ít iririi :i E s ii;è!!j' ÉÉ51;; È:;?l;! Hs!it 3.i rérél c! 3cÈ fsè;r d " P.E:ÈeA-:È P d!!t96=-d i J; * P E È se ;óh;èeèè 5 i-;;5ei E E8ÈEtr È ; EqùÈ e5 < à, Si íi:j: ;e^".0 óe:'3 r e:q i; E qèà 9::ó ;< ÈÈ à c4 ei.r.-j*f, Éi,je 3 i ;) È 5.: i! ^ e,- ^ 6> ' E3+ d 3 È".i S 3 :-=! 6t ; >ò i "!': E t " ;e j -d! : + : r E {É Ì È E ; { u '5 ;s:3 Eèe -cl 6: ÈÈ : ; >1 ó> f 'The lxnmicl psìlin 01 lh se irms is discussed by sm Aulhrs

4 >è àó,'t i:r -;l t" u i:l t:.t ì '1 '!_ : È;tl E.ó' íîl!1 E i Él ll ;É1 :,É 1 i l* t Èàt :î i iltl tl ttl ill ill ll ll sp:5ee9é$ {q,;;i t\pii! \4iS4;i:-!4q ièiè;c!:;islè5i! ui:=r! {: E i E CTEe;J;EE EÈ+èqt:è fti e; ; É E i:! ie É:!É:i 9+-à;Hàí È! ii d::[!! È* s.:è:flir 3;ip: ii iì: j;ph HEÈ;_,ì9tjE :.: i"=iì! É! ;3*3::îE i" E l5;i ri fqpeió*= ; É É É;EE ;:3iH;É';:iÈ' e!;; 1;;3! HÉ[:*',ii:É' ;Eà È É ÈE:8";-! Q=È à*: 5 *b à;d 14- e É6 - :-ì;ó-erf < ee ÉJi -i r::ó i< -- É;"- i Ed;" r. ut? 5:i;YÉ ; EE :59 -S 5 ò 3q"- : b> -::- 6* = '5! E -:r 9 i: ' 'ié:!; 6> 3 E =ò E É 1 te E + ;ù,: *; ú Èit! 3 i?i i i:, i liià:ii i i! ii iéiil i 3ÉiE {iir ÉÉE i i si Érf!t'E;EiE i =$"ti* THÈi-" # nj+,:'3 O'? Ésié**eÉti àtféc.béèhè H!Ì" 'ecèii "_ti EsrÀi : i 986 e ÉÀ ' The ixnm cl psli n t lhese ifms s disclssed by sme Aulhrs

5 Bichrnly f Pli-Pleistcene f mmmk, rnlluscs nd strcds in ltly -)/ f the upper Vldrn), where funl ssemble f the Trivers ljnit ws fund (Trre et l,, 1996; Albinelli et., 1997). n dditin t the bve mentined lclities, lcl funs referble t this.u. re present in different res f tly: S. Giust (Trre, 1987), Pnte Els (Arlí, 1992), Cse Nuve (Dminici et l. 1995; Benvenuti et l., 1995) in the lwer Vldrn; Splet in Umbri (Tuccimei, 1898; Srdell et., 1995) nd Arciile ner Grsset in suthern Tuscnv (Hùreler 6c Enesser, 1976; Msini &.Trre, 1987). n this unit there is the first recrded ppernce f the enus Pseuddm, Leptbs stenmetpn nd Stephnrhinus jenvireti, nd severl lre crnivres such s Acinnyx, Hmtherium nd Ursus minimus. There re ls frms f Viverride (Vierr nd Mevien) nd n extinct frm f pnd (Priluru). Tpirus druernensis nd Sus minr re still well represented whíle Mespithecus mnspessulns ccurs fr the lst time. The beinnin f the erly Villfrnchin (Trivers.U.) thus represents mrked funl turnver. The ccurrence f new tx enriches the fun which. s whle, hs mre dvnced chrcter cmpred with the Ruscinin nes. A mrked funl renewl mrks the trnsitin between this.u. nd the fllwin Mntpli.U.; tx typicl f wded hbitts dispper lzylphdn brsni, Thpirus rernensis, Sus minr (: Sus rernensis); Ursus minimusf nd in tly ne sees the spred f primitive elephnt (Mmmuthus rmwi), nd the hrse (Equus lienwensi). Bth f^x^ re indictive f mre pen lndscpes. At tht time Stephnrhinus etruscus ls prbbly ppers (R.ustini et., 1,995). The lcl Mntpli fun, in the Lwer Vldrn, ws cllected t the tp f mrine successin wìth Glbrtli crssfrmis nd is clibrted with the Guss/Mtuym trnsitin in the mnetstrtirphic scle (Benvenuti et l., t995; Lindsy et l., 1980; Trre, t987). The smll lcl fun t Clleprd, lcted in the Anni bsin in Ltium, is cmpsed f Menteren cultridens, Nyctereutes menxdstides, Stephnrhinus cf. S. jenireti, Ss sp. (f smll sie), Pseuddm lyr nd "Hernitrus cf.. H. stehlinl" (Cssli & Sere Nldini, 1994). \fith sme reservtins due t the scrcity f mteril, this smll fun culd be referred t the end f the erly Villfrnchin. Middle Villfrnchin funs re prly represented in ltly, nd they re represented by smll ssembles which hve nt yet been fully studied. One f these cmes frm the lclity f Cst S. Gicm, in in the Anni bsin. The Cst S. Gicm lcl fun is chrcterised by the first ccurrence f the enus Cnis s.s., Sws cí. S. strii, Leptbs furtitus, Hystrix cf.. H. refss nd Gellspir trticmis, nd the lst ccurrence f Anncus drernensls (Cssli & Sere Nldini, 1994; Msiní, 1.989; Rk & Trre, t996). The funl sscitin ls includes Stephnrhinus cí. S. etruscus while Nlrctereutes rrrcrnstides, Stephnrbinus cf. S. jenireti nd the smli sied suid re n lner present. The beinnin f the lte Villfrnchin (Olivl.U.) is mrked by the ppernce f Pclrycrcwt breirstris, Pnther mbseensis (: P. tscn), Prcmptcers britense nd new species f Eucldcers nd Pseuddm (E. dicrnis, P nesti). urrhermre mdern ds (Cnis etruscu) becme widespred. The mjrity f the Plicene species which hd survived the first crisis linked with the first rctic lcitin (istpic ste 100 nd fterwrds) re n lner present r re extremely rre. The mst imprtnt lcl fun is tht f Olivl (vl di M".") which is prbbly lcted in the Olduvi chrnne. n fct, reserch which is still in prress t Pi Rss (upper Vldrn; Trre, unpublished) suests tht the djcent site t Mrssin, which in the pst hd been ttributed t this.u. nd plemneticlly clibrted with the B event in the upper prt f the Olduvi chrnne (Trre et l., 1993, t996), hs funl ssemble with chrcteristics which re trnsitinl between the Olivl.U. nd the fllwin Tss.U. The Tss.U. is chrcterised by the rrívl f Htpppumus ntiquus, Prewibs, bisnlike kptbs (L. vllisml, Equus steblini, Cnis rnensis nd Cnis (Xencyn) flcneri, tether with the disppernce f Gllrl menebinii nd Gellspir trticrnis (De Giuli & Msini, 1987; Arli et l., 1988; Ambrsetti et L, 995). These events determine mrked chne in the funl sscitin, nd it cquires typiclly Quternry chrcter. After the turnver respnsible fr the frmtin f the Tss,J., mst f the typiclly Villfrnchin tx dispper r becme extremely rre, nd species which re phyleneticlly clser t the Middle Pleistcene species pper. This new situtin chrcterises the rnet.u. (Arí, 1977). kptbs nd Eucldcers re still present nd, fr the first time, deer f the Mecerides clde (M. bscwru) nd the vle Micrtus (Allpbirny) pper (Msini & Sntini, 1.991; Arli &M, 1993; Abbi, 1995). The richest lclity fr this.u. is Pietrfitt in Umbri (Ambrsetti er l., 1987; Ambrsetti et., 1992, b) where there re chrcteristic smll sied rhincerses (M et l., 1993). Pirr Nrd-Cv dell'erb, krst fissure fills in the Grn, (De Giuli et l., 1987; Abbi et l., 1,996 b), nd Cpen ner Rme (Petrni, 1,979) re tw lclities where the first ccurrence hs been recrded f primitive bisn, Bisn (Ebisn), nd f Mecerides species, prbbly ttributble t rhe M. slilbcus phyletic linee (Abbt & Msini, 1997). The presence f vibvine remins t Pirr Nrd is ls very sinificnt. These nvelties chrcterise the Pirr Nrd.U.

6 372 E. Glii et l. Glerin Ae. Tx which re cnsidered t be clssiclly Glerin (such s Equus ltidens, Equus r. bressnus-suessenbrnensis nd the enus Meceri.de) lredy ccur in the.u. f the Lte Villfrnchin nd the funl renewl tht prduced typiclly Glerin ssembles ws rdul phenmenn. The Clle Curti lcl fun is cnventinlly used t mrk the beinnin f this Mmml Ae. This funl ssemble (iccrelli & Silvestrini, 1991), which unfrtuntely is nt very rich, ws fund in sediments with nrml plrity nd which re crrelted with the bse f the Jrmill subchrnne (Trre et l., 1996; Albinelli et l., 1997). The fun is chrcterised by the presence f the first frms f Mecerides perticrnis. The smll number f species recrded in the ssemble des nt mke it pssible t verify hw mny typicl Glerin tx'were present t the timet fssils f medium-sied bvine culd be ttributed either t smll Bisn r, lterntively, t lcptbs. The presence f Plimls lenki tether with Micrtus (Allpbimys), which ws prbbly ncestrl t Micrtus ex r. ecnmus, cnltrms the innvtive chrcter f this funl ssemble nd strenthens its ttributin t the Glerin. Subsequently there were successive dispersl events f Asitic nd centrl Eurpen tx int the tlin peninsul, prbbly in reltin t phses f climtic clin. At the sme time new tx pper s result f lcl evlutin. \íithin the Glerin cmplex it is therefre pssible t recnise three successive sscitins chrcterisin the erly, middle nd lte Glerin respectively. The erly Glerin ssemble is chrcterised by the presence f Mecerides uerticrnis nd by the persistence f frms with Villfrnchin ffinities such s P clry cr cut bre ir stris, H mteriurn ex r, crentidensltidens, Mmmuthus (Archidiskdn) meridinlis, smll sied Stephnrhinus nd n dvnced frm f Pseuddm. The Clle Curti.U. is ttributed t the erly Glerin; it ws estblished n the bsis f the funs f the Clle Curti site nd thse frm Mnte Peli (vn der Meulen, 1.973; Pipern & Sere, 1984). At the ltter site the ire mmml ssemble, which hs yet t be studied nlyticllt suests the persistence f lre munt f Villfrnchin tx, while the smll mmmls indicte mre evlved ssemble thn the ne frm Pirr Nrd. t ppers therefre tht the erly Glerin ssemble in ltly, even thuh it is pr in species nd scrcely represented in terms f number f sites, cnstitutes cmplex which is indictive f the trnsitin between typicl lte Villfrnchin nd typicl Glerin funs. The middle Glerin ssemble, which is chrcterised by the presence f Centus elphus dcrntus, shws mre mrked funl renewl which cn lredy be seen in the Slivi.U. nd which is much mre nticeble in the serni.u.. Lin, plelxdnt elephnts nd the subenus Mmmuthus, typicl frm f Stepbnrhinws bundsheimensls, new meicerines nd new bvids such s Bs nd bisn belnin t subenus.blsn (Bisn) nd, mn the equids, Equus cbllus, ll pper. Sme Villfrnchin lre crnivres nd equids s well s n dvnced frm {. Pseuddmd re srill present. The ldest sites with middle Glerin funs be- nin t the Slivi.U. re Slivi (Ambrsetti et l., 1979; Bn et 1., 1992) nd Mnre Tend (Ps, 1947). Due t the persistence f sme Villfrnchin elements, the Selv Vecchi lcl fun (Bn et ^1., 99) seems r be slihtly mre rchic thn tht t Slivi. The ssembles frm serni (Sl, 1983, 7996), Cesi (lr4sini et l., 1991; iccrelli et l., in press) Vldemin (Sl, 1992) nd vrius sites in the surrundins f Pnte Gleri (Cpss Brbt & Petrni, 1984; Cli & Plmb, 1994) re ttributed t the fllwin serni.u. The fun frm Vislin (Cttni et l., 1991) nd Vens-Ntrchiric (Belli et 1., 1991) re prbbly trnsitinl with thse f the lte Glerin. The lte Glerin ssemble, t which the ntn Rnucci.U. is referred, is chrcterised by the ttl disppernce f typicl Villfrnchin rx with the exceptin f Hmtherium, whrch is represented by n evlved frm (Srdell, 1,994). The red deer subspecies Centus elphus estepl)ncers is present (Di Stefn & Petrni, 1993) nd Dm clctnin, E. ltidens nd E. suessenbrnensls cntinue t ccur. The mst representtive sites re ntn Rnucci (Sere, 1984) nd Cv Ner Mlinri (Di Stefn Ec Petrni, 1993). Aurelin Mmml Ae. A mrked renewl ccurs t the trnsitin t the fllwin funl ssemble, with the extinctin f sme Glerin frms such s the mecerine cervids f the Mecerides erticrnls rup, Melcers sini nd the mre rchic elphine deer, while tx which cnstitute the cre f the mdern mmml fun pper fr the first time. Very rich depsits re minly lcted in the Tyrrhenin cstl re Í Ltium, in the S.S. 1 Aureli. r this resn we prpse t use the term Aurelin e fr this funl ssemble. Aurelin funs cme frm depsits which dte t the lte Middle Pleistcene nd the Lte Pleistcene. Bievents such s the first ccurrence f the mdern lre wlf Cnis lupus nd f Ursus speleus hve been chsen t define this funl ssemble. n the Aurelin funl cmplex there re three different nd subsequent sscitins (erly, middle nd lte Aurelin) which re relted t the prressive ccurrences f diffe-

7 Bicbrnly f Pli-Pleistcene f mmmls, mlluscs nd strcds in ltfu J/ 3 rent tx f Asin riin r f ther frms which evlved in the Mediterrnen re. The erly Aurelin ssemble, which is represented by the Trre in Pietr.U., is chrcterised by the first ccurrence f the mdern wlf nd f the cve ber s well s by the first ccurrence f Melcers idnteus nd the diffusin f the mst rchic frms f mdern red deer Cerus elpbus riiensis. Crreltins between the type lclity f Trre in Pietr (Trre di Pliccett) nd the Middle Pleistcene Tyrrhenin shrelines refer this funl unit t istpic ste 9 (Cnt et l., 1980). The mst representtive depsits f this unl Unit, besides Trre in Pietr (wer level$ (Cii & Plmb, 1928; Csslí, 1978), re Cstel di Guid (Sl 6c Brbi, 1996), Mlrtt nd Rin (Cli Ec Plmb, 1994) nd l Plledrr di Cecnibbi (Anidei et 1., 1989; Arnldus-luyendveld & Anidei, 1993), lcted in Ltium. The next funl ssembles, referble t the middle Aurelin (istpic ste Z), re chrcterised by the ccurrence nd diffusin f the mst primitive subspecies f the mdern fllw deer Dm dm tiberin (Di Stefn & Petrni, 1997) nd Equus lrydruntinus, durin perid f climtic melirtin which is ls recnisbie in the mrine envirnment. The Vitini.U. hs been ttributed t the middle Aurelin; it is represented by the fun frm Vitini, Bucine, Cerveteri, Sedi del Divl nd Trre in Pietr (upper levels) (Cli & Plmb, 1994). The lte Aurelin ssemble, which beins with the Eemin nd termintes with the end f the lst lcitin, is difficult t define becuse it is minly chrcterised by the disppernce f lre nd medium sied mmmls. n the time spn crrespndin t this funl ssemble there ws series f imprtnt climtic events nd the different micrclimtic nd envirnmentl cnditins hve strnly influenced the funl cmpsitin ln bth the Tyrrhenin nd the Adritic csts f the tlin peninsul (Sl 1983b; Msseti et l., 1995; Trre et l., in press). Durin this perid the funl ssembles becme prer nd, in crrespndence with the beinnin f the lst nterlcil, re mre nd mre cnditined by nthrpic influence. Even thuh the depsits relted t this ssemble re bth numerus nd well studied, fr the resns mentined bve nne f them cn be cnsidered t be fully representtive f the lte Aurelin funl ppultin, nd s result it is nt pssible t chse lcl reference fun t define funl units within this ssemble. The ldest lcl funs (istpic ste 5) referble t the lte Aurelin ssemble re chrcterised by the first ccurrence f the mdern frm Dm dm dm. The mst imprtnt depsits re minly lcted in centrl nd suthern tly, such s S. Sider (De Giuli, 1983) nd Melpinn (Bin et 1., 1994), Gttt" Mscerini (lwer levels), Guttri nd ssellne (Circe) (Cli & Plmb, 1.994b). Subsequently, nt nly in Eurpe but ls within the tlin peninsul, there ws n enlrement f the distributin re f Cpr ibex nd the spred f lre cld envirnment stenthermic mmmls such s Celdnt ntiqwittis nd Mmmuthus primieniws. Severl rx f lre herbivres nd crnivres prressively disppered in crrespndence with the fllwin phses f climtic deterirtin. The mst recent lte Aurelin ssembles cme frm lre number f lclities ll ver ltly nd hve minly been fund in cve depsits. Amn these the sites with the mst cmplete plentlicl, pleeclicl nd pleclimtic dt re Grtt del Brin nd Ripr Tliente, (Bn et., 1991) in Nrthern tly, Grtt Breuil, Grtt Brbr nd the upper levels f Grtt dei Mscerini, Grtt ssellne, Grtt Guttri nd Grtt S. Astin in Centrl tly (Cli & Plmb, 1994b), nd Grtt del Cvll (Brtlmei, 1980), Grtt Plicci (Sl, 1983 b; Abbt et l., in press b) nd Grtt dell Serrtur (Bertlini et l., in press) in suthern tly. Smll mmmls. n cntrst with the lre mmmls, studies f ltlin fssil smll mmmls strted nly much mre recently. n fct, with the exceptin f the reserch crried ut by Bsc nd rsyth Mjr lmst century, it ws nly fter the end f the Secnd rwrld Wr tht reserch ws renewed by Ps nd Brtlmei nd nly since the seventies tht reter number f plentlists ben t wrk n this tpic. The Middle nd Lte Plicene nd the Erly Pleistcene smll mmml recrd is reltively scrce nd the number f sites is smll, while the Middle nd Lte Pleistcene depsits re s numerus s thse fr the lre mmmls, lthuh the number f sites with bth kinds f funl ssembles is reltively lw. The bichrnlicl ttributin f the ssembles fllws the scheme prpsed by ejfr & Heinrich (1989). wever, since the lre mmml dt fr the Plicene nd Erly Pleistcene in the tlin peninsul re cnsiderbly mre detiled thn tht f the smil mmmls, where pssible the fun hve been referred t the bichrnlicl scle f the lre mmmls. Villfrnchin Mmml Ae. A rich fun with insectivres, rdents nd lmrphs cmes frm Cscin Arndelli (Villfrnc d'asti, Piedmnt) nd is typicl f the Trivers.U. The ssemble is chrcterised by shrews such s Epis-

8 1 J74 E. Glii. et l. ;l 180 curue (Shckleln, 1 995) :: u '' '.' 5 q! ò È Clibrted tcl iuns unl Uniìs Selected lclities Grlt del Brin. R pr leen, Cv A Vei. Gr t Gh cc. Csielclvìl, Grit Sertur, S. S def S Befnrd n [,lqire E e É E tl tl ::-' -.::. - : t_'-r- :.- n =:-. --:- t _ -..-.= --t :'= ez --..i...::,- -=.= ll 1, ib i i, irc i,, 1.5 E - ì t- -i- ='l=.i O.+ - : l*.è - s',1'rp c Sl.9e M? Mqi (KArddn!-i lr '*"it!1j" t rrì: "l Sliv. U Clle Cudr Olv U Lr S. Aslin (C i Berict Bscchlesnuv, Sn Givnni di D!ìn. Spess ll Vens'Nlrchiric Mnle lend Slìvi fl nn. S lv Vecchi C le Cuill. Mnle Sve Cv Sud Cse lnjern (Uppef Csle frncó d Sòpr 1l :il èl Él iî il 99 i-=: ;= *î :l sl 6_9 ;; ;î 3 E ilt tl tl ill ltl ill ill rl tl,lr.<- = -t É.... :.] Csl S G Cst S Gicm, li t tl ltl tm i& = >16 -:.:-,'' -l- -jt'- :t_ -.'., A J2 33 "i l i,l : " [-* C leprd Mnlp i Arcii-. Sntr Brbr. Tr;vers (vri!s silesl :> tl ll tl tl ll ll tl lt ltj liîi? É E;àlÉ: i;:3é qi;: e; ir i ;r ÉsÉ: E íé ì si * s: = = iéi ÈrÈ léèiiéiéèiéiiiuiéi! à;! :,Èi: iié i i; i s t' i ;j* ;É,È, ";i j r.2 - Bichrnly f selected smll mmmls f tlin Peninsul frm Middle Plicene t Lte Pleistcene.

9 Bichrnly f Pli-Pleistcene f mrnmls, mlluscs nd. strcds in tly 375 riculus ibberdn (:Asriculus fter lutterer, 1994), Blrinides mrie, Petenyi bunric nd Beremendi fissidens, by the vles Mimmys hjnckenesis nd Mirnrlys stehlini nd by ther rdents nd lmrphs (Mwscrdinus cf. M. plicenicils, Glirulws pusillus, Prlus sei etc.) (Beri et., 1967; Michux, 1970; Msini 6c Trre, 1987). The fun frm Arcille (Grsset, Tuscny) with Mimm.ys hjnckensis nd frm Sn Giust (Pis, lwer Vldrn, Tuscny), rype lclity f Mimm)s stehlini, re ls ttributed t the Trivers.U. These three funs re referble t the erly Villnyin (Msini & Trre, 1987). Mny Eurpen depsits berin the remins f Mimmls hjnckensis hve been clibrted with the lwer prt f the Guss nrml mnetchrnne. The fllwin funl ssemble ws fund in the krst depsit t Rivli Vernese (Vern, Venet). n this fun there re severl vles which ccur fr the first time: Mitnmjs plicenicus, Mimm.ys pitymyides, Mirnrn)s trnensis, Villny cf. V. exilis, cf. Eilbius sp., primitive frm f the enus Dinrm.ys, D. llernii nd shrew f the subenus Srex (Drepnsrex). The specimens f the envs Mimmlts îund in the depsit llw this fun t be ttributed t the erly Villnyin nd the middle Villfrnchin (Sl et l., 1994; Sl, b; nfni 6c Msini, unpublished dt). Sme remins f Apdemus cf. A. dminns fund in Cv Tppetti (Tdi, Umbri) (Abbi et L, 1996) re ruhly cevl with thse fund t Rivli Vernese. The islted remins f Mimrnys plicenicus fund t Cstelfrnc di Spr (Upper Vldrn, Tuscny) nd Bcchinn (Sbin, Ltium) (lr4sini & Trre, 1987; Ktskis, 1988) re referred t fhe lte Villnyin nd t the lte Vilifrnchin (Olivl.U.) (n the bsis f the elicl cntext f the depsits nd the lre mmmls ssembles). Stiil in the Upper Vldrn (lclity Cse nfern, Tuscny), in depsits cntinin lre mmmls f the Tss.U. (lte Villfrnchin), there ws n islted find f Mimmys sini (:Crmerm.ys fter Nerudeu et l., 1995), species tht mrks the beinnin f the Bihrin (Trre, 1985; Msini &Trre,1,987). At Pietrfitt (Perui, Umbri) the remins f vles were fund sscited with lre mmmls f the rnet.u. (lte Villfrnchin): Mimmys pusillus nd the first frms knwn in tly f the subenus Micrtus (Allphimyt), M. (A.) ex r. M. (A.) rffii nd M. (A.) cblinei. The ssemble includes typicl species f the erly Bihrin (Gentili et 1.,7996). At Pirr Nrd (i, Apuli), the type lcliry f the Pirr.U. which is the lst.u. f the lte Viilfrnchin, n bundnt smll mmml fun ws fund sscited with lre mmmls which included M. (A.) cf. rffii nd the first ccurrence in tly f the shrew Crcidur nd f mdern type f muse f the subenus Apdernus (Sylvernus)(De Giuli et 1.,1987; Msini & Sntini, 99l; Abbi et L, 996b). The funs frm Plen (LAquil, Abrun) (Ktskis, unpubl. dt) nd Sve Cv Sud (Vern, Venet) (Ps, 1947 Brtlmei, 1980) cn ls be referred t the Pirr.U. All these micrmmmls ssembles indicte the erly Bihrin. Glerin Mmml Ae. n the type lclity f the first Glerin.U., Clle Curti (lr4rche), tether with lre mmmls there 'were the remins f Micrtus (Allpbirrryy' sp., which is prbbly primitive frm f the Micrtus ecn?nus rup, nd Plim.ys lenki Qrre et l., in press; Abbi Ec Msini, unpublished dt). The fun frm the krst depsit f Mnte Peli (Orviet, Umbri) cn be mributed t the sme.u., ithuh it my be slihtly lder. At this site there re tw fssiliferus levels, but the funl cmpsitin f ech is very similr. The rich ssemble f smll mmmls includes the lsr frms knwn in tly f the enus Berentendi, Glirulus, unrmjs. The subenus M. (Allphimls) ccurs with the mre evlved frms, M. (A.) nutiensis (Micrtus sp. A, in vn der Meulen, 1973) nd lr[. (A.) burndie (Micrtus sp. B in vn der Meulen, 1973). Mirnmlts sini, Mimrnys blnci nd Plimys episcplis re ls presenr (vn der Meulen, t973). The Clle Curti nd Mnte Peli funs indicte rhe finl phse f the erly Bihrin. An ssemble cmpsed f nly tw rdents, Prlurus pnnnicus nd (?) Predicrstnyx sp., sscited with mlluscn fun indictive f cld climtic cnditins, ws fund t ntinn (Pnte Gleri, Rme, Ltium). This fun cmes frm the clys beneth the cnlmertes where the Pnte Gleri lre mmmls were fund nd crrespnds with the finl phse f istpe ste 22 (Ktskis et l., 1992). n the type lclity f the secnd.u. f the Glerin, the Slivi.U. (Aurisin, Trieste, Yeneí Giuli), sme species f micrmmmls were fund s well s numerus lre mmml remins. n the brecci t this site there were Mimmys sini nd Dinrm),s sp., nd the first ppernce f Allcricetus burse nd Micrtus (Nedn) relides (Ambrsetti et l., 1979). This ssemble cn be referred t the lte Bihrin. The fun frm the krst fill t Mnte Tend (Sve, Vern, Venet) belns t the sme Mmml Ae (Lte Bihrin). t is chrcterised by the ccurrence f Mimmys sini nd Plimys episcplis nd by the first ppernce f the subener Micrtus (Micrtus) nd Miutus (lerricl) (Ps, 1947; Brtlmei, 1969). At the type lclity f the fllwin Glerin.LJ., serni (lr4lise), bth lre nd smll mmml remins were fund. n the fluvilcustrine depsits

10 376 E. Glii et l. Plimls episcplis, Plimys lenki, Miutus (Miutus) ff. M. (À4.) rlis nd Micrtus ([erricl) r. mwltiplex-swbterrneus ccur tether with Clethrinmys r. lrelus, Micrtus (bermys) brecciensis nd smll vle, nr very hypsdnt, with mimmin enmel nd t lest ne rdicted M1, which hs been clssified s cf. Aruicl cntin (Cltrti et l.,!982; Sl, 1983; Sl, 996). The fun f Vldemin cve (Bri Yerei, Svn, Liuri), where M. (.) brecciensis is very cmmn nd where new species f lmrph ws fund, Oryctlws buri (Sl, 1992; Ncchi & Sl, in press ), belns t the sme.u. but t different climtic phse. Bth the serni nd Vldemin funs my be referred t the beinnin f the Trinin. The fun f Ripr A di Vislin (Trieste, Venei Giuli), with its typicl lcil fetures, is prbbly mre recent thn the funs described bve, with Plim.ys episcplis, Dinrm.ys bdnwi, Anticl cntin nd Allcricetus burse nd the first ppernce f Micrtus (Stencrnius) relis, Sicis sp. nd Ochtn cf. O. pwsill. The fun cn be referred t the erly Trinin nd it is prbbly lcted between istpic stes L4 nd 10 (Brtlmei & Tí, 1,978; Cttni et., 1.991; Ti, 1995). The fun frm the fluvilcustrine depsit t Ntrchiric (Vens, Pten, Bsiiict) cn be ttributed t the sme perid; bether v/ith Micrtus fí. M. rlis which is dminnt, rhere re ls the remins f Plirnys episcplis, Ar"uicl cntidn nd Cbinm.ys nilis (Sl, 1991). inlly, tw funs cn be referred by indirect crreltin t the ntn Rnucci.U., the lst.u. f the Glerin: tht fund in the ssiferus brecci t Bscchiesnuv (Vern, Venet) vith vles such s Plimls episcplis, Plimys lenki, Dinrrnys bdnwi nd shrews (Srex runtnensis, Crcidur ri) (Brtlmei & Ps, 1970), nd tht fund in the krst depsit t Sn Givnni di Duin (Trieste, Yeneí Giuli), where Plimys episcplis, Dinrmys bdnwi, Micrtus (terrícl) stii, Alluicetus burse nd ther species ccur (Brtlmei, 1976).n bth depsits there is lre sricine, prbbly Episriculus sp.. The funs frm these depsits re referred t the finl phse f the lte Trinin. Aurelin Mmml Ae. The fun frm the uter levels t Grtt di Sn Bernrdin Mire (Clli Berici, Yicen, Venet) is chrcterised by the presence f Mrm mdrrnt, Cricetus cricetus, Dinrmys bdnui nd Micrtus r. redlis-restis (Brtlmei, 1960). These levels hve been referred t the penultimte lcitin by the rdimetric methd U/Th (luères et l., t996) nd prbbly crrespnd with the Vitini.U. f the Aurelin Mmml A.. The fun frm the krst fill depsits t Sn Sider 3 (lr4lie, Lecce, Apuli), where Micrtus (Terricl) svii ís the dminnt species sscited with Micrtus rlis, Lepus eurpeus nd Oryctlus cuniculus (De Giuli, 1983; Ncchi & Sl, in press b), cn be referred t the lst nterlcil. There is lre number f smll mmml funs which dte t the lst Glcil nd the sires re distributed ll ver the tlin peninsul. Durin this perid tly represented refue re fr mny species f mmmls, nd fr this resn in sme lclities temperte frms survived durin the phses f mximum climtic deterirtin. The erphicl nd mrphlicl [etures f the tlin peninsul resulted in subdivisin int res chrcterised by very different ciimtes nd lndscpes. Therefre the smll mmml ssembles vry nt nly s result f the climtic chnes in iven re but ls vry sinificntly s resulr f the erphicl lctin nd ltitude. Ntble differences existed between the Adritic side f the peninsul, which ws chrcterised by clder nd mre rid Mediterrnen climte, nd the Tyrrhenin side which ws mre temperte, nd between the funl sscitins frm cstl nd fthill res nd thse fr the muntins. The Lst Glcil smll mmml funs re enerlly chrcterised by reàrer bundnce f Micrtus (Micrtus) rplis nd Miutus (Micrtus) restis cmpred with the ther vles nd the mice f the enus r4pdemus (e.. nrn, i; Cpss Brbt et 1., 1.992;Petrni et l., 1996). Drmice re represented by Myxus lis, Muscrdinus ellnrius, Elimys quercinus nd DrTmys nitedul (which were ls fund t Grtt Breuil - Mnte Circe, Ltium; Ktskis, 1991; Alhique et l.,!995 - lclity situted between the tw res f tly where it is currently fund) nd re usully very rre. Amn the vles, Micrtus (Tenicl) lwys ccurs in the Suthern tlin reins which represented refue ^re, nd it becme mre bundnt durin the mre temperte phses (e.. t Grtt di Cstelcivit, Slern, Cmpni, Cini et 1.,1.979; Msini tt Abbi,1997). Amn the insectivrus, besides the nerly cnstnr presence f Tlp nd Erinceu.s, the shrews f the enus Srex re reltively widespred, while Crcidur seems t be bsent frm Nrthern tly. n the Nrth tlin sites nd thse f the Adritic cst there re sprdic ccurrences f species with "steppic" ffinities durin clder nd mre rid perids which cme frm Nrthern r Estern Eurpe. The min species is Micrtus ecntnu.s, numerus ppultins f which re smetimes fund in sites in the P vlley (Znld, 1994). The ccurrence f thers "steppic" species (Sicist cf. S. betwli n nd Ochtn pusill) is very rre nd s fr hs been discvered nly in the reins f Venet nd Mrche (Brtlmei, 1980; Sl, 1990).

11 Bicbrnly f Pli-Pleistcene f mmrnls, mlluscs nd strcds in ltly )/ / Amn the mst imprtnt strtirphic sequences frm the pint f view f the bundnce f the fssil recrd nd the strtirphicl cntinuity is tht t Grtt del Brin (Clli Berici, Yícen; Znld, 1994). The first prt f the Lst Glcitin is well dcumented t Grtt di Vej, Ripr Meen, Ripr dell Ghiccii (Mnti Lessini, Vern) nd the lwer levels f Ripr Tliente (Gren, Vern) (Bn et l., 1991). The recent prt f the inter-plenilcil is clerly dcumented in the depsits t Grtt dell Cl (lr4rin di Cmert, Slern; Brtlmei et. l., 1,975) nd Grtt di Cstelcivit (Mnti Alburni, Slern; Cini et 1., 1,979; Msini & Abbí, 1997) ín Suthern tly, while the Lte Plenilcil nd the Lte Glcil re well dcumented in mny sites: these include the upper levels f the depsit t Ripr Tliente (Gren, Vern), Grtt Plicci (Rinn Grnic, i; Brtlmei, 1975), Grtt dell Serrtur (Mrin di Cmert, Slern; Bertlini et l., in press), the upper levels t Grtt dell Cl $4rin di Cmert, Slern) nd Grtt di Pri Mre (Csen, Clbrí; Cpss Brbt & Glíi, 1995). Mlluscs. Key fr the distributin chrt f the nn-mrine mlluscs. The Pli-Pleistcene ssembles f lnd nd freshwter mlluscs f the tlin peninsul re chrnliclly relted t the mmml es shwn in the mmml distributin chrt in this pper. Only extinct species nd thse livin species which hve sme bichrnlicl sinificnce in cnnectin with their ppernce nd extinctin in tly hve been tken int cnsidertin. n the distributin chrt the species f mlluscs re pltted inst their site(s) f prvennce which, in turn, re rdered ccrdin t the mmml es, since mst f the sites re the sme fr bth the mlluscs nd the vertebrtes. n the cse f sites which re imprtnt frm mlclicl pint f view but lck vertebrtes, the chrnstrtirphic psitin ws btined by mens f crreltins with mrine successins, plemnetic dt, istpic stes r with ther cntinentl mlluscn funs. The first prt f the chrt, which cntins the list f cevi sites rrned in rder, ives the impressin fr sme species f mre extensive strtirphic rne thn is ctully the cse, while fr thers their ppernce is scled ver time. The secnd prt f the chrt, which is relted t the mmml es nd nt t the sites, ives mre relistic picture f the strtirphic distributin f the species: clumn L cntins the species f depsits referred t the erly Villfrnchin Mmml Ae; clumn 2 shws the lte Plicene species which cn be crrelted with the middle Villfrnchin Mmml Ae; species frm sites referred t the funl units f Olivl, Tss, rnet (te Villfrnchin Mmml Ae) re listed in clumns 3, 4 nd 5 respectively; species relted t the Glerin Mmml Ae re shwn in clumn 6 nd, finlly, clumn 7 rups the species f the Aurelin Mmml Ae. Villfrnchin Mmml Ae. Erly Villfrnchin Mmml Ae. The nn-mrine mlluscs f this mmml ^e re referred t the Middle Plicene nd t the Trivers.U. These ssembles re chrcterised by n bundnt stck f species, 35 in number, belnin minly t lnd pulmntes, tw thirds f which d nt seem t rech the Middle Villfrnchin. Whilst sme species re limited t the Piedmnt bsin (Viiprus pllneri, Emnterici plicenic, Discus pntnellii, Jnulus nustiumbilictus, Retinell (Lyrdiscus) cf. R.(L.) jurdni, Tiiptychi mstdntphil, etc.), thers re ls widespred in centrl tly (Lminifer (Lminiplic) villfrnchin, Crychium (Srphi) pseudtetrdn, Neulus illfrnchinus, Gstrcpt (Albinul) cumint fssnensis, Gstrcpt (Wrtipsis) debrni, kistyl ttschicki, Estrbilps lisii, Plelndin lunensls, etc.) (Esu & Girtti, 1991; Cinhertti et., 1996). Scc (1887, 1892) cites numerus ther tx frm Plicene sediments t vrius sites in the Piedmnt bsin (Tssrl, Ceresle d'alb, etc.). Jnfrtunteiy the exct strtirphic psitin f these sites is unknwn nd therefre these species re nt listed'in the chrt. Nevertheless the e my be cnsistent with the Trivers.U. (see ls Esu et., 1993).t is impssible t cmpre these ssembles with lder nes (Ruscinin, Erly Plicene) becuse f the lmst cmplete bsence f dt n mlluscs frm this perid resultin frm the scrcity f cntinentl sediments in itly which dte t the Lwer Plicene (Esu, 1982). rm plebierphic pint f view sme species f the Piedmnt bsin re ls present in the bsins f L Bresse (Rhne, rnce) nd in the Rhine (Germny) (Esu et 1.,1993). Middle Villfrnchin Mmml Ae. Tw Umbrin sites, Cv Tppetti ("upper beds') nd Dunrbb, hve been referred t the Lte Plicene (middle Villfrnchin Mmml Ae) fr vrius resns. At Cv Tppetti the lwer leve1s, which re prt f the ss Binc rmtin (Bsilici, in Ambrsetti et l., 1995 b) nd cntin Prsstheni sp. n. nd Zilcheuchilus sp. n,, hve been referred t the Middle nd Lte Plicene n the bsis f plemnetic dt (frm Guss t Reunin, see Albinelli & Nplene, in press); the tw

12 378 E. Glii et l. MOLLUSC BOCHRONOLOGY m r ḡ T f t -m r t - i m -t c ī m r t t : ]' c r m r T t l m t q Tm = i. 3 - Bichrnly f the extinct nn-mrine mlluscs f tlin Peninsul frm Middle Plicene t Lte Pleistcene. The sterisc indictes selected livin specres.

13 Bichrnly f Pli-Pleistcene f mmrnls, mlluscs nd strcds in ltly 379 Prsbrnchi mentined bve cme frm the levels with Guss nrml mnetic plrity. The "upper beds"(: Pnte Nj rmtin, Bsilici, 1995) re referred t the Lte Plicene (middle Villfrnchin Mmml Ae) by Abbi et l.(1996) n the bsis f their mmml cntent. The mlclicl ssemble in this frmtin is quite different frm tht f the "lwer beds" nd the species re mstly cmmn t thse f the Dunrbb site where the sediments beln t the tp f the ss Binc rmtin (Ambrsetti et l., 1995b). The Prsbrnchi f the tw ssembles frm limited stck f freshwter species which rech the erly Pleistcene in the Tiberin bsin, wheres the lnd pulmntes dispper t the end f the Plicene prt frm G(V) dehmi which hs recently been recrded in Upper "Villfrnchin" sediments ner ihille (Anhiri, Are) (unpubl. dt). Lte Villfrnchin Mmml Ae. Nn-mrine ssembles frm severl sites, mst f which re in centrl tl;r, re referred t the lte Villfrnchin Mmml Ae, divided between the vrius.u. f the mmmls. The rich micfunl cmplex is cnstituted by numerus species f prsbrnchs nd pulmntes (Esu & Girtti, 1975; Cnti & Esu, 1981; Esu 6r Girtti, 1991; Ambrsetti et l., 1995; Cinhertti et l., in press b). Amn the pulmntes sme Plicene species survive: N. villfrnchinus, P lwnensis, Ebni ermicwlri survive up t the Olivl.U. whrle Crychium (Crychiell) puisseuri nd G(V) dehmi suwle up t the Tss.U. The ltter species ws ls fund in qurry ner ihille sscited with remins f bvid, Leptbs merli-furtir.tr (pers. cm.. Msini). The prsbrnchs re the mst chrcteristic rup f the lte Villfrnchin Mmml Ae: ener (such s Stepbni, Prsstlteni, Turnuerin, Neumyri) nd endemic species f centrl tly (Tyrrhenin side) which dispper t the end f the Erly Pleistcene. A smli rup f species is f Plicene rià (Thedxus (Nerite) rynus, Prsstheni pdt, Turnuerin belnensis, Micrmelni (Gnichilus) itteli, Emmerici umbr, Melnpsis ffinis). Sme species f prsbrnchs nd pulmntes ccurrin in the Tss.U. d nt seem t be present in the rnet.u.: 7 (N.) rynus, Stephni brnni, Prsstheni menebinin, Tànusi litblyphides, Neumyri priscille, M. (G.) itteli, C.(C) puisseuri, G(V) de h rni, A n cy lus prmpb rus. Tw species, Lithlyphus ff. L. jhni nd Thedxus ff. T dnubilis, ccur fr the first time in sediments f the Stirne nd Crstl Rivers (nrthern tly) f Emilin e; these species survive in the sme re vp t the Glerin Mmml Ae. The Stirne nd Crstl Rivers hve n Adritic drine, but the Emilin sediments they cut ls cntin sme elements chrcterisin the lte Villfrnchin bsins f the Tyrrhenin side, such s T (N) rynus nd Viiprus mpullceus (Crispin & Esu, 1995; Cinhertti et l., in press, ). The Leffe bsin in Nrthern tly (Berm) hs cmpletely different fun, with still livin species nd nly ne exrincr species, Vl"tt chlinei, which hs been fund utside f tly in the Lte Plicene nd Erly Pleistcene f the L Bresse bsin (rnce) (Esu et l., L993). Glerin Mmml Ae. The mlcfunl cmplex referble t the Glerin Mmml Ae is chrcterised by minly mdern prsbrnchs nd pulmntes; the Villfrnchin species re cmpleteiy extinct nd nly few f the Glerin species. Of these, s fr is we knw; L.ff. L. jhni nd T Íf. T dnubilis re nly present in the upper beds f the Stirne sequence (erly Glerin Mmml Ae); Tbedxus isseli nd Jmini mltestl extinuish durin the Aurelin Mmml Ae (Esu & Girtti, 1991). The extinctin f severl Villfrnchin endemic tx f centrl tly is due t the climtic nd elicl histry f these bsins t the end f the Erly Pleistcene. At the beinnin f the Glerin Mmml Ae litypic ssembles similr t thse recrded in the less depsits f centrl Eurpe (Lek, 1964) pper in centrl tly (Ktskis et l., 1992). They re cmpsed f species which nw live in nrthern cuntries (s Helicell itl) r f species still livin in tly but which re less cmmn r widespred in nrthern tly (s Tiichi hupid). t is cler tht the nn-mrine mllusc ssembles f the Middle (nd Lte) Pleistcene re influenced by climtic scilltins. Generlly the plitypic ssembles re relted t temperte nd temperte-wrm climtic phses, whilst durin cld perids strictly litypic ssembles re recrded (Esu et l., 1989). Aurelin Mmml Ae. The trnsitin frm the Glerin mlcfuns t the Aurelin nes is nt detectble. Assembles f cld-temperte nd wrm-temperte chrcter similr t thse f the Glerin Mmml Ae cntinue t lternte. They re minly cmpsed f livin species. n sme Aureiin depsits f centrl-suthern tly vrius extinct species re recrded Hydrbi melii, Tnwsi subt, Belrndi ilchi (Esu & Girtti, 1991; Esu & Ktskis, i996). Tw species, T isseli nd J. mltesti, which were present in the Glerin ssembles, becme extinct durin this intervl. At the beinnin f the Aurelin Mmml Ae, durin wrm climtic scilltin, trpicl iivin species, Melnides tubercul (see trsu 6c Girtti, 1991), ws present in the suthern prt f the tiin peninsul, prbbly trnsprted there by

14 380 E. Glii et l. birds. n enerl the Aurelin mlclicl ssembles re mre useful fr ecstrtirphic studies thn fr bistrtirphic r bichrnlicl purpses. presence,/bsence in ne prticulr depsit des nt represent nly bichrnlicl dt but must be filtered thruh n envirnmentl interprettin. n fct it is knwn tht strcds re used fr bth the definitin f clssicl nes inside cnventinl bistrtirphy (r bichrnly), tkin int ccunt the sme envirnmentl cnditins (e.. littrl mrine envirnment bines), nd fr the cretin f eclicl cmmunity units (r ecnes) inside n ecstrtirphy (Mrtinssn, 1973).n bth cses, in rder t vid the vertici distributin f the tx bein ltered by eclicl bckrund nise, it is necessry t wrk with lre number f sites pertinin t different envirnments fr ech time perid. Despite the scrcity f sites where strcds hve been nlysed s fr in tly, i. 4 shws tenttive chrnlicl rderin f tlin strcd funs knwn frm freshwter nd brckish (li-meshline) envirnments. {ere, the presence f severl selected tx frm different tlin depsits is reprted. Only thse species nt recrded prir the Middle Plicene hve been tken int cnsidertin; lmst ll these species re still livin in the tlin Peninsul, s i. 4 is nly indictive f their first ccurrence. The sites re referred t the Mmml Aes nd, when pssible, ls t the unl Units which represent the dtum-pint fr the Pli-Pleistcene bichrnly. t is imprtnt t pint ut tht this tble represents nly strtin pint nd tht ther studies n different strcd funs re necessry in rder t mke ut distributin chrt which culd be f use fr bichrnlicl purpses. Ostrcds. Distributin chrt. Studies n the tlin Pli-Pleistcene cntinentl (fresh wter nd brckish) strcd funs re scrce. Only in the lst few yers prticulr ttentin hs been pid t their nlysis, since they prved t be very useful fr detiled pleeciicl recnstructins. {wever, t the mment it is still nt cler whether strcd funs re ls vlid tl fr Pli-Pleistcene bichrnly, minly fr tw resns: 1.) in severl cses the life f the species is much lner thn the time perid under cnsidertin (fr exmple, nd nly t cite sme species which re very frequent in Pli-Quternry fresh wter depsits, Cyclcypris levis is knwn frm the Dnin t the present dy, Druinul steensnl frm the Olicene t the present dy, nd Cndn nelect, Ptmcypris ful nd Pseudcndn mrchic frm the Lte Micene t the present d^ù; Z) strcds re benthic rnisms which re prticulrly sensitive t the envirnmentl chrcteristics, nd cnsequently their Villfrnchin Mmml Ae. There re few sedimentry series berin strcds referble t the erly nd middle Villfrnchin Mmmi Aes: the R.D.B. qurry (Villfrnc d'asti, Piedmnt), Arndelli nd Rtt (Piedmnt) (Ghi, 7996) referble t the Trivers.J., Cv Tppetti-ss Binc Unit (Tdi, Umbri) (prbbly referble t the Mntpli.U.), Cv Tppetti-Pnte Nj Unit (Cst S. Gicm.U.) (Glii, unpubl. dt) nd, finlly, Dunrbb (Umbri) which is referble, n the bsis f the mllusc ssemble (Ambrsetti et l., 995), t eneric middle Villfrnchin (Gíì, unpubl. dt). All these sequences pertin t limnetic dmin, except fr Cv Tppetti-Pnte Nj Unit which recrds brckish envirnment. n this wide time intervl, crrespndin t the Middle nd Lte Plicene, Cndn cndid, Stencypris sp. Cndn fbefrmis, Prlimnrythere cmpress, Pseudcndn prllel, Cyprideis sp. 1 nd Cypridpsis pr"u seem t pper. All these species chrcterise the mdern tlin strcd fun, except fr: 1) Stenqpris sp., enus t present wideiy distributed t trpicl ltitudes, which ws fund s fssil t the R.D.B. qurry; t present it is recrded s bein present in tly by Ghetti & McKenie (1981) represented by the species S, fntinlis e S. mjr nd is nly fund in lkes with ht wter input. Accrdin t McKenie & Mrni (1986), these frms cn be cnsidered "frein uests", thr is frms which hve been ccidentlly intrduced by mn with frein d trnsprts;2) Cyprideis sp. L, n extinct frm fund nly t Cv Tppetti-Pnte Nj Unit, which seems t be limited r the Lte Plicene. This frm pertins t the Cyprideis trs rup, but differs frm C. trs in severl mrphmetric chrcters such s its dimensins nd enerl shpe (Glii, unpublished dt). Three mre species, Limnrythere ff. L. inpint, Metcypris crdt nd Pseudcndn cmpress, were recrded by Di Npli (1962) in the linite depsits t Mrnn, Trin, Spin, lci, nd Cllne (Tiberin Bsin), which he referred t the Lte Plicene. At present these depsits re n lner visible, nd cnsequently strtirphic cntrl usin the recent Pli- Pleistcene chrnlicl sequence is impssible; these species hve therefre nt been put int the sbck f species which ppered durin the Middle-Lte Plicene. The depsits referble t the lte Villfrnchin Mmml Ae (Erly Pleistcene) re mre numerus, but they re minly lcted in centrl tly: severl lclities in the Tiberin Bsin (Umbri) [Bstrd nd,

15 Bicbrnly f Pli-Pleistcene f mmmls, mlluscs nd strcds in lly Derut nd Cllne, enericlly referred t the first prt f the Erly Pleistcene; Bstrd, ry V nd V, referble t pst-snternin (Di Npli, 1962), nd Vill S. ustin (Ambrsetti et l., 995b) which cn be crreited with the Tss.U.], Pietrfitt (Umbri) (Glir, unpublished dt), severl lclities f the Rieti Bsin (Ltium) [Cse Strinti, ss ilundici, Aplei, Vicchinne, crrelted with the Tss.U. nd Belmnte Sbin, prbbly referble t yuner Erly Ch rnstrtirphy PLOCENE MDDLE LATE EARLY PLESTOCENE MDDLE PLESTOCENE LATE )LESTOCENE Mmml es Erly Villf rn. Middle Vilìf rn. Lte Villfrnchin Aurelin lr È l Pleistcene (pst-snternin) (Brberi et l., 1995)l nd Mntller (Sicily) (Decim, 1963) crrelted with the Emilin. The mjrity f these series recrd brckish, li-meshline envirnments. Durin the lte Villfrnchin Mmml Ae the present livin species f Cyprideís trs, Cytberiss lcustris, Herpetrypris chereuxi, Cndn lenderi, Lirnnclthere snctiptricii nd Pmcypris schkkei ll seem t pper. Mrever Limncythere inpint, Metcypris crdt nd Pseud- tr î '- 6 u) U c l c.6 c0 q È uj fl 0- d) É. f ii q () c 6 '6 c (L uj fl l 6 U) ff) É. u c.ì t! q c tl _ U) c É. U î lr q l.9 c0 LJ cc î lt =r tr E. l ul J J ts cc C' ts (-) (r L J À = Cndn cndìd c{. -Stencvpris sp, Cndn fbefrmis Prlimnc!,there cmpress cî. cî. t uvpnders sp. 1 Cypridpsis pru vprideis trs Lr m nc!,the re rnpr nl 'Metcvris crdt C)4heriss lcustris HerDetcvDns chevreuxl c cf. Cndn levnderì Limncvthere snclrtricli Plrncvris schkkei cl Cndn pubescens Cypridpsis vidu t Leptcythere (Amnicythere) tllx '"Cndn cudt '"Cndn lbipes 'Sctti brwnin ""tucypns prr uypncercus relrcurtus Princypris enkeri Cypris puber rltmncvînere messnl Eucyprìs clvt Eucvpris serrl '6 = 6 m U cc f ' ō c cc!! ts lr l-.; U O = - J u U Í. ( E r.e.p tr - Ò. t (, lt) uj J N ) tt 5 'c! f.- 6 É = tr fr È È.. 'c! U = tr U) O LU E E -c N c 'È (, l J.D 5 'È c tḡ _) tr - b (t) 6 5 c Lrl =q U) O - t! J cl u, c c uj =(, - U J) i 4 Bichrnly f selected livin nd extinct brckish nd freshwter strcds f tlin Pleistcene. Leend: 'f species t present distributed t circumtrpici ltitudes; Peninsul frm Middle Plicene t Lte species t present livin ut f t y t hiher ltitudes; species t present widely distributed t hiher ltitudes; + estinct species.

16 J82 E. Glii et l. cndn c?rrpress re nw recrded with certinty. Vithin this rup, the presence f Cyprideis trs. needs t be explined better: Cyprideis trs ís recrded in the literture s bein frequent in the Eurpen Plicene, nd smetimes it is reprted s species while t ther times it is reprted s C. r. trs. t is nt pssible t exclude the fct tht this species my re y hve ppered durin the Plicene, lthuh ll the tlin depsits in which its presence hs been cntrlled re f Pleistcene e. Aurelin Mmml Ae. At present n studies n strcd funs referble t the Glerin Mmml Ae hve been crried ut, while there re numerus studies n Aurelin depsits, unfrtuntely ll f which re lcted in Ltium (centrl tly): Trre in Pietr (Miinri Pnelli, 1978) (crrelted with Oxyen stpe Ste 9), Pinur Pntin (lr4ilir 47 brehle) (Brbieri et 1., in press) referble t Stes 9,?7, 6 nd 5, severl lclities ln the Liri Vlley (Devt, 1965) referble t Stes 9-8 nd, tentti vely, 7, Vlle di Cstiline brehle, referble t Stes 7,5 nd 3 (Glií &Mini, in press), nd L Lun brehle, referble t Ste 5 (Clderni et., 1994). The species tht seem t pper t the beinnin f the Aurelin Mmml Ae re numerus, nd this is prtly due t the imprtnt txnmic studies which hve been crried ut n Aurelin strcd funs by Mlinri Pneili (1978) nd Devt (1965): Cndn pubescens, Herpet cypris reptns, Cypridpsis vidu, Leptqttbere fllx, Cndn cudt, Pseudcndn lbipes, Sctti brunin, Eurypris pir, Prinrypris enkeri, Cypricercus reticultus, Herpetcypris intermedid, Psychrdrmus licews, Cypris puber nd Prlimncythere messni. Actully, this nticeble bundnce f frms is minly due t the fct tht sme depsits f the Liri Vlley crrespnd t cld perids (stpe Ste 6) nd mny f the frms which ccur durin this ste re nrthern frms, presently fund in centrl nd nrthern Eurpe (Sctti brnin, Herpetqtpris intermedi e Psycbrdrmus liceu) (which cn be cnsidered s "cld uests") r frms which, lthuh they re still fund in tly (nly in scttered lclities f Piedmnt nd Lmbrdi), shw their min distributin relcted in mre nrthern res (Cndn cwdt, Cndn lbipes, Eucypris pi).tkin inr ccunt the fct tht the Liri Vlley strcd funs re the nly cld srrcd ssembles knwn in tly, it is nt pssible t scertin whether these cld species entered tly nly frm Ste 6 nwrds, r whether they mirted suthwrds durin ther erlier cld perids. t is interestin t underline the presence durin the Aurelin Mmml Ae f n extinct frm, cptcytbere fllx, nd the first ccurrence Í Pr imncythere mess ni. The beinnin f the lte Aurelin Mmml Ae (stpic Subste 5e) recrds the ppernce f the present-dy livin species Pseudcndn srsi, Eucypris clt nd, perhps, Eucypris sey'rt, chrcterised by hvin niy ne summer reprductive sesn (Brnshtein, 1e47). Achnuledemmts. This pper ws supprted by CNR rnts, Centr Studi per il Quternri e l'evluine Ambientle, Rin. REERENCES Lbbt L. (1995) - Mecerides bscurus frm the Vl di Chin (Cv Libertri, Tuscnn Centrl ltly, rnet f. u., erly Pleistcene). Remrks n erly evlutin nd systemtic f Mecerides. Bll. Sc. Plent. tl., v.3 Q), pp , Mden. Abbí L.& Msini. (1,997) - Mecerides slilhcus nd ther deer frm the middle Pleistcene site f serni L Pinet (Mlise, tly). Bll. Sc. Plent. tl., v. 35 Q) 1996, pp , Mden. Abbi L., Ambrsetti P., Arenti P., Bsilici G., Bertini A., Gentiii S., Msini. Ec Pntini M.R. (1996) - Plentlicl nd sedimentlicl recrds in Plicene distl lluvii fn depsits (Cv Tppetfi ner Tdi, Centrl tly). Abstr. X Meetin Sc.Plent.1t., Prm 7A-13 Sept. 1996, pp. 1-3, Prm. Abbr L., Benvenuti M., Bschin G., Dminici S., Msini., Mebtt C., Rk L., Vlleri G. & Trre D. (1996b) - The Neene nd Pleistcene successin nd the mmml funl ssembles f ^î re between Apricen nd Pi mperile (i, tly). Mem. Bll. Sc. Gel. t., v. 51, pp , Rm. Lbbi L., Albinelli A., Ambrsetti P., Arenti P., Bsilici G., Benini A., Gentili S., Msini., Nplene G. & Pntini M.R. (in press ) - Plentlicl nd sedimentlicl recrds in Plicene distl lluvil fn depsits f Cv Tppetti (Tdi, Centrl hly). Bll. Sc. Plent. tl., MderL^. Abbi L., Deld Huerts A., cumin P., Lninelli A., iccrelli G., Msini. & Trre D. (in press b) - Mmml chnes nd istpic biechemistry. An interdisciplinry pprch t climtic - envirnmentl recnstructins durin the lst Pleini lc i / Lte-Gcil ::'r'- sitin in the Plicci cve sectin (Grn, Apuli). 1/ Quternri, rrr'. Auirre E. & Psini G. (1985) - The Plicene-Pleistcene bundry. Episdes, v. 8, pp A, Ottw.

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