Palaeoenvironment and palaeoclimate in the western Liguria region (northwestern Italy) during the Last Glacial. The small mammal sequence of Riparo Mochi (Balzi Rossi, Ventimiglia)

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ContentslistsavailableatScienceDirect

Comptes

Rendus

Palevol

www . s c ie n c e d i r e c t . c o m

General

Palaeontology,

Systematics,

and

Evolution

(Palaeoecology)

Palaeoenvironment

and

palaeoclimate

in

the

western

Liguria

region

(northwestern

Italy)

during

the

Last

Glacial.

The

small

mammal

sequence

of

Riparo

Mochi

(Balzi

Rossi,

Ventimiglia)

Paléoenvironnement

et

paléoclimat

en

Ligurie

occidentale

(Nord-Ouest

de

l’Italie)

au

cours

de

la

dernière

période

glaciaire.

La

séquence

de

petits

mammifères

du

Riparo

Mochi

(Balzi

Rossi,

Vintimille)

Claudio

Berto

a,b,c,∗

_,

_Fabio

_Santaniello

d,e

_,

_Stefano

_Grimaldi

d,e

a_Università_degli_Studi_di_Firenze,_Dipartimento_di_Storia,_Archeologia,_Geograﬁa,_Arte_e_Spettacolo_{(SAGAS)–Archeologia}_preistorica,_Via

S.Egidio21,50122Firenze,Italy

b_Museo_e_Istituto_Fiorentino_di_Preistoria,_Firenze,_Italy

c_Università_degli_Studi_di_Ferrara,_Dipartimento_di_Studi_Umanistici,_Sezione_di_Scienze_preistoriche_e_{antropologiche,}_C.so_Ercole_I_d’Este,

32,44121Ferrara,Italy

d_Università_degli_Studi_di_Trento,_Dipartimento_di_Lettere_e_Filosoﬁa,_via_Tommaso_Gar_14,₃₈₁₂₂_Trento,_Italy e_Istituto_Italiano_di_{Paleontologia}_Umana,_Anagni,_Italy

a

r

t

i

c

l

e

i

n

f

o

Articlehistory:

Received4January2018

Acceptedafterrevision29April2018 Availableonline4July2018

HandledbyLarsW.vandenHoekOstende Keywords: UpperPleistocene Paleoclimatology LateGlacial Gravettian Insectivores Rodents

a

b

s

t

r

a

c

t

RiparoMochiisconsideredakeysitefortheMiddle–UpperPalaeolithicchrono-cultural sequenceintheMediterraneanarea.Climaticandenvironmentalconditionshavebeen reconstructedafterthestudyofthesmallmammalassemblagebymeansoflinear regres-sionmethodandtheHabitatWeightingmethod.TheclimateproxiesforthelateMousterian andProto-Aurignacian(culturalunitsItoG)indicateanenvironmentwitharelativehigh percentageofforestcomponentduringtwocoldandonerelativewarmoscillations.Open environmentswithlowtemperaturesareregisteredfortheGravettian(culturalUnitD) allowingapossiblerelationtotheHeinrich3Event.TheﬁnalGravettian(culturalUnit C)showsarelativewarmpeak,tentativelyrelatedtoGreenlandInterstadial2.Thelate Epigravettian(cultural UnitA)ischaracterizedbyahighpercentage offorest-related environmentsuggesting thatthishorizonformedduringtheBølling–Allerød Intersta-dial.Finally,theRiparoMochisequencecontributestoreconstructtheprocessesoffaunal changesalongthenorthernTyrrheniancoastinaperiodofclimaticinstability,especially duringtheendofMarineIsotopeStage3whenforaginggroupsadopteddifferentmobility strategiesbetweensouthernFranceandtheItalianpeninsula.

Motsclés:

Pléistocènesupérieur Paléoclimatologie

r

é

s

u

m

é

Le Riparo Mochi est un site clé pour la séquence chrono-culturelle Paléolithique moyen–supérieurdelarégionméditerranéenne.Lesconditionsclimatiqueset environ-nementalesontétéreconstruitesàpartirdel’étudedel’assemblagedepetitsmammifères

∗ Correspondingauthor.UniversitàdegliStudidiFirenze,DipartimentodiStoria,Archeologia,Geograﬁa,ArteeSpettacolo(SAGAS)–Archeologia preis-torica,ViaS.Egidio21,50122Firenze,Italy.

E-mailaddress:claudio.berto@unife.it(C.Berto).

https://doi.org/10.1016/j.crpv.2018.04.007

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DernierGlaciaire Gravettien Insectivores Rongeurs

parlesméthodesdemoyennesderégressionlinéaireetHabitatWeighting.Lesdonnées cli-matiquespourleMoustérienfinaletleProto-Aurignacien(unitésculturellesIàG)indiquent unenvironnementcomposé,d’unepartrelativementimportantedeforêtaucoursdedeux oscillationsfroidesetuneoscillationrelativementchaude.Unenvironnementouvertavec destempératuresbassesestenregistrépourleGravettien(unitéculturelleD),permettant unecorrélationpossibleavecl’évènementHeinrich3.LeGravettienfinal(unitéculturelle C)présentedespicschauds,pouvantêtrereliésauGreenlandInterstadial2.L’Épigravettien final(unitéculturelleA)estcaractériséparuneproportionimportanted’environnement forestier,suggérantquecethorizons’estformépendantl’interglaciaireBølling–Allerød.La séquenceduRiparoMochicontribueàlareconstructiondesprocessusdechangement fau-niquelelongdelacôteNord-Tyrrhéniennependantunepérioded’instabilitéclimatique, particulièrementàlafinduMIS3,lorsquelesgroupesdechasseurs-cueilleursadoptent différentesstratégiesdemobilitéentreleSuddelaFranceetlapéninsuleItalienne.

1. Introduction

TheenvironmentandclimatereconstructionsofMIS3 and2ofcentralandsouthernEuropehavebeenwidely dis-cussed(BarronandPollard,2002;Monegatoetal.,2011; Pinietal.,2010aamongothers).Mostresearchfocuseson thepossibleclimaticcausesthatledtotheNeanderthal extinction and that could have driven the diffusion of AnatomicallyModernHumansinEurope(e.g.,Fedeleetal., 2002;Highametal.,2009;López-Garcíaetal.,2015).

Thirteencontinuousandrepeatedclimatecycles,such asDansgaard–Oeschger(DO=GI)andHeinrichEvents(H), havebeenrecognizedbetween50,000and20,000years BPinGreenlandandNorthAtlanticcores(Andersenetal., 2006;Bondetal.,1993;Dansgaardetal.,1993;Rasmussen etal.,2014,amongothers).Bycontrast,continentalrecords supportingtheseeventsarerareandusuallyincomplete, mainlybecauseofhiatusesinsedimentationand/orpoor dating (Spötl and Mangini, 2002). Nevertheless, several studiesallowedustocorrelatetheMIS3climatecycles to changes in the small mammal communities found throughoutthestratigraphicsequences,especiallyinthe Mediterraneanarea(e.g.,Bertoetal.,2017a;Burjachsetal., 2012;López-Garcíaetal.,2012,2014b,2015;Royeretal., 2013)contributingtoreconstructtheclimateconditionsin whichprehistorichunter-gathererslived.

Inthis context, little isknownabout smallmammal variationsinnorthwesternItalyduringMIS3and2, espe-cially in thecoastal region (Berto, 2013).This areahas beenrecentlyrecognizedaskeytobetterunderstandthe land use strategies of the last Neanderthals as wellas thediffusionoftheUpperPalaeolithictechnocomplexes suchas theProtoaurignacian, Aurignacian,and Gravet-tian(GrimaldiandSantaniello,2014;Grimaldietal.,2014,

Santaniello,2016).ThenorthernTyrrheniancoast,the so-calledLiguro-Provenc¸alArc(Fig.1a),formsanarrowlittoral corridor,400kmlongandafewkilometreswide, delim-itedbytheApenninesandthewesternAlpstothenorth, andtheTyrrhenianSeatothesouth,furtherlinking cen-tralTyrrhenianItalytotheRhôneValleyinFrance.During theMIS3,thelandscapeoftheLiguro-Provenc¸alArcwas notvery differentfromtoday’s(Arobbaand Caramiello, 2009;Pons-Branchuetal.,2010;Wattsetal.,2000).This

region was a narrow corridor even during glacial time according to the bathymetry of the sea bottom (IBCM, 2014).Consequently,fromanarchaeologicalperspective, theLiguro-Provenc¸alArcshouldhavebeenanaturalaxis channelling the exchange of both humans and animals betweencentralItalyandsouthernFrance(Porrazetal., 2010).

Here,wepresentfortheﬁrsttimeastudyoftheentire collection of smallmammals (insectivores and rodents) fromtheRiparoMochisite(BalziRossi,Ventimiglia,Italy). Thissiteholdsoneofthemostimportantsectionsinthe Tyrrhenianarea,spanningfromMousterianto Epigravet-tian,anditisconsideredakeysitefortheinvestigations ontheMiddle-UpperPalaeolithicboundary(Doukaetal., 2012; Higham et al., 2014). This paper aims to recon-structthelocalenvironmentandclimatechangesbasedon thesmallmammals.Thereconstructionwillbediscussed taking in account other northern Italy small mammal sequences–suchasGrottadelBroion(Colamussi,2002) andGrottadiFumane(López-Garcíaetal.,2015)–aswell aspollenarchivesfromsouthernFranceandnorthernItaly withtheircorrelationtoOxygenIsotoperecords.

2. RiparoMochi

RiparoMochi(Mochirockshelter)isabroad,shallow rocksheltersituatedintheBalziRossiarchaeological com-plex,inwesternLiguriaontheborderbetweenItalyand France(43◦473.66N,7◦324.18E,30ma.s.l.).Itisoneof severalprehistoricsites,alsoknownas“Grimaldicaves” (deVilleneuveetal.,1919)thatprovided archaeological depositsrangingfromtheearlyMiddletotheﬁnalUpper Palaeolithic(Fig.1a).

Thesite wasdiscovered in1938byA.C.Blancand L. Cardini of theItalian Instituteof Human Palaeontology (IsIPU). They performed ﬁrst a small-scale testing over threetrenches(A,B,C)(Blanc,1938);trenchA,latercalled “CentralTrench”,wasenlargedandexcavated systemati-callyin1941,1942and1949.In1959,anotherexcavation carriedoutbyCardiniyieldedtheUpperPalaeolithic lay-erslocatedattheeasternandwesternsideoftheCentral Trench.Fornearlyfourdecadesthesitewasabandoned, butfrom1995to2005,A.Bietti(LaSapienzaUniversityand

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Fig.1. A:LocationofRiparoMochi(redsquare);locationofthemostcompleteItalianPeninsulasmallmammalsequences(blacksquares);locationof citedpollenrecords(bluedots).B:RiparoMochistratigraphyandselectedradiocarbondatescalBP(originaldrawingfromSegreandmodiﬁedfromBlanc, 1954;foracompleteframeworkaboutradiocarbondatesseeDoukaetal.,2012).

Fig.1. LocalisationduRiparoMochi(carrérouge);localisationdesséquencesdepetitsmammifèreslespluscomplètesdelapéninsuleItalienne(carrés noirs);localisationdesdonnéespalynologiquescitées(pointsbleus).B:StratigraphieduRiparoMochi(dessinoriginaldeSegremodiﬁéparBlanc,1954; Doukaetal.,2012).

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IIPU)resumedtheexcavations.Since2007,theUniversity ofTrento,incollaborationwiththeformerSoprintendenza ArcheologicadellaLiguriastartednewresearchonthissite, whichisstillongoing.Thankstothisproject,new radio-carbondatesofthelowerpartoftheUpperPalaeolithic sequencehavebeenpublished(Doukaetal.,2012).Based onthisevidence,thebottomoftheculturalUnitG (Protoau-rignacian)ofRiparoMochi,datingtoaround37–36.5kaBP, istheoldestdirectly-datedUpperPalaeolithicassemblage inItaly.

Thechronoculturalsequence,approximately10mdeep, consistsofnineunits,mainlyinferredinretrospectupon the characteristics of the embedded cultural remains (Grimaldi and Santaniello, 2014; Grimaldi et al., 2014; Laplace,1977;Tomassoet al.,2014among others),and namedAtoI,fromtoptobottom(Fig.1b).Eachculturalunit wasexcavatedin10to5cmspits.Geoarchaeological obser-vationsontheupper5.5mofthesequence(UnitC–H)allow ustosubdivideitinto3facies.Theﬁrst(Facies1)consists ofthicklayerscomposedoflimestonerubbleina sandy siltyloammatrix;thesecond(Facies 2)iscomposedof siltyclayloamunitswithheartsinterbeddedbetweenthem (Facies3)(Doukaetal.,2012).Thelithological characteris-ticsofthesequenceinassociationwiththesitechronology, alreadydescribed in Doukaetal. (2012),showthat the sequencewasdepositedmainlybetweenMIS3and2. Six-teenradiocarbondatesareavailablefromthetopofthe culturalUnitI(Mousterian)tothebottomofculturalUnit D(Noaillian–Gravettian)providingachronologicalrange between41,630–40,410cal.BPand29,380–28,510years cal.BP.Theupperpartofthesequencehasnotbeendated yetandtherelativechronologyisinferredbylithic evi-dence:theculturalUnitAyieldedlateEpigravettiantools whiletheculturalUnitC,coveredbythesterileUnitB,has beenattributedtoFinalGravettian.

2.1. Previouspaleoenvironmentanalyses

Severalpaleoenvironmentanalyseshave been previ-ouslycarried out on Riparo Mochi. These studies have focusedonthepollensequence(Renault-Miskovsky,1972), large mammals (Alhaique, 2000; Arellano, 2009, 2004; Zeppieri,2009,summarisedinTagliacozzoetal.,2012),and asampleofsmallmammals(Abbassi,1999).

TwelvepollensamplescollectedfromculturalUnitsG andF–E(1959Cardinisequence,Eastsector)allowedus toreconstructtheenvironmentoftheregionatthe begin-ningoftheUpperPalaeolithic(Renault-Miskovsky,1972). Theidentiﬁedpollenessencescamefromplantscurrently livinginthereliefsnexttothesite(startingfrom600m a.s.l.),suchasconifersanddeciduoustrees,andfromtypical MediterraneanplantssuchasmaritimepineandOleaceae, nowadayspresentalongthecoast.Fourenvironment oscil-lationsweredetected:achangefromanopenenvironment (ArborealPollen=5%),witharegionalcoldanddryclimate, toanincrease of wooden areas(AP=40–65%) with ter-mophilicessences(Oleaceae10–30%),suggestingawarmer phaserelatedtotheArcyinterstadial.Othertwo oscilla-tions,testiﬁedonlybytwosamplesinUnitsFandE,show acoldandaridclimatefollowedbyawarmerphase.

The faunalevidence comingfrom Riparo Mochiwas onlypartiallystudiedbyseveralauthors.Arellano(2009)

reported the largemammal species for theMousterian sequence(culturalUnitI)withoutprovidinganydetailed paleoenvironmentandpaleoclimaticreconstruction; how-ever,threeenvironmentalchangeswereobserved.Inthe lowerspits,cervidsaredominantandwildboar(Susscrofa) is very abundant while the presence of roe deer sug-gestsa relativelyhumidclimate.Thisconditionchanges in the medium part of the unit where mammals such asRupicaprarupicapraand Capraibex increaseandwild boar–amongtheforestungulates,themostsensitiveone toclimaticchanges–showsanabruptdecline,suggestinga colderphaseinassociationwithmorearidconditions.In theupperpartoftheunit(i.e.ﬁnalMousterian),theforest faunabecomesdominantinassociationwithDamadama remains.

Alhaique(2000)analysedthefaunalassemblage com-ingfromthe1996–1996Bietti’sexcavation(GtoCUnits). TheUpperPalaeolithicsequence,althoughpoorin mate-rial,providedsomeinformation.Reddeerandroedeerare presentinthewholesequence,butroedeershowsahigher frequencyinunitG.Marmotamarmotaisalsopresentin UnitsCandD;accordingtotheauthor,theonlyindividual foundinUnitFmightbeintrusive.

Amoredetailedstudyofthelargemammalsevidence comingfromUnitD(Gravettian, 1959Cardini’s excava-tion)ispresentedinZeppieri(2009),latersummarizedin

Tagliacozzoetal.(2012).ThefaunalremainsfromUnitD weredividedintophases(fromIVtoI,fromthebottom totop ofthe Unit)that have beenassociatedwith sea-sonalityofseveraloccupations;thissubdivisionshouldno longerbeconsideredcorrectasitwasestablished follow-ingtheartiﬁcialstratigraphydeﬁned byCardinihimself withoutanyviablecorrelationwiththegeo-archaeological evidence(comparewithSantaniello,2016).Broadly speak-ing,atthebaseoftheUnitD(PhaseIV),thefaunalspectrum indicatesawoodedenvironment,withconifersand decid-uoustrees,characterizedbyatemperatetocool,climate. Such anecosystemchangeddue tocoldertemperatures andadryclimateinthecentralphaseofUnitD(PhaseIII andII),wheretheupperlimitoftheforestleftspacefor alpinemeadows.Finally,ontopofUnitD(PhaseI),the for-estcoveragewasthicker,withtemperatetocoolandmore humidclimaticconditions.However,inallperiods,there is acontemporaneous presenceof temperateand forest fauna(Cervuselaphus,Capreoluscapreolus,Susscrofa)that probablylivedintheimmediatesurroundingsofthesite, togetherwithcoldandopenenvironmentspecies(Capra ibex,Rupicaprarupicapra,Marmotamarmota)whichwere presentinmountainareasabovethetreeline.

Asampleofsmallmammalfauna,collectedbythe “Lab-oratoiredépartementaldePréhistoireduLazaret”in1993 (Paunescoet al., 2010), wasstudied byAbbassi (1999). ThesamplewasdividedinculturalunitsfollowingBlanc’s (1954) stratigraphy withthe only exception for theso called“UnitJ”,probablyrepresentingthebaseoftheUnit I. The rodents identiﬁed along the sequence are Mar-motamarmota,Eliomysquercinus,Muscardinusavellanarius, Microtusarvalis,Microtus(Terricola)multiplex,Chionomys nivalis,Arvicolaamphibius,Arvicolasapidus,Clethrionomys

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glareolus, and Apodemus ﬂavicollis, while the specimen identiﬁed as Microtus (Terricola) savii was later revised as Microtus (Terricola) multiplex (Paunesco et al., 2010).

Abbassi(1999)suggestedaMediterraneanclimate,similar topresentconditions,atthebaseofthesequence(possibly Unit“J”),whichchangestowardacolderandcontinental climatebecauseofthepresenceofChionomysnivalisand Microtus (Terricola)multiplex.Theevidencefoundatthe Middle–UpperPalaeolithictransitionindicatesanarid cli-matewhichbecomesmorecontinentalintheupperUnits GtoC.

3. Materialandmethods

ThesmallmammalremainsfromRiparoMochiare rep-resented by disarticulated bone fragments collected by water-screeningusing1mmmeshsievesduringthe Car-dini(1938–1949,1959)andBietti(1995–2005)campaigns. Thesmall mammalassemblageincludesa total of1244 remains,correspondingtoaminimumnumberof795 indi-viduals (Table1, Fig. 2).The speciﬁc attribution ofthis materialwasmainlybasedonthebestdiagnosticelements: mandible,maxillaandisolatedteethforrodents,mandible andmaxillaforshrews,mandible,maxilla,isolatedteeth andpostcranialbonesforTalpidae.

ThetaxonomicclassiﬁcationfollowsWilsonandReeder (2005),exceptforClethrionomysglareolus(forthepriority overMyodes,seeTesakovetal.,2010).Dataonthe distri-butionandhabitatofthespeciesweretakenfromAmori etal.(2008),Boitanietal.(2003)andMitchell-Jonesetal. (1999).

We calculatedthepalaeodiversityusingtheSimpson indexofEvenness=1–

(pi2),wherepiistheproportion

ofindividualsintheith_species₍_Harper,_2005;_Magurran,

2004).Theevennessindexisconstrainedbetween0and 1.TheindexhasbeencalculatedusingPAST3.04avoiding

redundant determinations(i.e. for Microtus agrestis, the individuals determinedas Microtus cf. agrestishave not beenincludedintheSimpsonindexcalculation)(Hammer etal.,2001).ARattussp.ﬁrstlowermolarfoundontop of UnitDhasbeen consideredintrusive fromHolocene sediments,thusthisspecieshasbeennottakeninaccount for paleoenvironmentand paleoclimatereconstructions. Finally,despitethecriticisms outlinedabove,a compar-isonwith theseasonal phases described by Zeppieriin

Tagliacozzoetal.(2012)isalsodiscussed.

3.1. Paleoenvironmentreconstruction

InordertoreconstructthepaleoenvironmentatRiparo Mochi,weusedtheHabitatWeightingmethod(Andrews, 2006;Evansetal.,1981), assigningeachsmallmammal taxontohabitat(s)whereitcanbefoundtodayinEurope. Tothispurpose,habitatsweredividedupintosixtypes (Cuenca-Bescós et al., 2010, 2009; López-García et al., 2014a):openlandwitheitherdryorwetmeadows(OD and OH,respectively); woodland environments, divided intoopenwoodland,woodlandmarginsandforestpatches (OW)andwoodlandandmatureforesthabitat(W);water, areasalongstreams,lakesandponds(Wa);andhabitats withasuitablerockyorstonysubstratum(R)(Table1).

3.2. Climatereconstruction

PaleoclimaticdatafromRiparoMochihavebeen calcu-latedusingthebioclimaticmodeldescribedbyHernández Fernández(2001a,2001b).First,themammalassemblage wasassignedtotheclimatictypesdescribedinHernández Fernández(2001b) and in Walter (1970), following the values establishedin Hernández Fernándezand Peláez-Campomanes (2005): IV–subtropical with winter rains and summer droughts; VI–typical temperate;

VII–arid-Table1

SmallmammalpercentagesofMinimumNumberofIndividuals(MNI)andsmallmammaldistributionbyhabitat. Tableau1

Pourcentagesdenombreminimumd’individus(NMI)depetitsmammifèresetdistributiondespetitsmammifèresparhabitat.

A C DI DII DIII DIV E F G H I OD OH OW W R Wa Arvicolaamphibius 16.1 22.7 26.9 10.2 8.7 8.6 4.3 10.7 13 31.4 8.9 1 Chionomysnivalis 7.6 5.8 6.8 4.6 4.3 2.7 4.3 5.7 8.9 1 Microtusarvalis 19.4 37.9 30.8 48.9 60 56.2 60.9 32 43.5 28.6 15.6 1 Microtusagrestis 12.9 5.7 3.1 4.3 2.2 0.5 0.5 Microtuscf.agrestis 1.9 1.1 0.5 0.5 Microtus(Terricola)gr. multiplex-subterraneus 6.5 6.1 3.8 6.8 8.2 10.5 8.7 9.3 4.3 11.4 20 0.5 0.5 Clethrionomysglareolus 0.6 4.3 2.9 15.6 0.25 0.75 Apodemussp. 16.1 6.1 5.8 4.5 1.5 3.1 13.0 18.7 8.7 5.7 11.1 1 Apodemussylvaticus 1.5 1.9 0.5 2.7 4.4 1 Apodemusﬂavicollis 3.2 1.5 2.3 0.5 1.2 2.7 8.7 2.9 1 Rattussp. 1.9 - -Eliomysquercinus 3.2 1.5 0.6 1.3 2.2 0.75 0.25 Talpasp. 1.9 1.5 0.6 0.5 0.5 Talpaeuropaea 22.6 12.1 15.4 11.4 8.7 6.2 8.7 16 17.4 11.4 6.7 0.5 0.5 Erinaceuseuropaeus 3.8 2.3 2.5 4 2.2 0.25 0.75 Sorexgr.araneus 2.1 1.2 2.2 0.75 0.25 Sorexsp. 0.5 0.75 0.25 Neomyssp. 3.0 0.75 0.5 TotalMNI 31 66 52 88 195 162 23 75 23 35 45 TotalNISP 52 114 79 160 306 244 40 105 28 50 66

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Fig.2. SomeidentiﬁedsmallmammalsfromRiparoMochi,allscalesare1mmlong.1and2:Arvicolaamphibius,leftandrightm1;3and4:Chionomysnivalis, rightm1;5,6,and7:Microtusarvalis,onerightm1andtwoleftm1;8:Microtus(Terricola)gr.multiplex-subterraneus,rightm1;9and10:Clethrionomys glareolus,leftandrightm1;11:Apodemussylvaticus,leftm1;12:Apodemusﬂavicollis,rightm1;13:Eliomysquercinus,leftm1,m2,andm3;14:Sorexgr. araneus,leftmandible;15:Neomyssp.,mandibularcondyle;16:Erinaceuseuropaeus,rightm1.

Fig.2. ExemplesdepetitsmammifèresidentiﬁésauRiparoMochi;toutesleséchellessontde1mmdelong.1et2:Arvicolaamphibius,m1gaucheet droite;3et4:Chionomysnivalis,m1droite;5,6et7:Microtusarvalis,unem1droiteetdeuxm1gauches;8:Microtus(Terricola)gr.multiplex-subterraneus, m1droite;9et10:Clethrionomysglareolus,m1gaucheetdroite;11:Apodemussylvaticus,m1gauche;12:Apodemusﬂavicollis,m1droite;13:Eliomys quercinus,m1,m2,m3gauches;14:Sorexgr.araneus,mandibulegauche;15:Neomyssp.,condylemandibulaire;16:Erinaceuseuropaeus,m1droite.

temperate; VIII–cold-temperate (boreal); IX–polar. The faunalassemblagewasanalysedusingtheClimatic Restric-tionIndex(CRIi=1/n,wherenisthenumberofclimatic

zonesinhabitedbythespeciesandiistheclimaticzone wherethespeciesappears).

Afterwards,theBioclimaticComponent(BC)was calcu-latedusingthefollowingformula:BCi=(CRIi)×100/S,

where S is the number of species by unit at Riparo Mochi.FromtheBC,climaticparameterswereestimated withmultiplelinearregressionmethodusingthevalues givenbyHernándezFernándezandPeláez-Campomanes (2005). Mean Annual Temperatures (MAT), Mean Tem-peratures of the Coldest and Warmest month (MTC and MTW respectively) and Mean Annual Precipitation (MAP)wereobtained.Thedatawerecomparedwiththe present conditions registered at Sanremo meteorologi-cal station(Imperia, Liguria, 43◦47’N, 7◦46’E, 9ma.s.l.), approximately 20km east from the site. The current data are: MAT=16.4◦C, MTW=23.5◦C, MTC=9.5◦C and MAP=773mm.

4. Resultsanddiscussion

4.1. TheRiparoMochisequence

ThesmallmammalassemblageofRiparoMochiis char-acterizedbyhighpercentageofopenenvironment-related arvicolidssuchasMicrotusarvalisandMicrotus(Terricola) gr.multiplex-subterraneus.Speciesthatliveinforestorin

forestmargins(Apodemusgr.sylvaticus-ﬂavicollis, Clethri-onomysglareolus)reachrelativehighpercentagesonlyin UnitI,F,andA.

Arvicolaamphibius,aspeciesthatintheItalian penin-sulalivesnearwaterstreams,lakesandponds,ispresent throughoutthesequence.AllArvicolaspecimenshave posi-tiveorMicrotus-typeenamel(Fig.2,number1and2).Thus, thepresenceofArvicolasapidus,aspeciescharacterizedby anegativeorMimomys-typeenamel,signalledbyAbbassi (1999)atRiparoMochiisnotconﬁrmed.

The BCi values (Table 2), related only to

pres-ence/absenceofspecies,aredominatedbycomponentVI because most the rodents found at Riparo Mochi cur-rentlyliveintemperateclimateareas(Mitchell-Jonesetal., 1999).Nevertheless,theHabitatWeightingvariations, con-nectedtotheabundantorpoorpresenceofinsectivores androdentsintheassemblage,indicatethatseveral envi-ronmentchangeshappenedduringtheformationofthe sequence.

AstheUnitsfromItoE(Table1)arecharacterizedbya lowMNI,conclusionsonenvironmentandclimatechanges duringthetransitionfromtheMousteriantothe Aurigna-cianmustbeconsideredwithcaution.

In Unit I (NMI=45), the biodiversity is high and equallydistributed(Simpsonindex=0.865;Table2)and the presence of forest-related species is relatively high (OW=22.68%;W=30.56). Theprecipitation valueis the lowest of the entire sequence (MAP=872mm), while temperature proxies indicate the presence of very cold

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Table2

Biodiversity,ClimateandLandscapevalues.Percentagerepresentationofsmallmammaltaxaassociatedwithopendrymeadows(OD),openhumid meadows(OH),openwoodlandenvironments(OW),woodlandenvironments(W),rockyenvironments(R)andlandscapesconstitutedbyriver,lakesand ponds(Wa);bioclimaticcomponents(BCi);temperatureandprecipitationreconstructionforRiparoMochisequence:MAT:MeanAnnualTemperature,

MTW:MeanTemperatureoftheWarmestmonth,MTC:MeanTemperatureoftheColdestmonth,MAP:MeanAnnualPrecipitation.Numberoftaxaineach layer;valuesobtainedforevenness:Simpsondiversityindex=1–

(pi2).

Tableau2

Valeursdebiodiversité,climatetpaysage.Représentationdespourcentagesdestaxonsdepetitsmammifèresassociésàuneprairiesècheouverte(OD), uneprairiehumideouverte(OH),unenvironnementboiséouvert(OW),unenvironnementboisé(W),unenvironnementrocheux(R)etdespaysages constituésderivières,lacsetétangs(Wa);composantsbioclimatiques(BCi);reconstructiondestempératuresetdesprécipitationsdelaséquencedu

RiparoMochi:MAT,moyennedestempératuresannuelles,MTW,moyennedestempératuresdumoislepluschaud,MTC,moyennedestempératuresdu moisleplusfroid,MAP,moyennedesprécipitationsannuelles.Nombredetaxonspourchaqueniveau;valeursobtenuespourlarégularitédesespèces: indicedediversitédeSimpson=1–

(pi2).

%HabitatWeighting BCi Climaterecords Biodiversity

OD OH OW W R Wa IV VI VII VIII IX MAT MTW MTC MAP Ntaxa 1-D

A 19.35 20.97 20.97 21.77 0.81 16.13 12.50 54.17 4.17 10.71 0.00 13.2 19.0 8.4 1822 7 0.826 C 37.88 11.36 12.12 10.23 7.95 24.24 25.00 75.00 4.17 7.14 3.13 9.7 16.6 4.3 1345 8 0.771 DI 30.77 10.58 15.38 10.58 5.77 26.92 16.67 58.33 4.17 14.29 3.13 9.7 17.4 3.1 1465 8 0.780 DII 48.86 12.50 12.50 8.52 6.82 10.23 8.33 66.67 4.17 14.29 3.13 9.2 16.2 3.3 1695 8 0.714 DIII 59.69 11.48 9.95 2.55 4.59 8.67 16.67 75.00 4.17 14.29 3.13 7.4 15.5 0.7 1381 8 0.598 DIV 56.17 11.42 10.65 7.10 4.48 8.64 16.67 83.33 4.17 21.43 3.13 3.4 13.7 -5.5 1102 11 0.651 E 60.87 8.70 18.48 16.30 4.35 4.17 45.83 4.17 10.71 15.0 19.7 11.0 2136 6 0.594 F 32.00 12.67 16.33 28.00 3.00 10.67 25.00 75.00 4.17 7.14 3.13 9.7 16.6 4.3 1345 8 0.792 G 43.48 10.87 19.57 17.39 4.35 13.04 8.33 58.33 4.17 7.14 3.13 13.2 18.0 9.6 1974 6 0.730 H 28.57 11.43 17.86 10.71 5.71 31.43 8.33 66.67 4.17 14.29 3.13 9.2 16.2 3.3 1695 7 0.782 I 15.56 16.11 22.78 30.56 9.44 8.89 25.00 75.00 4.17 21.43 3.13 3.9 14.9 -5.8 872 11 0.865

conditions (MAT=3.9◦C).Such climatedataare directly relatedtothehighpercentageofChionomysnivalis(8.9%), theabsenceofforestspeciessuchasGlisglis,thelow per-centage ofErinaceuseuropaeus,andthehighpercentage of Microtusarvalisand Microtus (Terricola)gr. multiplex-subterraneus.Thus,itispossibletosuggestthepresenceof anenvironmentcharacterizedbytreepatchesinanopen androckyarearatherthanadenseforestedlandscape.

The cold climate evolves to slightly warmer condi-tionsthroughthesequence(UnitsHandG)reﬂectedina moreopenenvironment.Theriseintemperatureandopen environmentmayberelatedtoadifferentprecipitation dis-tributionthatcausedthelossofbiodiversityandtheforest environmentreduction(Fig.3).

UnitF(earlyAurignacian)isthemostreliablelayerof this portionof thesequence(NMI=75): itis dominated byMicrotusarvalis(32%),butthegenusApodemusiswell represented (Apodemussp.and Apodemus gr. sylvaticus-ﬂavicollis>20%).Theenvironmentwasprobablymadeup ofsparsewoodenareaswithoutbushandopenmeadows. Thisreconstructioncanbeimplementedthankstothe paly-nologicalanalysescarried outattheRiparo Bombrini,a rockshelterlocatedfewtensofmeterseastwardtoRiparo Mochi,whichyieldedlayersrelatedtoMiddle-Upper Palae-olithictransition.Theauthorssuggestedthepresenceof spacepinewood,mixedoakwood,andthermophiloustaxa relatedtothecoastalareaintheﬁnalMousterianlayers; warmertemperatureswithareducedtreecoveringwere recordedduringthedepositionoftheearlyUpper Palae-olithiclayers(ArobbaandCaramiello,2009).

Thelossofequalityinthesmallmammalassemblage distributioncontinuesinUnitEofRiparoMochi.Here,few smallmammalshavebeenrecovered(NMI=23)toreach thelowest values in UnitD. PhasesDIV, III, and IIare dominatedbyonespecies,Microtusarvalis;the biodiver-sityis relatively low when compared tothelowermost

phase(1-Dvalues from0.598to0.714).Theforest com-ponentsdecreaseinfavouroftheoneindicativeofmore openenvironmentsduetoacoldanddryclimate. From phaseIVtoII,aslightclimateimprovementisregistered; itspeakisreachedduringarelativetemperatecyclewhich characterizesthephase I(topof UnitD)aswellas the UnitC,evenifthegeneralcontextisdominatedbycold climateconditions.Intheseunits,theariseofawater com-ponentrepresentedmainlybyArvicolaamphibiussuggests thepresenceofaﬂuvialorawater-relatedenvironment, probablyduetothepresenceoftheSanLuigiriver,about 450mtothewestofthesite.

After the thin and sterile Unit B, the Riparo Mochi sequence ends withtheculturalUnit A (Epigravettian). ThisUnitalsohasalowNMIbutthehighpercentageof forest-related indicatorsand therelative warm temper-aturessuggest that the unit formedin an environment similartothepresentone.

4.2. CorrelationbetweenRiparoMochisequenceand otherclimateproxies

Therelativelyhighpercentageofforestrelatedspecies inUnitsItoFsuggeststhat,intheperiodoftimebetween DO(=GI)12andH4(followingDoukaetal.,2012),the west-ernLiguriaenvironmentwasconditionedbythe Mediter-raneanSeainfluence.Thisisalsoconfirmedbypollendata indicatingthepresenceofMediterraneanplants(Oleaceae) throughtheRiparoMochisequence(Renault-Miskovsky, 1972).Incontrast,thepercentagedecreaseofMicrotus (Ter-ricola)multiplex-subterraneusandClethrionomysglareolus suggestsanenvironmentmodificationfromaforestwith shrubsandawell-formedsubstratum(UnitI,Mousterian) toanopenforest(UnitF,earlyAurignacian).Thechangeis probablyduetoaclimatedeteriorationrelatedtoastadial episodeofMIS3.Unfortunately,itisnotpossibletoargue

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Fig.3.RepresentationofthesmallmammalSimpsondiversityindex(1-D),themeanannualprecipitation(MAP),themeantemperatureofthecoldestmonth (MTC),themeanannualtemperature(MAT),themeantemperatureofthewarmestmonth(MTW)andthelandscapepercentages(OD:openmeadows; OH:openhumidmeadows;W:woodlands;R:rockyareas;Wa:areasalongstreams,lakesandponds).GI:GreenlandInterstadial;GS:GreenlandStadial. Fig.3. Représentationdel’indicedediversitédeSimpsondespetitsmammifères(1-D),moyennedesprécipitationsannuelles(MAP),moyennedes températuresdumoisleplusfroid(MTC),moyenneannuelledestempératures(MAT),moyennedestempératuresdumoislepluschaud(MTW)et pourcentagesdepaysage(OD:prairiesouvertes;OH:prairieshumidesouvertes;W:bois;R:zonesrocheuses;Wa:zoneslelongderuisseaux,lacset étangs).GI:Interstadegroenlandais.GS:Stadegroenlandais.

whetherthischangeisrelatedonlytoH4ratherorany otherstadials(GS12,11,10,and9).

RiparoMochiUnitsfromItoFcanbecomparedto lay-ersSUA5+A6toA1oftheFumaneCave(Verona,Italy). There,adecreaseinpercentageofwoodlandcoverageis registeredandthisreductionculminatesinlayersA3-A1 thatareattributedtoHeinrichEvent4(López-Garcíaetal., 2015).Differencesbetween thetwo sequencesmust be noticed:attheRiparoMochi,thebiodiversityislowerthan intheFumanecaveandboreal/continentalclimate indica-tors,aswellaseasternEuropespeciessuchasDinaromys bogdanovi,Microtusoeconomus,Sicistabetulina,and Crice-tuscricetusareabsent.ThecompositionoftheRiparoMochi assemblageisbetterrelatedtosouthwesternItalian penin-sula coastal sites, at Grottadella Cala and Grotta della Serratura (Bambini, 1996; Bertoliniet al., 1996), where suchcoldclimaterelatedspeciesareabsent,ratherthan tonorthernItalianPeninsulasites(Berto,2013;Bonand Boscato, 1996; López-García et al., 2014a; Masini and Abbazzi,1997).

Threeshort and not yet chronologically determined interstadialscharacterizedbysparsetreecover are also foundintwoFrenchsitepollenrecords:LesÉchets,zoneG (BeaulieuandReille,2008)andBouchet(Reilleetal.,2000). Finally,theRiparoMochiculturalUnitsItoFcanberelated tothepalaeoenvironmentalreconstructionssuggestedfor FimonLake(FP17a,17b,and17c)andforAzzanoDecimo (AZ65-67),inthesouthernAlps.ThesouthernAlpspollen recordsshowathicktreecoverageand,evenduringstadial periodsbecausetheenvironmentwasdominatedbyPinus

andBetulaprobablyduetoorographicprecipitations(Pini etal.,2010b,2009).Availabledates(Doukaetal.,2012) allowustorelatethebaseofUnitDofRiparoMochi (Zep-pieri’sphaseIV)toHeinrichEvent3.Theeffectsofthecold andaridclimateareclearlyvisibleonthesmallmammal assemblage–especiallywhenphasesIVandIIIofUnitDare considered–andtheycanberelatedtoLesÉchets(zone O),FimonLake(FP18),andAzzanoDecimo(AZ68).

TheUnitD(phaseIII)temperatureincreasefollowedby anopenenvironmentdecreasevisibleatthetopofUnit D ofRiparo Mochican berelatedtoepisodes following the H3. No abrupt changes are visible in thetransition fromthetop ofUnit DtoUnitC, althoughthelatter is attributedtotheﬁnalGravettiantechnocomplex, chrono-logicallyrelatedtoLastGlacialMaximum(LGM)andDO2 (Berto,2013;Doukaetal.,2012;PalmadiCesnola,1993). Noradiometricdatesareavailableyet,buttwo hypothe-sescanbeproposedassumingthattheopenenvironment decreaseinUnitCisrelatedtotheDO2:A)theupperpartof UnitDcanberelatedtoDO4-3,twocycleschronologically closeandhardlydistinguishablewithoutradiometric dat-ingsinacontinentalsequence,ahiatusinsedimentationis alsopresentbetweenLGMandDO2;B)thedepositionof UnitDtookplaceinperiodbetweenH3andLGM.

ItaliansmallmammalsequencesrelatedtoH3and fol-lowingDOcyclesareveryscarce;theclimaticdeterioration related tothis eventin northern Italy isvisible only at GrottadelBroion(layersG2–F),whereadecreaseofforest relatedspeciesisregistered(Colamussi,2002).In south-ernItaly,theforestindicatorsdecreaseatGrottadellaCala

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(3m-1)(Bambini,1996)andtheincreaseinpercentageof MicrotusarvalisatGrottaPaglicci(22bto21c)canberelated toH3(Bertoetal.,2017a).

ThelateEpigravettianUnitAofRiparoMochiis char-acterizedbythehighpercentageofforestrelatedspecies. The Unit probably accumulated after theturnover reg-istered during the Late Glacial in the small mammal communities.InsouthwesternFrance,cold-climatespecies suchas Microtus oeconomus,Microtus gregalis and Sper-mophilussp.werereplacedbyforest-relatedspecies(Royer et al.,2016)whilein thesouthern Alps,species related tocold-continentalclimatessuchasDinaromysbogdanovi, MicrotusoeconomusandCricetuscricetusdisappeared(i.e. atRiparoTagliente,Bertoetal.,2017b).Finally,insouthern Italythesmallmammalcommunities,dominatedduring the MIS 3 and 2 by Microtus arvalis, changed to forest relatedassemblages(dominatedbyGlisglis)ortoastrictly Mediterranean-relatedcommunity dominated by Micro-tus(Terricola)savii(Bertoetal.,2017a;López-Garcíaetal., 2014a).ThechangesarerelatedtoBølling–Allerød Intersta-dialafforestationregisteredalsoinpollensequences(Berto etal.,2017b;Ravazzietal.,2007).

5. Conclusions

ThesmallmammalanalysisfromRiparoMochiallows ustoreconstructtheenvironmentanditschangesinthe regionsurroundingthesite fromtheﬁnal Middle Palae-olithictotheUpperPalaeolithic.Thechangesaredueto D/OcyclesandHeinrichEventsregisteredduringMIS3and 2anddetectedinnumerousclimateproxiesofEuropean continentalsequences.

Duetoitsparticularposition,thesmallmammal com-munity of Riparo Mochi has been inﬂuenced by the Mediterraneanclimateand,evenduringthecoldeststadial periodofMIS3,speciesrelatedtoborealregionsusually documentedinnorthernItalydidnotreachthisarea.

DuringtheMiddletoUpperPalaeolithictransition,two cold and one slightly warm oscillations were detected depicting a forest and bush reduction near theshelter. It wasprobablyduetostadialmomentsnothighlighted throughoutthelowsmallmammalsMNIregisteredinthe collection.

DuringtheGravettian,theenvironmentchangedfrom onedominatedmainlybyopengrasslands,relatedtoacold anddryclimate,toonecharacterizedbyopenforestswith anincreaseofwaterstreams,thelatterprobablyduetothe ﬂowrateofthenearSanLuigiriver.Theradiometricdate atthebaseofUnitDandthepresenceoflithictoolsrelated toﬁnalGravettianinUnitCindicatethatthisportionofthe sequenceformedbetweenH3andD/O(GI)2.

ThesmallmammalassemblageoftheuppermostUnit A,relatedtoﬁnalEpigravettian,indicatesthepresenceofa forestedenvironment.ThisUnitprobablyformedafterthe faunalturnoverregisteredinsouthernEuropeduringthe LateGlacial.

ThesequenceofsmallmammalassemblagesatRiparo Mochi ﬁlls a gapbetween southern Italy and southern Francesmallmammalsequences,conﬁrmingthesequence as a reference one not only for the Middle to Upper

PalaeolithictransitionbutalsoforthetransitionfromMIS 3andMIS2.

Acknowledgements

TheDipartimentodiLettereeFilosofia,Universitàdegli studidiTrento(Italia)partiallysupportedthisstudy.The formerSoprintendenzaperiBeniArcheologicidella Lig-uriagrantedthepermissionforthestudy;inparticular,we thankDr.AngioloDelLuccheseandDr.ElisabettaStarnini. Wearegratefultomanypeopleandfriendswhohelped usduringthefieldwork,especiallytoM.A.Segréandthe staffof the MuseoPreistorico dei BalziRossi. We want tothanktheeditorialsecretaryFathiaLemhemdiandthe reviewersJuanManuelLópez-GarcíaandLarsvandenHoek Ostendefortheircommentsthatimprovedthefinal ver-sionofthemanuscript.TheauthorsalsothankBenedetto SalaandElisa LuzifortheirusefulsuggestionsandJulie ArnaudfortheFrenchspellingcheck.Finally,weareindebt withA.Segrewhogenerouslygaveusdirectinformation andoriginalfielddocumentationabouttheRiparoMochi excavation.

AppendixA. Supplementarydata

Supplementarydataassociatedwiththisarticlecanbe found, in the online version, at https://doi.org/10.1016/ j.crpv.2018.04.007.

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