An overview of the crocodylian fossil record from Sardinia (Italy)

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Original

article

An

overview

of

the

crocodylian

fossil

record

from

Sardinia

(Italy)

Un

aperc¸

u

sur

les

crocodiles

fossiles

de

Sardaigne

(Italie)

Daniel

Zoboli

a,∗

_,

_Luigi

_Sanciu

b,c

_,

_Gian

_Luigi

_Pillola

a

_,

_Massimo

_Delﬁno

d,e

a_Dipartimento_di_Scienze_Chimiche_e_Geologiche,_Università_di_Cagliari,_51,_via_Trentino,₀₉₁₂₇_Cagliari,_Italy

b_Centro_Studi_di_Storia_Naturale_del_{Mediterraneo,}_Museo_di_Storia_Naturale_Aquilegia_–_GeoMuseo_Monte_Arci_“Stefano_Incani”,_Via_San_Francesco,₀₉₀₉₀ Masullas,Italy

c_P.AR.C.,_loc._S._Antine,₀₈₀₃₀_Genoni,_Italy

d_Dipartimento_di_Scienze_della_Terra,_Università_di_Torino,_35,_via_Valperga_Caluso,₁₀₁₂₅_Torino,_Italy

e_Institut_Català_de_{Paleontologia}_Miquel_Crusafont,_Universitat_Autònoma_de_Barcelona,_Ediﬁci_ICTA-ICP,_Carrer_de_les_Columnes_s/n,_Campus_de_la_UAB, 08193CerdanyoladelVallès,Barcelona,Spain

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Articlehistory:

Received21November2018 Accepted29April2019 Availableonline29May2019 Keywords: Crocodylidae Tomistominae Tomistomacalaritanum SystematicPalaeontology Cenozoic Italy

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InthepresentcontributionwefocusonanoverviewoftheCenozoiccrocodyliansofSardinia(Italy). CrocodyliansfromthisItalianislandhavebeenpublishedsincethesecondhalfofthe19thCenturyand anumberofpapersreportedonthepresenceofthesevertebratesin16differentlocalities.Theremains, someofwhichapparentlylost,aremainlyrepresentedbyisolatedteethhousedinhistoricalcollectionsof differentmuseums(inItalyandTheNetherlands)thatinmostcaseshaveneverbeenfiguredordescribed. Weillustrateforthefirsttimeandprovidebriefdescriptionsofallcrocodylianteethcollectedintheisland aswellasofthestillavailableholotypeofTomistomacalaritanum,theonlycrocodyliantaxonestablished onmaterialfromSardinia.Inaddition,wereportcrocodylianremainsfromtwonewlocalities,oneof whichtestifiesthepresenceofthesevertebratesalreadyintheEocenerecordoftheisland.

Motsclés: Crocodylidae Tomistominae Tomistomacalaritanum PaléontologieSystématique Cénozoïque Italie

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Danslaprésentecontribution,nousnousconcentronssurunaperc¸udescrocodylienscénozoïquesde Sardaigne(Italie).Lescrocodiliensdecetteîleitalienneontétépubliésdepuislasecondemoitiédu 19e_siècle_et_un_certain_nombre_d’articles_ont_fait_état_de_la_présence_de_ces_vertébrés_dans₁₆_localités

différentes.Lesrestes,dontunepartiesembleavoirétéperdue,sontprincipalementreprésentéspardes dentsisolées,conservésdansdescollectionshistoriquesdedifférentsmusées(enItalieetauxPays-Bas) qui,danslaplupartdescas,n’ontjamaisétéﬁgurésnidécrits.Nousillustronspourlapremièrefois etfournissonsunebrèvedescriptiondetouteslesdentsdecrocodilienscollectéessurl’îleainsiquede l’holotypeencoredisponibledeTomistomacalaritanum,leseultaxondecrocodilienétablisurdumatériel deSardaigne.Enoutre,noussignalonsdesrestesdecrocodiliensdedeuxnouvelleslocalités,dontl’une témoignedelaprésencedecesvertébrésdéjàdanslesdépotsdel’Éocènedel’île.

1. Introduction

1.1. Palaeogeographiccontext

TheSardinia-CorsicaMassifrepresentsamicroplatelocatedin thewesternMediterraneanbasin,betweentheLiguro-Provenc¸al

∗ Correspondingauthor.

E-mailaddress:zoboli.a@tiscali.it(D.Zoboli).

basinandtheTyrrenianSea.ItrepresentsasegmentoftheAlpine orogenicsystemoriginallylocatedincontinuitywiththesouthern Europeancontinentalmargin.FromtheearlyMioceneonwards,the Sardinia-CorsicaMassifstartedtoseparatefromthemainlandwith anticlockwise rotation (Vardabasso, 1962; Cherchi,1974).Thus, the Sardinian Neogene fossils representthe oldestknown case ofendemicinsularvertebratefaunasfromtheMediterraneanSea (Mennecartetal.,2017).However,thepalaeogeographicevolution ofthisareaalloweddifferentfewimmigrationphasesofmainland vertebratesduringtheNeogene(AzzaroliandGuazzone,1979;Van

https://doi.org/10.1016/j.annpal.2019.05.001

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124 D.Zobolietal./AnnalesdePaléontologie105(2019)123–137

derMade,1999;Abbazzietal.,2004,2008a,2008b)andespecially duringtheQuaternary(Azzaroli,1983;Palombo,1985,2009;Van derMade,1999;PillolaandZoboli,2017;ZoboliandCaddeo,2016; ZoboliandPillola,2016a,2017a;Zobolietal.,2016,2018). 1.2. CenozoicSandinianPaleoherpetology

TheSardinianfossilrecordofCenozoicherpetofaunasisrather interesting,thoughnotabundantandcertainlyincompletebecause manyextantspecieshavenotbeentracedyet.However,members ofeachofthehighertaxacurrentlyinhabitingthenorthern hemi-spherehavebeenidentified:Squamata(Serpentes,Amphisbaenia, “Lacertilia”),Testudines,andCrocodylia.Squamateswerereported fromtheearlyMiocene(Oschiri;VenczelandSanchiz,2006)tothe Holocene(Delfino,2002).Amongsnakes,worthmentioningisthe “exotic”presenceofanextinctaniilidsnake,Eoanilius oligoceni-cus,atOschiri(VenczelandSanchiz,2006)andtheholotypeand onlyspecimenofthebooidsnakePalaeopythonsardusPortis,1901 (Portis,1901a),thatwasrecentlyidentifiedasanundeterminedfish (Delfinoetal.,2014).Booidsnakeswereinanycasepresentin Sar-diniaduringthelatePlioceneastestifiedbyEryxfromCapoMannu (D1LocalFauna;Delfinoetal.,2011).ThePleistocenekarstfissures ofMonteTuttavista(Abbazzietal.,2004)yieldedthousandsof dis-articulatedsnakeremains(mostlyisolatedvertebrae)belongingto aviperid,astillundescribednewcolubridspecies,andagrasssnake ofthegenusNatrix.TheviperofMonteTuttavistaalongthoseoftwo othersites(VenczelandSanchiz,2006;Delfinoetal.,2011) repre-sentstheonlySardinianevidenceforthisgroupofsnakescurrently extinctontheisland(Sindacoetal.,2006).Colubridsnakes (colu-brinesandNatrix)werealsoidentifiedatCapoMannu(D1Local Fauna;Delfinoetal.,2011)andinsomeQuaternarysites(Delfino, 2002).

LizardsarerepresentedatMonteTuttavistabythemost abun-dantandinformativeEuropeanremainsofagamids(Delfinoetal., 2008), bythefirstevidence inSardiniaof alarge-sizedlacertid lizardthatrecentlyturnedouttobelongtoTimon(Tschoppetal., 2018),aswellasbysmallerlacertidlizardsandgeckos(Abbazzi etal.,2004).AnguidlizardsreferredtothegenusDopasiahavebeen reportedfromtheearlyMioceneofOschiri(VenczelandSanchiz, 2006)andthePlioceneofCapoMannu(D1LocalFauna;Delfino etal.,2011);thelatterwasrecentlyidentifiedasOphisaurusspinari byKlembaraandRummel(2018).Skinkshavebeenalsocitedor describedfromtheMioceneofOschiri(VenczelandSanchiz,2006) totheHolocene(Delfino,2002),buttheirrecordisveryscarce.

InthePlioceneofCapoMannu(D1LocalFauna;Delﬁnoetal., 2011)andPleistoceneofMonteTuttavista(Abbazzietal.,2004) amphisbaeniansarealsopresentandlikelybelongtotheBlanidae. They were already present in the early Miocene of the island (Oschiri)withBlanusgracilis Roˇcek 1984(Venczeland Sanchiz, 2006).

FossilturtlesarerathercommoninSardiniaand,witha sin-gle exception, are all represented by Neogene and Quaternary cryptodires.Oneexceptionconsistsof astillunderscribed pleu-rodirefromtheEoceneofCixerrithatwaspreliminaryreferredto Erymnochelys(DelﬁnoandRook,2008a).Amongthecryptodires, softshellturtlesarepresentinafewEoceneandMiocene locali-ties.ArevisionofProcyclanorbissardus(Portis,1901b)revealedthat thenameisanomendubiumandthetypeisactuallyan indeter-minatepan-trionychine(seeGeorgalisetal.,2017,andreferences therein).Testudinidsarerepresentedbyanextinctspecies, Tes-tudopecoriniiDelﬁno,2008(inAbbazzietal.,2008a),fromthelate PlioceneofCapoMannuD4localfauna(SanVeroMilis)andby Testudocf.T.hermannifromthePleistoceneofMonteTuttavista (Abbazzi et al., 2004).Zoboli(2017) reported a large-size pha-lanxofa probabletestudinidfromtheMajor’s Collectionofthe NaturhistorischesMuseumBasel.Thefossilwascollectedinthe

bonebrecciasofMonteSanGiovanni(southernSardinia)andcould testifyforthepresenceofagianttortoiseinthe?LatePleistocene ofSardinia.

Freshwater turtles have been described in a few localities. FragmentaryremainsrepresentingthelastItalianevidenceof Mau-remyscomefromtheLatePleistoceneofSanGiovannidiSinis(Chesi etal.,2007),whereasa well-preservedcarapaceofEmys orbicu-lariscomesfromGrottadiMonteMeana,nearSantadi(Chesietal., 2008).Marineturtles,referredtoCheloniidaeindet.,arepresent intheearlyMioceneofNoragugume(Chesi,2008)andinthelate MioceneofCagliari(ZoboliandPillola,2016b).Trachyaspislardyi wasidentiﬁedintheMioceneofBosa(Chesi,2008)andCaretta, wasreportedonlyinHolocenearchaeologicalsites(Wilkens,2003; Delussu,2007).

Theﬁrst fossilcrocodylians ofSardinia havebeen published sincethesecondhalfofthe19thCentury(Gennari,1868;Capellini, 1890a,1890b)andsincethenanumberofpapersreportedonthe presenceordescribedremainsinlocalitiesofdifferentsubregions ofthe Island(Fig.1).The remains,all Miocenein age,are rep-resentedinmostofthecasesbyisolatedteeth(seebelow),but alsoanearlycompleteskull andlowerjaw(andassociatedfew postcranialelementssuchasosteodermsand vertebrae) onthe basisof whicha newspecies oftomistomine,Tomistoma calari-tanumCapellini,1890,wasdescribed(asTomistomacalaritanus,see below;Capellini,1890b).Duetothenumberofpaperspublishedon thesubject(ofteninoldItalianlocaljournals,seereferencesbelow), thedamageandlosssufferedbythetypematerialofT.calaritanum anditsconvolutedtaxonomicandnomenclaturalhistory,itishere presentedanoverviewonthecrocodylianfossilrecordofSardinia, withareassessmentoftheidentiﬁcationspublishedinthepast.

2. ThecrocodilianfossilrecordofSardinia

TheSardiniancrocodylianfossilshereinillustratedarehousedin historicalcollectionsofdifferentmuseums.Unpublishedmaterials fromtwonewlocalities,oneofwhichpushingbacktotheEocene thepresenceofcrocodyliansintheisland,arealsodescribed.

InstitutionalAbbreviations:MDLCA,MuseoSardodiGeologia e Paleontologia Domenico Lovisato, Cagliari, Italy; PARC, Paleo ArcheoCentro,Genoni,Italy;MACPL,GeoMuseoMonteArci “Ste-fano Incani”, Masullas, Italy; MGPT-PU, Museo di Geologia e Paleontologiadell’UniversitàdegliStudidiTorino,Turin,Italy;UU, UniversityofUtrecht,Utrecht,TheNetherlands.

2.1. Ypresian 2.1.1. Locality

Escalaplano(Sarrabus-Gerreisubregion,southernSardinia).

Ageandgeologicalsetting:MonteCardiga Formation(early

Eocene, Ypresian; Cherchi, 1985). The Monte Cardiga Fm. is a lithostratigraphicunitdiscontinuouslycroppingoutinthe south-easternSardinia(SaltodiQuirra,Sarrabus-Gerrei,Ogliastra).Inthe Escalaplano and Perdasdefogu areasthis formation overliesthe Palaeozoic basementand theMesozoiclithologies (Vardabasso, 1962).Atthebottom,theMonteCardigaFm.isrepresentedby poly-genicconglomerates,sandstoneswithsiliceouscementandmarly sandstoneswithscarceplantremains. Intheupperpart,marls, calcarenites and bioclastic limestones with a rich foraminiferal faunaarepredominant;locally,layersofpolygenicconglomerates arepresent.Furthermore,severallevelsofcalcareniterichin gas-tropods(cerithids)andothermarineinvertebratesarepresentin themedium-basalportionofthesequence.Thecrocodiletoothwas collectedinoneoftheselevelscroppingoutatthesouthern periph-eryofthevillageofEscalaplano.TheMonteCardigaFm.isreferred

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Fig.1.SardinianCenozoiccrocodylian-bearinglocalities:1,Escalaplano;2,Cava Duidduru(Genoni);3,StrintudeMelonisandComidiano(Nurri);4,Laerru;5, Oschiri;6,LaCrucca(Sassari);7,MonteOrria(Nulvi);8,Tresnuraghes;9,Bingia Farg-eri(Cagliari);10,Florinas;11,FiumeSanto(PortoTorres);12,CapoFrasca(Arbus); 13,Tuvumannu;14,BonariaandMontixeddudiBonaria–CavaTimon;15,Cimitero MonumentalediBonaria;16,IsMirrionis–Piazzad’Armi.Whitesquare: Ypre-sian,triangles:?Chattian-Burdigalian,blacksquares:Langhian-Serravallian,circles: Tortonian-Messinian.

Localitéscénozoïquesayantlivréderestesdescrocodiles:1,Escalaplano;2,Cava Duid-duru(Genoni);3,StrintudeMelonisetComidiano(Nurri);4,Laerru;5,Oschiri;6,La Crucca(Sassari);7,MonteOrria(Nulvi);8,Tresnuraghes;9,BingiaFargeri(Cagliari); 10,Florinas;11,FiumeSanto(PortoTorres);12,CapoFrasca(Arbus);13,Tuvumannu; 14,BonariaetMontixeddudiBonaria–CavaTimon;15,CimiteroMonumentaledi Bonaria;16,IsMirrionis–Piazzad’Armi.Carréblanc:Yprésien,triangles: ?Chattien-Burdigalien,carrésnoirs:Langhien-Serravallien,cercles:Tortonien-Messinien.

totheYpresianonthebasisofthemicropalaeontologicalcontent (Cherchi,1985).

Taxon:Crocodyliaindet.

Material:MACPL1364,oneisolatedtooth(Fig.2A).

Descriptionandremarks:Theisolatedtoothisstillembedded

inthecalcarenitematrixandonlywhatappearstobeamoderately alteredlingualsurfaceisvisible.Theapexisbrokenoffandtheroot ismissing.Thepreservedportionofthecrownis7.7mmlong.The associatedmarinefaunasuggeststhetransportofthecrocodylian remainsfromtheemergedareastotheseaside.

2.2. ?Chattian-Burdigalian 2.2.1. Locality

“Cava Duidduru” (Genoni, Sarcidano subregion, central Sar-dinia).

Ageandgeologicalsetting:NurallaoFormation(?Chattian–

earlyBurdigalian,Funeddaetal.,2011).Theisolatedcrocodylian toothwascollectedinthe“CavaDuidduru”,anabandonedquarry currently reconverted in a geosite. The quarry is opened in the?Chattian–earlyBurdigalianmarinesuccessionoftheNurallao Fm.(Funedda et al.,2011)belongingto theﬁrst Cenozoic sed-imentary cycle of Sardinia.The Nurallao Fm. is representedby marine-littoralsedimentsand isdividedintwo members: Con-glomeratodiDuiddurumember(atthebottom)andArenariedi SerraLongamember(atthetop).Thelowermemberisconstituted by polygenic and heterometric conglomerates mainlyresulting fromtheerosionoftheunderlyingPalaeozoicbasementand sub-ordinatelybyCenozoicvolcanites.Thefossilcontentisrepresented especially byfragments ofbivalve shells.The lithologychanges bothlaterallyandverticallyinthesandstoneandsilicoclasticsands of the Arenarie di Serra Longamember. Thislatest member is representedbysandsandmicroconglomeratesthatlaterallyand verticallychangeincoarsesandstoneswithcarbonatecement. Fur-thermore,decimetric-conglomeratelevelswithelementsofquartz, metamorphitesand granitoids,andtanatocenosisdominatedby bivalvesandechinidsarepresent.IntheDuidduruquarryispresent onlythislastmemberoftheNurallaoFm.Thecrocodyliantooth wasfoundinapoorly-cementedgreysandstonelevelrichin frag-mentsofcarbonizedplants,turritellinegastropodsandcrab-shell fragments. This level seems to testify a proximal depositional environment characterized by ﬂuvial transport and deposition oforganicmaterialinacoastallagoon.Todate,thecrocodylian remains anda mandible ofa smallruminant(Mennecartetal., 2017)aretheonlycontinentalvertebratescollectedinthemarine depositsofthelatestOligocene–earlyMioceneofcentralSardinia.

Material:PARC205626,oneisolatedtooth(Fig.2B).

Descriptionandremarks:Thetoothisrepresentedbyaportion

ofthecrownmissingitsapexand,probably,itsbase.Thepreserved portion,22.3mmlong,isrelativelystout.Mesiodistalcarinaeare presentbutweaklydeveloped(Fig.2B5).

2.2.2. Localities

StrintudeMelonisandComidiano,betweenStrintuArgioluand thetrainstationofNurri(Nurri,Sarcidanosubregion,central Sar-dinia).

Ageandgeologicalsetting:NurallaoFormation(?Chattian–

earlyBurdigalian,Funeddaetal.,2001,seeabove).

Taxon:Crocodyliaindet.(=?Tomistomasp.inLovisato,1892;

Tomistoma calaritanus in Del Vecchio, 1921, Spano, 1985; Crocodyliaindet.inKotsakisetal.,2004;Tomistomacalaritanusin theoriginallabelofMGPT-PU).

Material:MGPT-PU17336,oneisolatedtooth(Fig.2C).

Descriptionandremarks:Twoincompletecrocodileteeth,

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Fig.2.Crocodyliaindet.,isolatedteethfromdifferentlocalitiesofSardinia.A,MACPL1364fromtheYpresianofEscalaplano(southernSardinia).B,PARC205626from the?Chattian–earlyBurdigalianof“CavaDuidduru”,Genoni(centralSardinia);B1,labialview;B2,linguaview;B3-4,lateralview;B5,occlusalview.C,MGPT-PU17336from the?Chattian–earlyBurdigalianofComidiano,betweenStrintuArgioluandthetrainstationofNurri(centralSardinia);C1,labialview;C2,lingualview;C3-4,lateralview. D,MDLCA14114afromtheearly-middleBurdigalianofLaerru(northernSardinia);D1,labialview;D2,lingualview;D3,lateralview;D4,occlusalview.E,MDLCA14114b fromtheearly-middleBurdigalianofLaerru;E1,labialview;E2,lingualview;E3-4,lateralview;E5,occlusalview.F,MDLCA14114cfromtheearly-middleBurdigalianof Laerru;F1,labialview;F2,lingualview;F3-4,lateralview;F5,occlusalview.G,MDLCA14114dfromtheearly-middleBurdigalianofLaerru;G1,labialview;G2,lingual view;G3-4,lateralview;G5,occlusalview.H,UUOS501fromtheBurdigalianofOschiri(northernSardinia);H1,labialview;H2-3,lateralview.I,MGPT-PU17337fromthe upperBurdigalianofMonteOrria,betweenNulviandChiaramonti(northernSardinia);I1,labialview;I2,lingualview;I3,occlusalview.J,MGPT-PU17334fromthe?middle Langhian–lowerSerravallianofBingiaFargeri,Cagliari(southernSardinia);J1,labialview;J2,lingualview;J3–4,lateralview.K–L,fromthe?middleLanghian–lower SerravallianofBingiaFargeri,Cagliari(fromDelVecchio,1921).M–P,fromtheSerravallianofFlorinas(northwesternSardinia)(A.Pigaprivatecollection,fromMarrasand Ventura,1985).ScalebarforA,E,F:0.5cm;scalebarforB,C,I,J:1cm;scalebarforD:0.6cm;scalebarforG:0.3cm;scalebarforH:1.5cm;scalebarforK–P:2cm. Crocodyliaindet.,dentsisoléesprovenantsdedifférenteslocalitésdeSardaigne.A,MACPL1364del’Ypresiend’Escalaplano(suddelaSardaigne).B,PARC205626du?Chattien– Burdigalieninférieurde“CavaDuidduru”,Genoni(Sardaignecentrale);B1,vuelabiale;B2,vuelinguale;B3-4,vuelatérale;B5,vueocclusale.C,MGPT-PU17336du?Chattien –BurdigalieninférieurdeComidiano,entreStrintuArgioluetlagaredeNurri(Sardaignecentrale);C1,vuelabiale;C2,vuelinguale;C3-4,vuelatérale.D,MDLCA14114adu Burdigalieninférieur-moyendeLaerru(norddelaSardaigne);D1,vuelabiale;D2,vuelinguale;D3,vuelatérale;D4,vueocclusale.E,MDLCA14114bduBurdigalieninférieur-moyen deLaerru;E1,vuelabiale;E2,vuelinguale;E3-4,vuelatérale;E5,vueocclusale.F,MDLCA14114cduBurdigalieninférieur-moyendeLaerru;F1,vuelabiale;F2,vuelinguale;F3-4, vuelatérale;F5,vueocclusale.G,MDLCA14114dduBurdigalieninférieur-moyendeLaerru;G1,vuelabiale;G2,vuelinguale;G3-4,vuelatérale;G5,vueocclusale.H,UUOS501 duBurdigaliend’Oschiri(norddelaSardaigne);H1,vuelabiale;H2-3,vuelatérale.I,MGPT-PU17337duBurdigaliensupérieurdeMonteOrria,entreNulvietChiaramonti(nord delaSardaigne);I1,vuelabiale;I2,vuelinguale;I3,vueocclusale.J,MGPT-PU17334du?Langhienmoyen–SerravallieninférieurdeBingiaFargeri,Cagliari(suddelaSardaigne); J1,vuelabiale;J2,vuelinguale;J3–4,vuelatérale.K–L,du?Langhienmoyen–SerravallieninférieurdeBingiaFargeri,Cagliari(d’aprèsDelVecchio,1921).M–P,duSerravalliende Florinas(SardaigneduNord-Ouest)(collectionprivéeA.Piga,deMarrasetVentura,1985).Barred’échellepourA,E,F:0,5cm;barred’échellepourB,C,I,J:1cm;barred’échelle pourD:0,6cm;barred’échellepourG:0,3cm;barred’échellepourH:1,5cm;barred’échellepourK–P:2cm.

byLovisato in a marlylimestone cropping outin the southern suburbsofthevillageofNurri, nearthecemeteryofthevillage (StrintudeMelonis)(Lovisato,1892).DelVecchio(1921)listing thecrocodileteethhousedinCagliari reportedasingletoothof

TomistomacalaritanusfromComidiano,betweenStrintuArgiolu andthetrain stationofNurri. Currently,theteeth fromStrintu deMelonisarelostwhereasthetoothfromComidianoishoused inTurinandaccessionedasMGPT-PU17336.Itisaslenderand

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pointedtoothcrown,21.6mmlong,withsparse,weaklongitudinal ridgesonthelabialandlingualsurfaces.

2.2.3. Locality

Laerru(Anglonasubregion,northernSardinia).

Ageandgeologicalsetting:PerfugasFormation(early-middle

Burdigalian, between 18.83±0.13 and 18.29±0.13 Ma; Oudet et al.,2010; Sowerbutts, 2000).Thefossiliferous site islocated onthenorthernperipheryofthevillageofLaerru.Anepiclastic pomiceus-cineriticlevelunderliessiliciﬁedlacustrinecarbonates ofthelowerpartofthePerfugasFormation(Sowerbutts,2000). Thisfossiliferousepiclasticlevelisdividedintothreedifferent lay-ers,themajorityofthevertebrateremainscomingfromthetopof thelowerlayer(ZoboliandPillola,2017b;Mennecartetal.,2019). IntheAnglonasubregion,thelowerpartofthePerfugasFm. com-prisescontinentallacustrinedepositsrepresentedbyﬁnelybedded limestonesandtuffs(Sowerbutts,2000).

Taxon:Crocodyliaindet.(=cf.Diplocynodonsp.inZoboliand

Pillola,2017b).

Material:MDLCA14114a-d,fourisolatedteeth(Fig.2D–G).

Descriptionandremarks:ZoboliandPillola(2017b)recently

described an interesting early Miocene vertebrate fauna from Laerru.Fourtiny(crownlengthofthebiggesttooth=0.9cm, bucco-labial diameter=0.3) and incomplete isolated crocodilian teeth werepreliminaryreferredtocf.Diplocynodonsp.However,dueto thegeneralized shapeoftheteethandthereforetheabsenceof diagnosticcharacters,thematerialisherereferredtoCrocodylia indet.NoteworthyisthatDiplocynodoniscurrentlyconsidereda Europeanendemicandwasneverreportedininsularenvironments asexpectedforanalligatoroidthatcouldhavesharedsalt-water intolerancewithextantalligatorids(TaplinandGrigg,1989;Delﬁno etal.,2007;DelﬁnoandRossi,2013).

2.2.4. Locality

Oschiri(Monteacutosubregion;northernSardinia).

Age and geological setting: “Formazione lacustre”

Auct.=Perfugas Fm. (Burdigalian, MN3; Van der Made, 2008; ZoboliandPillola,2017b).Thevertebratesite isrepresentedby asmallexposureoflacustrineclayinaroadcutnearthevillage ofOschiri(DeBruijnandRümke,1974).Theageofthemammal assemblageofOschirihasbeendiscussedbyseveralauthors(e.g.

DeBruijnandRümke,1974;AzzaroliandGuazzone,1979;Azzaroli, 1983;VanderMade,2008).Themicromammalsarequitediverse, withninetaxa includingsevenendemic species(DeBruijn and Rümke,1974).TheadvancedendemismofCtenodactylidaeandthe presenceofGliridae,TalpidaeandSoricidaeareverysimilartothe mainlandtaxa,andsuggestdifferentphasesofcolonizationofthe CorsicaSardiniaMassif(Ferrandinietal.,2000;Mennecartetal., 2017,2019).TheOligo-Miocenepalaeogeographicalcontextofthe westernMediterraneanindicatesthatthelatestadditionstothe insularassociationarrivedinSardiniaduringtheEuropeanLand MammalUnitMN3,whichiscoincidentwiththeearlyBurdigalian marineregression,around20Maago(VanderMade,2008;Zoboli andPillola,2017b).

Material:UUOS501,oneisolatedtooth(Fig.2H).

Descriptionandremarks:Theoccurrenceofcrocodyliansat

OschiriisalreadyknownintheliteratureaccordingtoDelﬁnoetal. (2011:379,tab.1)whoreportedthepresenceof“anunpublished isolatedtoothstoredinthecollectionsofUtrechtUniversity (spec-imenunnumbered)”.Thisisolatedtooth,nowUUOS501,israther largebecauseevenifdevoidofmostoftheroot,is28.6mmtalland hasalabio-lingualdiameterof11.3mm(themesiodistaldiameter isslightlysmallerbutthiscouldbeduetopreservationalreasons). Thetoothisconicalandslightlybentinlingualdirection.The sur-faceisvariablydamaged,neverthelessthemesiodisalcarinaiswell

visibleandsmooth.Theconicaldepressionthatislikelypresentat thebaseisstillﬁlledwithsediment.

2.2.5. Locality

LaCrucca(Sassari,Sassaresesubregion,northwesternSardinia).

Age andgeologicalsetting:MoresFormation (upper

Burdi-galian).Thecrocodylianfossilremainswerecollectedinacompact limestoneleveloftheMoresFm.(upperBurdigalian)croppingout inthelocalityofLaCrucca,abouteightkilometersWestofSassari (Spano,1985).

Taxon: cf. Tomistoma sp. (=Tomistoma calaritanus in Spano,

1985;Crocodyliaindet.inKotsakisetal.,2004).

Material:twolumbarvertebrae,twosacralvertebrae,two

cau-dalvertebrae,ilium,fragmentofleftischium,someribfragments, rightfemur,someosteodermsfromthecaudalregion(Fig.3).

Descriptionandremarks:ThematerialfromLaCruccabelongs

toasingleindividualbrieﬂydescribedandonlypartiallyﬁguredby

Spano(1985).Despitethefactthatwehavenotbeenabletolocate thismaterial,thankstothepublishedillustrationsitispossibleto referittoatomistominecrocodylianonthebasisofthebroadand anteroposteriorlyshortosteodermswhoseornamentationis rep-resentedbyrelativelyfewbutlargepits(compareFig.3Awiththe typeosteodermsofTomistomacalaritanuminFig.8;seealsoPiras etal.,2007;NojimaandItoigawa,2017;Shanetal.,2017). 2.2.6. Locality

MonteOrria(betweenNulviandChiaramonti,Anglona subre-gion,northernSardinia).

Age andgeologicalsetting:MoresFormation (upper

Burdi-galian).AmiddleMioceneagehasbeenattributedbySpano(1985)

tothematerialfromMonte Orria,insteadKotsakisetal.(2004)

indicatedalateMioceneage.However,onlytheupperBurdigalian littoralcarbonaticlithofaciesoftheMoresFm.isreportedinthe MonteOrriaarea.TheoriginallabelofMGPT-PU17337indicates thattheonlyavailabletoothfromMonteOrriawascollectedina brecciouslimestone.

Taxon: Crocodyliaindet.(=Crocodyliaindet. in DelVecchio,

1921;Spano,1985;Kotsakisetal.,2004;Tomistomacalaritanusin theoriginallabelofMGPT-PU).

Material:MGPT-PU17337,oneisolatedtooth(Fig.2I).

Descriptionandremarks:DelVecchio(1921)listingthe

mate-rialcollectedinCagliarireportedseveralsmallfragmentsofsmall teethfromMonte Orria,betweenNulviand Chiaramonti. How-ever,onlyoneisolatedtoothfromthislocalityiscurrentlyhoused inTurin.MGPT-PU17337isasmall,andpoorlypreservedcrown, 9.5mmlongthatisnearlycompletelyenamel-free.

2.3. Langhian-Serravallian 2.3.1. Locality

Tresnuraghes(Planargiasubregion,westernSardinia).

Age and geological setting: “Crabs-rich” marly limestone

(Langhian).

Taxon: Crocodyliaindet. (=Tomistoma sp.in Lovisato, 1892;

TomistomacalaritanusandCrocodyliaindet.inDelVecchio,1921; Crocodyliaindet.inKotsakisetal.,2004).

Material:notavailable.

Remarks:Lovisato(1892)reportedtwoisolatedteethfromthis

localityandreferredthemtoTomistomasp.DelVecchio(1921) list-ingthematerialhousedinCagliarireportedtwoteethdifferentin size(thesmallestreferredtoTomistomacalaritanusandthelargest toCrocodyliaindet.).Theseteetharecurrentlynotavailableandit isthereforeimpossibletoprovideadescription.

2.3.2. Locality

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Fig.3. cf.Tomistomasp.fromthelateBurdigalianofLaCrucca(Sassari),Italy.A,osteodermsfromthecaudalregion.B,articulatedvertebrae(twosacralsandonelumbar) andiliacbones,ventralview.C,incompleteleftischium.D,rightfemur.Scalebar:10cm.AllﬁguresmodiﬁedfromSpano,1985.

cf.Tomistomasp.,BurdigaliensupérieurdeLaCrucca(Sassari),Italie.A,ostéodermesdelarégioncaudale.B,vertèbresarticulées(deuxsacralesetunelombaire)etosiliaques,vue ventrale.C,ischiumgaucheincomplet.D,fémurdroit.Barred’échelle:10cm.Touslesﬁguresd’aprèsSpano,1985.

Age and geological setting: Argille di Fangario Formation

(?middleLanghian–lowerSerravallian).TheArgillediFangario Fm.isrepresentedbyclayeyand/orsandygray-yellowishmarls, developingupwardtomarlysandstones,withforaminifers,corals, brachiopods,echinoids,molluscsandichthyoliticfragments,The micropalaeontologicalcontentallowstorefer thissuccessionto the?middleLanghian–lowerSerravallian(Cherchi,1974).

Taxon:Crocodyliaindet.(=Tomistomacf.Tomistomacalaritanus

inCapellini,1890b; Tomistomasp.in Lovisato,1892;Tomistoma calaritanusinDelVecchio,1921;TomistomacalaritanusinSpano, 1985;Crocodyliaindet.inKotsakisetal.,2004;Crocodylus calari-tanusintheoriginallabelofMGPT-PU).

Material: MGPT-PU 17334, one incomplete isolated tooth

(Fig.2J).TwoisolatedteethﬁguredbyDelVecchio,1921(Fig.2K andL).

Description and remarks: Capellini (1890b) reported an

incompletetoothcollectedbyLovisatoatBingiaFargeri.Theauthor referredthefossiltoTomistomacf.calaritanusbutLovisato(1892)

referredthetoothtoTomistomasp.Lateron,DelVecchio(1921)

describedand ﬁgured two elongatedteethattributedto Tomis-tomacalaritanusfromBingiaFargeri(Fig.2Kand L).Inaddition, theauthormentionedathirdelongatedtoothfromthesame local-ity.TheonlyavailabletoothfromBingiaFargeriisnowMGPT-PU 17334.ItisnotpossibletocorrelatethesamplehousedinTurin withthematerialmentionedbutnotﬁguredbyCapellini(1890b)

andLovisato(1892).Furthermore,MGPT-PU17334istoodifferent fromthoseillustratedbyDelVecchio(1921).MGPT-PU17334isa massivecrown,21.5mmtall,withawornapexandthesurfaceof thebaseonlypartlypreserved.However,mostofboththelabial andlingualsurfaceispreservedanddistinctlysmooth.

2.3.3. Locality

Florinas(Sassaresesubregion,northwesternSardinia).

Ageandgeologicalsetting:FlorinasFormation(Serravallian).

TheteethwerefoundinasandquarryabouttwokilometersSouth ofthevillageofFlorinas.

Taxon:Crocodyliaindet.(=?TomistomalusitanicainMarrasand

Ventura,1985;Crocodyliaindet.inKotsakisetal.,2004).

Material:fourisolatedteeth.AccordingtoMarrasandVentura,

1985(pp.25),theseteethwerehostedinaprivatecollection:“Si ringraziailsig.AlbertoPigaperavercigentilmenteprestatoil mate-riale[Mr.AlbertoPigaisthankedforhavingkindlyloanedusthe material]”.Wehavenotbeenabletolocatethematerial.

Remarks:Theteethwerepreviouslydescribedandﬁguredby

MarrasandVentura(1985).Theauthorsdoubtfullyascribedthe fossilstoTomistomalusitanica(ViannaandMoraes,1945).

2.4. Tortonian-Messinian 2.4.1. Locality

FiumeSanto(PortoTorres,Nurrasubregion,northwestern Sar-dinia).

Ageandgeologicalsetting:Alluvialdeposit(Tortonian–early

Messinian).Severalisolatedcrocodylianteethwerecollectedin thesiteofFiumeSanto,inupperMiocenealluvialsedimentsof thethirdMiocenesedimentarycyclerecognizedinSardinia.The detailedstratigraphicdataandthegeologicalcontextofthesiteof FiumeSantoarenotsatisfactorilyknown,however,alate Torto-nian–earlyMessinanageissuggestedbytheassociatedTurolian mammalfauna(Abbazzietal.,2008b;Casanovas-Vilaretal.,2011).

Taxon:Crocodyliaindet.(=Crocodyliaindet.inKotsakisetal.,

2004;Abbazzietal.,2008b).

Material:49isolatedteeth(unumbered)hostedinthe

collec-tionsoftheSoprintendenzaArcheologia,belleartiepaesaggioper leprovincediSassarieNuoro(Fig.5MandN).

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Fig.4.Crocodyliaindet.,isolatedteethfromthe“Tramezzario”lithofacies(earlyMessinian)ofCagliari(southernSardinia).A,MGPT-PU17332/1fromIsMirrionis;A1,labial view;A2–3,lateralview.B,MGPT-PU17332/1fromIsMirrionis;B1,labialview;B2-3,lateralview.C,MGPT-PU17332/3fromIsMirrionis;C1,Labialview,C2,lingualview; C3,lateralview.D,MGPT-PU17332/2fromIsMirrionis.E,MGPT-PU17332/4fromIsMirrionis;E1,labialview;E2,lateralview.F,MGPT-PU17332/5fromIsMirrionis(near Piazzad’Armi);F1,labialview;F2,lateralview.G,MGPT-PU17332/5fromIsMirrionis(nearPiazzad’Armi);G1-2,linguo-lateralview.H,MGPT-PU17331/1fromPiazza d’Armi;H1,labialview;H2-3,lateralview.I,MGPT-PU17331/2fromPiazzad’Armi;I1,labialview;I2,lateralview.J,MGPT-PU17331/3fromPiazzad’Armi,labialview.K, MGPT-PU17333/2fromTuvumannu(alsocalledMonteSanGiuseppeorMontedellaPace);K1,labialview;K2,lingualview;K3-4,lateralview;K5,crosssection.Scalebar forA-C,E,F,H:1cm;scalebarforD,K:0.6cm;scalebarforG,I,J:1.5cm.

Crocodyliaindet.,dentsisoléesissusdeslithofaciès«Tramezzario»(Messinieninférieur)deCagliari(suddelaSardaigne).A,MGPT-PU17332/1,IsMirrionis;A1,vuelabiale;A2-3, vuelatérale.B,MGPT-PU17332/1,IsMirrionis;B1,vuelabiale;B2-3,vuelatérale.C,MGPT-PU17332/3,IsMirrionis;C1,vuelabiale,C2,vuelinguale;C3,vuelatérale.D,MGPT-PU 17332/2,IsMirrionis.E,MGPT-PU17332/4,IsMirrionis;E1,vuelabiale;E2,vuelatérale.F,MGPT-PU17332/5,IsMirrionis(prèsdelaPiazzad’Armi);F1,vuelabiale;F2,vue latérale.G,MGPT-PU17332/5,IsMirrionis(prèsdelaPiazzad’Armi);G1-2,vuelinguo-latérale.H,MGPT-PU17331/1,Piazzad’Armi;H1,vuelabiale;H2-3,vuelatérale.I,MGPT-PU 17331/2,Piazzad’Armi;I1,vuelabiale;I2,vuelatérale.J,MGPT-PU17331/3,Piazzad’Armi,vuelabiale.K,MGPT-PU17333/2,Tuvumannu(égalementappeléMonteSanGiuseppe ouMontedellaPace);K1,vuelabiale;K2,vuelinguale;K3-4,vuelatérale;K5,coupetransversale.Barred’échellepourA-C,E,F,H:1cm;barred’échellepourD,K:0,6cm;barre d’échellepourG,I,J:1,5cm.

Description and remarks: The general morphology of the

crocodylian teeth from Fiume Santo was already described in

Abbazzietal.(2008b)andthetwoisolatedteethfiguredherefor thefirsttimefitwellwiththatdescription.Thesmallest(Fig.5M; 22.10.1996,AreaIngressoNord;ZonaB),11.0mmtall,comesfrom theposteriorsectionofamaxillaryordentarytoothrow;ithas well-markedmesiodistalcarinaethatareevenmorehiglithened bya groovethat runsparalleltothem onboth thelingual and labialsurfaces.Thelargest(Fig.5N;23.10.1996,AreaIngressoNord; A1/BDIV;terradisetacciatura)ismassiveand22.1mmtall,itwas probablylocatedinoneofthelargestalveoliofthetoothrow.

2.4.2. Locality

CapoFrasca(Arbus,Monrealesubregion,westernSardinia).

Ageandgeologicalsetting:CapoSanMarcoFormation(early

Messinian).ThefossilwascollectedbyLovisatoinacompact lime-stone.

Taxon:Crocodylia indet.(=Tomistomasp. in Lovisato, 1892;

Crocodyliaindet.inDelVecchio,1921;Tomistomacalaritanusin

Spano,1985;Crocodyliaindet.inKotsakisetal.,2004;Tomistoma calaritanusintheoriginallabelofMGPT-PU).

Material:MGPT-PU17335,oneisolatedtooth(Fig.5L).

Descriptionandremarks:Thissmall,conicalfragmentcan

ten-tativelybeconsideredaportionofthecrownofacrocodyliantooth missingboththetipandthebase.

2.4.3. Localitiy

Tuvumannu=Monte San Giuseppe or Monte della Pace (Cagliari).

Ageandgeologicalsetting:CalcaridiCagliariFormation(late

Tortonian–earlyMessinian).ThetypematerialofTomistoma calar-itanumandseveralisolatedcrocodylianteethwerefoundinthe carbonatesuccessionoftheCalcaridiCagliariFm.(Gandolﬁand Porcu,1967;Cherchi,1974),croppingoutonlyintheCagliariarea. This successionis represented by three main lithofacies which are,fromthebottomtotop:“PietraCantone”,“Tramezzario”and “Pietra Forte”. Note that the names, informallyadopted in the mostrecentofﬁcialgeologicalmap(Barcaetal.,2005),arederived fromthenamesusedbyquarrymen(Georgalisetal.,2017).The bottom lithofacies (“Pietra Cantone”) is represented by yellow marly-arenaceouslimestonewithcommonintensebioturbation. The“Tramezzario”(or“Tramezzara”)isrepresentedbycalcarenites andmarlswithabundantbioclasticcomponents.Thislithofacies showswidespreadphenomenaofsynsedimentarybreccias, slump-ings, erosional surfacesand faulting. The uppermostlithofacies (“PietraForte”or“PietradiBonaria”)consistsofmassivecoarse biostromallimestones(Barcaetal.,2005).Thecarbonatesuccession ofCagliariisconsideredtobelateMioceneinage(Kotsakis,1985; ZoboliandPillola,2016b).TheTortonian-Messinianboundaryis tentativelyplacedwithintheupperpartofthe“PietraCantone” (Georgalisetal.,2017).

Crocodyilianremains werecollectedin differentlocalitiesof Cagliari since the beginning of the 19th Century: Tuvumannu, Piazzad’Armi,IsMirrionis(=MirioniinGennari,1868;IsMeriones

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Fig.5. Crocodyliaindet.,isolatedteethfromthelateMioceneofdifferentlocalitiesofSardinia.A,MGPT-PU17333/1fromthe“PietraForte”lithofaciesofTuvumannu (Cagliari),earlyMessinian,lateralview.B,MGPT-PU17331/4fromthe“PietraForte”lithofaciesofCimiteroMonumentalediBonaria(Cagliari),earlyMessinian;B1,lingual view;B2-3,linguo-lateralview.C,MGPT-PU17330/4fromthe“PietraForte”lithofaciesofMontixeddudiBonaria-CavaTimon(Cagliari),earlyMessinian;C1,labialview;C2, lingualview;C3,lateralview.D,MGPT-PU17330/8fromthe“PietraForte”lithofaciesofMontixeddudiBonaria(Cagliari),earlyMessinian;D1,crosssection;D2,longitudinal section.E,MGPT-PU17330/6fromthe“PietraForte”lithofaciesofBonaria(Cagliari),earlyMessinian;E1,labialview;E2,lateralview.F,MGPT-PU17330/3fromthe“Pietra Forte”lithofaciesofBonaria(Cagliari),earlyMessinian.G,MGPT-PU17330/5fromthe“PietraForte”lithofaciesofMontixeddudiBonaria-CavaTimon(Cagliari),early Messinian;G1,labialview;G2,lateralview;G3,crosssection.H–I,MGPT-PU17330/2fromthe“PietraForte”lithofaciesofBonaria(Cagliari),earlyMessinian.J,MGPT-PU 17330/1fromthe“PietraForte”lithofaciesofBonaria(Cagliari),earlyMessinian;J1,labialview;J2–3,lateralview.K,MGPT-PU17330/7fromthe“PietraForte”lithofacies ofMontixeddudiBonaria–CavaTimon(Cagliari),earlyMessinian;K1,lingualview;K2,latealview.L,MGPT-PU17335fromtheearlyMessinianofCapoFrasca(western Sardinia).M,Unnumberedtooth(22.10.1996,AreaIngressoNord;ZonaB)fromtheTortonian–earlyMessinianofFiumeSanto(northwesternSardinia);M1,labialview; M2,lingualview;M3-4,lateralview.N,Unnumberedtooth(23.10.1996,AreaIngressoNord;A1/BDIV;terradisetacciatura)fromtheTortonian–earlyMessinianofFiume Santo(northwesternSardinia);N1,labialview;N2,lingualview;N3-4,lateralview.O,fromanunknownlocalityofSardinia,indeterminateageandgeologicaldata(from DelVecchio,1921).ScalebarforA–E,G–K,O:1cm;scalebarforF:0.5cm;scalebarforL:0.3cm;scalebarforM,N:0.7cm.

Crocodyliaindet.,dentsisoléesduMiocènesupérieurdedifférenteslocalitésdeSardaigne.A,MGPT-PU17333/1,lithofaciès«PietraForte»deTuvumannu(Cagliari),Messinien inférieur,vuelatérale.B,MGPT-PU17331/4,lithofaciès«PietraForte»deCimiteroMonumentalediBonaria(Cagliari),Messinieninférieur;B1,vuelinguale;B2-3,vue linguo-latérale.C,MGPT-PU17330/4,lithofaciès«PietraForte»deMontixeddudiBonaria–CavaTimon(Cagliari),Messinieninférieur;C1,vuelabiale;C2,vuelinguale;C3,vuelatérale. D,MGPT-PU17330/8,lithofaciès«PietraForte»deMontixeddudiBonaria(Cagliari),Messinieninférieur;D1,sectiontransversale;D2,coupelongitudinale.E,MGPT-PU17330/6, lithofaciès«PietraForte»deBonaria(Cagliari),Messinieninférieur;E1,vuelabiale;E2,vuelatérale.F,MGPT-PU17330/3,lithofaciès«PietraForte»deBonaria(Cagliari),Messinien inférieur.G,MGPT-PU17330/5,lithofacies«PietraForte»deMontixeddudiBonaria–CavaTimon(Cagliari),Messinieninférieur;G1,vuelabiale;G2,vuelatérale;G3,coupe transversale.H–I,MGPT-PU17330/2d’aprèsleslithofaciès«PietraForte»deBonaria(Cagliari),Messinieninférieur.J,MGPT-PU17330/1d’aprèsleslithofaciès«PietraForte»de Bonaria(Cagliari),Messinieninférieur;J1,vuelabiale;J2–3,vuelatérale.K,MGPT-PU17330/7deslithofaciès«PietraForte»deMontixeddudiBonaria–CavaTimon(Cagliari), Messinieninférieur;K1,vuelinguale;K2,vuelatérale.L,MGPT-PU17335duMessinieninférieurdeCapoFrasca(ouestdelaSardaigne).M,Dentnonnumérotée(22.10.1996,Zone IngressoNord;ZonaB)duTortonien–PremierMessiniendeFiumeSanto(SardaigneNord-Ouest);M1,vuelabiale;M2,vuelinguale;M3-4,vuelatérale.N,dentnonnumérotée (23.10.1996,zoneIngressoNord;A1/BDIV;terradisetacciatura)duTortonien–MessinieninférieurdeFiumeSanto(nord-ouestdelaSardaigne);N1,vuelabiale;N2,vuelinguale; N3-4,vuelatérale.O,dentprovenantd’unelocalitéinconnuedeSardaigne,âgeetdonnéesgéologiquesindéterminées(d’aprèsDelVecchio,1921).Barred’échellepourA–E,G–K,O: 1cm;barred’échellepourF:0,5cm;barred’échellepourL:0,3cm;barred’échellepourM,N:0,7cm.

inCapellini,1890b;IsMirionesinKotsakisetal.,2004),Bonaria, CimiteroMonumentalediBonaria,CavaTimon–Montixeddudi Bonaria(Gennari,1868;Capellini,1890a,1890b;Lovisato,1892; DelVecchio,1921).ThetypeofTomistomacalaritanumwas col-lected in the “Tramezzario” lithofacies cropping out in the Is Mirrionisneighbourhood(Piazzad’Armi). Theassociatedreptile faunafromIs Mirrionisconsistsofthetypematerialofthe tur-tle Procyclanorbis sardus Portis, 1901, currently referred to an

indeterminatepan-trionychine(Georgalisetal.,2017),andofother fewisolatedpan-trionychineremains(ComaschiCaria,1958).

Taxon:Crocodyliaindet.(=Crocodyliaindet.inCapellini,1890b,

Lovisato,1892;DelVecchio,1921;Crocodylussp.in theoriginal labelofMGPT-PU).

Material:MGPT-PU17333/2,fromthe“Tramezzario”lithofacies

(Fig.4K);MGPT-PU17333/1,oneisolatedtoothfromthe“Pietra Forte”lithofacies(Fig.5A).

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Fig.6.TheholotypeskullofTomistomacalaritanumCapellini,1890fromthelateMioceneofPiazzad’Armi,Cagliari(fromtheoriginalplatesofCapellini,1890b).A,skull beforetherestorationindorsalview.B,interpretativedrawingsoftherestoredskull;B1,rightlateralview;B2,dorsalview;B3,leftlateralview.Scalebar:10cm. CrânedeTomistomacalaritanumCapellini,1890,holotype,issuduMiocènesupérieurdePiazzad’Armi,Cagliari(d’aprèslesplanchesoriginalesdeCapellini,1890b).A,crâneen vuedorsaleavantlarestauration.B,dessinsd’interprétationducrânerestauré;B1,vuelatéraledroite;B2,vuedorsale;B3,vuelatéralegauche.Barred’échelle:10cm.

Descriptionand remarks: Lovisato (1892) reported a large

size tooth from the “Tramezzario” of Monte San Giuseppe (=Tuvumannu)andreferredittoa“truecrocodile”.Lovisato(1829) andDelVecchio(1921)didnotprovidetheﬁguresofthe mate-rialfromTuvumannu,thereforeasurecorrelationwiththeteeth housedinTurinisnotpossibile.MGPT-PU17333/1(Fig.5A)isa par-tialcrown,about26mmlong,stillembeddedinablockofmatrix whereasMGPT-PU17333/2(Fig.4K)istheapex,14.9mmlong,ofa slenderandthintoothwithsharpmesiodistalcarinaedelimitinga broadlabialsurfaceandarelativelysmalllingualsurface(Fig.4K5).

2.4.4. Locality

Bonaria,MontixeddudiBonaria–CavaTimon(Cagliari).

Ageandgeologicasetting:CalcaridiCagliariFormation(late

Tortonian–earlyMessinian,GandolﬁandPorcu,1967,seeabove).

Taxon:Crocodylia indet.(=Crocodyliaindet.in Del Vecchio,

1921andKotsakisetal.,2004;Tomistomacalaritanusand “Crocody-lus”sp.intheoriginallabelofMGPT-PU).

Material:Allteetharefromthe“PietraForte”lithofacies.

MGPT-PU17330/1, oneisolatedtoothfromBonaria(Fig.5J);MGPT-PU 17330/2,twoincompleteteethfromBonaria(Fig.5HandI); MGPT-PU17330/3,oneisolatedtoothfromBonaria(Fig.5F);MGPT-PU 17330/4,oneisolatedtoothfromMontixeddudiBonaria–Cava Timon(Fig.5C);MGPT-PU17330/5,oneisolatedtoothfrom Mon-tixeddudiBonaria–CavaTimon(Fig.5G);MGPT-PU17330/6,one isolatetoothfromBonaria(Fig.5E);MGPT-PU17330/7,one iso-latedtoothfromMontixeddudiBonaria–CavaTimon(Fig.5K). MGPT-PU17330/8,oneisolatedtoothfromMontixeddudiBonaria (Fig.5D).

Descriptionandremarks:Theseisolatedandincompleteteeth

haveatotal lengthvaryingfrom5.4mm(MGPT-PU17330/3)to about42mm(MGPT-PU17330/7)oreventoabout45mminthe caseofMGPT-PU17330/8whosestateofpreservationcastssome doubtonitsidentityascrocodylian.AsshowninFig.5,someof themarestillembeddedinasmallblockofmatrix.Curiously,the onlypreservedmesiodistalcarinaofMGPT-PU17330/4bifurcates proximally(Fig.5C3).

2.4.5. Locality

CimiteroMonumentalediBonaria(Cagliari).

Taxon: Crocodylia indet.(=Crocodyliaindet. in DelVecchio,

1921andKotsakisetal.,2004;Tomistomacalaritanusintheoriginal labelofMGPT-PU).

Material:MGPT-PU17331/4,oneisolatedtoothfromthe“Pietra

Forte”lithofacies(Fig.5B).

Descriptionandremarks:theexposedportionof theparlty

embeddedcrownMGPT-PU17331/4isverywell-preservedand shows the wrinkles of the enamel surface extending onto the mesiodistalcarinae;consideringalsothebasalportionofthecrown thatisonlyminimallypreserved,theremainis23mmlong.

2.4.6. Locality

IsMirrionis–Piazzad’Armi(Cagliari).Theoriginallabelsofthe materialhousedatMGPT-PUindicatetwodifferentlocalities:“Is Mirrionis” and“Piazzad’Armi”. However,Piazza d’Armiis actu-allyincludedintheIsMirrionisneighborhood,therefore,weprefer togrouptogetheralloftheisolatedcrocodylianteethfromthese toponyms.

Taxa:Crocodyliaindet.(=Crocodyliaindet.inDelVecchio,1923

andKotsakisetal.,2004;Tomistomacalaritanusintheoriginallabel ofMGPT-PU).

Material: all fossils are from the “Tramezzario” lithofacies.

MGPT-PU17332/1,twoisolatedteeth(Fig.4AandB);MGPT-PU 17332/2,oneisolatedtooth(Fig.4D);MGPT-PU17332/3,one iso-latedtooth(Fig.4C);MGPT-PU17332/4,oneisolatedtooth(Fig.4E); MGPT-PU17332/5,twoincompletetooth(Fig.4FandG).MGPT-PU 17331/1,oneisolatedtooth(Fig.4H);MGPT-PU17331/2,one iso-latedtooth(Fig.4I);MGPT-PU17331/3,oneisolatedtooth(Fig.4J).

Description and remarks: Is Mirrionis: the sample of

iso-latedteethfromIsMirrionisischaracterizedbyarelativelysmall size,beingthesmallest(MGPT-PU17332/2)just9mmlongand thelargest(oneofthetwoteethaccessionedunderthenumber

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Fig.7.ThelosttypematerialofTomistomacalaritanumCapellini,1890fromthelateMioceneofPiazzad’Armi,Cagliari(fromtheoriginalplatesofCapellini,1890b).A, premaxillaeandmaxillaeinventralview.B,dentariesandsplenialsinventralview.C,sixthcervicalvertebrainlateralview.D,eighthcervicalvertebrainposteriorview. E–F,cervicalribs.Scalebar:5cm.

MatérielperdudutypedeTomistomacalaritanumCapellini,1890provenantduMiocènesupérieurdePiazzad’Armi,Cagliari(d’aprèslesplanchesoriginalesdeCapellini,1890b). A,premaxillaeetmaxillairesenvueventrale.B,dentairesetsplénialesenvueventrale.C,sixièmevertèbrecervicaleenvuelatérale.D,huitièmevertèbrecervicaleenvuepostérieure. E–F,côtescervicales.Barred’échelle:5cm.

MGPT-PU17332/5)about28mmlong(includingtheportionthat isparltyhidden).Someoftheteetharestillembeddedinawhitish matrix.Piazzad’Armi:MGPT-PU17331/1(Fig.4H)isanisolated, fragmentary crown, 27.2mm long, whereas MGPT-PU 17331/2 (Fig.4I)isa large,massivecrown,about36mmlong;thelatter isstillembeddedinthematrixandshowsprominentmesiodistal carinae.AlsoMGPT-PU17331/3(Fig.4J)isstillembeddedinasmall blockbutitisheavilydamaged;itisanelongatedtoothwithatotal lengthofapproximately39mm.Theseisolatedteeth,despitethe absenceofdiagnosticcharacters,maybelongtothetomistomine crocodyliandescribedbelow.

2.4.7. Locality

Piazzad’Armi(Cagliari).

Taxon:TomistomacalaritanumCapellini,1890(Fig.6).

Material:oneincompletecranium(MDLCA148)(Fig.9BandC),

eightincompleteosteoderms(MDLCA14401)(Fig.8A2,E–G,I,L3, M3,N2),castsoftherestoredcranium(Fig.9A)andsixosteoderms (Fig.8D,H2,J2,K2,L2,M2[collectionofDipartimentodiScienze ChimicheeGeologicheoftheUniversityofCagliari;notethatthese

castsareacopyoftheoriginalcastshousedintheMuseoGeologico G.CapelliniofBologna]).Partofthecranium(Fig.7AandB),some osteoderms(Fig.8BandC),cervicalvertebrae(Fig.7CandD)and cervicalribs(Fig.7EandF)werelostordestroyedduringtheSecond WorldWar.

The holotype of Tomistoma calaritanum Capellini, 1890:

Thecomparisonofthedescriptionsandillustrationsprovidedby

Capellini(1890b)withthecurrentstateofpreservationofMDLCA 148(Fig.9B)clearlyindicatesthatmostoftheskeletalelements andthemorphologicalinformationtheyprovidedisnowlost.As emphasizedbyCapellini(1890b,p.514),theﬁguresinhis publica-tiondonotclearlyidentifywhatisrepresentedbyplasterversus realbones.However,thecoloroftheplastercastoftheoriginal restoredskull(Fig.9A)representstheonlyway todiscriminate therealextentoftherestorationworkperformedbyCapellini.The mainskeletalelementsstillpreservedaretheposteriorregionof boththemaxillae,partofthenasalsandquadratojugals(Fig.10). Thelowerjawstillpreservestheposteriorregionofthedentaries, partof thesurangular and angular (the latter only onthe left side).Thesurfaceoftheblockpreserves themorphologyofthe innersurfaceofpartoftheelementsofthedorsalarea(the poste-riorareaofthenasalsandmaxillaeinparticular).Moreover,very

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Fig.8. TypeosteodermsofTomistomacalaritanumCapellini,1890,fromthelateMioceneofPiazzad’Armi(Cagliari),dorsalview.A,cervicalosteoderm;A1,asinCapellini, 1890b(tav.IV,Fig.1);A2,ascurrentlypreserved(MDLCA14401).B,cervicalosteoderm(lostelement)asinCapellini,1890b(tav.IV,Fig.2).C,nucalosteoderm(lostelement) asinCapellini,1890b(tav.IV,Fig.3).D,incompletenucal(?)osteoderm,plastercastofalostelement.E,incompletenucal(?)osteoderm(MDLCA14401).F,incompletenucal (?)osteoderm(MDLCA14401).G,fragmentofosteoderm(MDLCA14401).H,dorsalosteodermofthemedianseries(lostelement);H1,asinCapellini,1890b(tav.IV,Fig.4); H2,plastercast.I,dorsalosteodermofthemedianseries(MDLCA14401).J,dorsalosteodermofthemedianseries(lostelement);J1,asinCapellini,1890b(tav.IV,Fig.5);J2, plastercast.K,dorsalosteodermoftherightlateralseries(lostelement);K1,asinCapellini,1890b(tav.IV,Fig.6);K2,plastercast.L,dorsalosteodermofthemedianseries; L1,asinCapellini,1890b(tav.IV,Fig.7);L2,plastercast;L3,ascurrentlypreserved(MDLCA14401).M,dorsalosteodermofthemedianseries;M1,asinCapellini,1890b (tav.IV,Fig.8);M2,plastercast;M3,ascurrentlypreserved(MDLCA14401).N,dorsalosteodermoftherightlateralseries;N1,asinCapellini,1890b(tav.IV,Fig.9);N2,as currentlypreserved(MDLCA14401).Scalebar:5cm.

OstéodermesdutypedeTomistomacalaritanumCapellini,1890,duMiocènesupérieurdePiazzad’Armi(Cagliari),vuesdorsales.A,ostéodermecervical;A1,d’aprèsCapellini,1890b (pl.IV,Fig.1);A2,telqu’ilestconservéactuellement(MDLCA14401).B,ostéodermecervical(élémentperdu)d’aprèsCapellini,1890b(pl.IV,Fig.2).C,ostéodermenucal(élément perdu)d’aprèsCapellini,1890b(pl.IV,Fig.3).D,ostéodermenucal(?)incomplet,moulageenplâtre,élémentperdu.E,ostéodermenucal(?)Incomplet(MDLCA14401).F,ostéoderme nucal(?)Incomplet(MDLCA14401).G,fragmentd’ostéoderme(MDLCA14401).H,ostéodermedorsaldelasériemédiane(élémentperdu);H1,commedansCapellini,1890b(pl.IV, Fig.4);H2,mouleenplâtre.I,ostéodermedorsaldelasériemédiane(MDLCA14401).J,ostéodermedorsaldelasériemédiane(élémentperdu);J1,commedansCapellini,1890b (pl.IV,Fig.5);J2,mouleenplâtre.K,ostéodermedorsaldelasérielatéraledroite(élémentperdu);K1,d’aprèsCapellini,1890b(pl.IV,Fig.6);K2,mouleenplâtre.L,ostéoderme dorsaldelasériemédiane;L1,d’aprèsCapellini,1890b(pl.IV,Fig.7);L2,mouleenplâtre;L3,danssaformeactuelle(MDLCA14401).M,ostéodermedorsaldelasériemédiane; M1,d’aprèsCapellini,1890b(pl.IV,Fig.8);M2,mouleenplâtre;M3,telqu’ilestactuellementconservé(MDLCA14401).N,ostéodermedorsaldelasérielatéraledroite;N1,d’après Capellini,1890b(pl.IV,Fig.9);N2,telqu’ilestactuellementconservé(MDLCA14401).Barred’échelle:5cm.

fragmentaryskeletalelementsinsectionarepreservedonthe ante-rior,posterodorsal,and posteriorsurfacesoftheblock.Theydo notproviderelevantmorphologicalinformationandinmanycases theycannotbeidentiﬁedwithcertainty.

The currentlyavailable eight osteoderms, collectively acces-sionedasMDLCA14401(Fig.8),arevariablyincompletebutshare thesameexternalornamentationcharacterizedbyrelativelyfew andverylargepits.Theanteriorsmoothsurfaceis anteroposteri-orlyshortandthelongitudinalkeelisratherlow.Thelargestofthe osteoderms(plastercastFig.8H2)is68mmwideand92mmlong.

Capellini(1890b) reportedthepresenceofabouttwenty osteo-derms,butﬁguredonlyninespecimens(theoriginalphotographs byCapelliniarehereshowninFig.8A1,B,C,H1,J1,K1,L1,M1, N1).Fiveofthesenine osteodermsﬁguredbyCapellini(1890b)

arecurrentlylost(Fig.8B,C,H1,J1,K1),buttheplastercastsof threespecimensarestillavailable(Fig.8H2,J2,K2).Moreover,four

incompleteosteodermsoriginallynotﬁguredbyCapellini(1890b)

arecurrentlyavailable(Fig.8E,F,G,I)andtheplastercastofalost specimenisavailable(Fig.8D).

Capellini(1890b) listedand brieflydescribedseveral,mostly incompletevertebrae(fragmentsoftheI–IIcervicalvertebrae,the III,IV,VI,VIII,IXcervicalvertebrae,fragmentsofthefirstdorsal vertebrae,onelumbarvertebraandonecaudalvertebra)but fig-uredonlytwoofthem(tavIV,Figs.10and11inCapellini,1890b;

Fig.9C,Dinthispaper).Inaddition,Capellini(1890b)ﬁguredtwo cervicalribs(tav.IV,Figs.12and13inCapellini,1890b;Fig.7E,F inthispaper).Allvertebraeandcervicalribsarecurrentlylost.

Remarksandnomenclaturalissues:Gennari(1868)

condition-allyproposedthenameCroc.Caralitanus(=Crocodyluscaralitanus) forthetaxonfromPiazzad’Armi.Sincethepaperwaspublished before1961,accordingtotheArticle11.5.1ofICZN(1999)thename isnot tobeexcludedjustbecauseoftheconditionalcontextin

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Fig.9. TomistomacalaritanumCapellini,1890(MDLCA148).A,plastercastoftherestoredskull;A1,dorsalview;A2,leftlateralview;A3,rightlateralview.B,thecurrent stateofpreservationoftheskull;B1,anteriorview;B2,dorsalview;B3,lateralleftview;B4,lateralrightview.C,detailoftherightteethrow;C1,buccalview;C2,anterior view.ScalebarforA:10cm,scalebarforB:5cm,scalebarforC:1cm.

TomistomacalaritanumCapellini,1890(MDLCA148).A,moulageenplâtreducrânerestauré;A1,vuedorsale;A2,vuelatéralegauche;A3,vuelatéraledroite.B,l’étatactuel deconservationducrâne;B1,vueantérieure;B2,vuedorsale;B3,vuelatéralegauche;B4,vuelatéraledroite.C,détaildelarangéededentsdroite;C1,vuebuccale;C2,vue antérieure.Barred’échellepourA:10cm,barred’échellepourB:5cm,barred’échellepourC:1cm.

whichitwasproposed.Itisactuallyanavailablenamebecause Gen-nari(1868:128–129)providedabriefdescriptionfocusingonthe elongatedsnout,onitslengthversusthelengthoftherestofskull, andonthenumber(24/20=84),thedisposition,andtheshapeof theteeththatresembleGavialisgangeticusandCrocodylusacutus. Fewdecadeslater,Capellini(1890a,b)clariﬁedthatthespecimen wasactuallyatomistominecrocodylian.

Regardlessofthegenericreferral,evenifthespeciﬁcepitheh CrocodyluscaralitanusGennari,1868isavailable,itishereproposed areversalofprecedenceinfavorofTomistomacalaritanusCapellini, 1890because:

• theseniorsynonymwasneverusedafter1899(article23.9.1.1; ICZN,1999)(apparentlyitwasusedonlybyGennariin1868;and thenmentionedbyCapellini,1890,asthenamewrittenonthe tagofthespecimenbyGennarihimself),and;

• thejuniorsynonymhasbeenusedinthelast50yearsinatleastin 25works(sometimesasT.calaritanum,seebelow;Kotsakisand Palombo,1979;EsuandKotsakis,1983;Spano,1985;Comaschi Caria,1986;AntunesandGinsburg,1989;Rossmannetal.,1996; Brochu,1997,2001;Crespo,2001;Kotsakisetal.,2004;Weems, 2006;Pirasetal.,2007;MasandAntunes,2008;Delfino,2010; Delfinoand Ragazzini,2010; Fanti, 2010;Gerali, 2012, 2014; Delfinoand Rossi, 2013; DelfinoandRook,2008b; Abate and Fanti,2014;Delfinoetal.,2014;Jouveetal.,2014;Georgalisetal.,

2016,2017;ZoboliandPillola,2016b)publishedbyatleast10 authorsinatimespanencompassingmorethan10years(article 23.9.1.2;ICZN,1999).

Therefore,itishereformallyproposedtomaintainthe prevail-ingname Tomistomacalaritanus Capellini,1890that becomes a nomenprotectum,whereasCrocodyluscaralitanusGennari(1868)

isanomenoblitum.However,worthnotingisthatKotsakisetal. (2004)andPirasetal.(2007),consideringthatthegenderofthe epithetTomistomaisneutral,correctlymodiﬁedthespeciﬁc epi-thetinTomistomacalaritanum.Areassessmentofthevalidityofthis specieswillbethesubjectofanotherprojectthatrequiresabroad comparisonoftheholotypewithseveralotherextincttomistomes thatwerediagnosedandnamedinthepast.

2.5. Unknownage 2.5.1. Locality

unknown.

Ageandgeologicalsetting:unknown.

Taxon: Crocodylia indet. (=Tomistoma calaritanus in Del

Vecchio,1921).

Material: one isolated tooth ﬁgured by Del Vecchio, 1921

(Fig.5O).ThespecimenisnotpresentinthecollectionofMDLCA andMGPT-PU.

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Fig.10.TomistomacalaritanumCapellini,1890(MDLCA148),preservedskeletal elementsoftheskull.A1,dorsalview;A2,rightlateralview;A3,leftlateralview. Abbreviations:An,angular;Su,surangular;De,dentary;Ju,jugal;Ma,maxilla;Na, nasal.Scalebar:5cm.

TomistomacalaritanumCapellini,1890(MDLCA148),élémentssquelettiquespréservés ducrâne.A1,vuedorsale;A2,vuelatéraledroite;A3,vuelatéralegauche.Abréviations: An,angulaire;Su,surangulaire;De,dentaire;Ju,jugal;Ma,maxillaire;Na,nasale. Barred’échelle:5cm.

3. Conclusions

ThecrocodylianremainsfrommostoftheSardinianlocalities areexclusivelyrepresentedbyisolatedteeththatarenotdiagnostic beyondtheordinalleveldespitetheiroriginalreferral(inmuseum labelsorpublishedpapers)toCrocodylus,Diplocynodonor Tomis-toma.Therelevantexceptionisrepresentedbytheremainsfrom Piazzad’Armiin CagliarireferredbyGennari(1868)tothenew speciesCrocodyluscaralitanusandlateronbyCapellini(1890a,b)

toTomistomacalaritanus.Hereweprovideevidenceforthe rever-salofprecedenceinfavorofthelattername,that,however,being Tomistomaneutral,shouldbeTomistomacalaritanum.Theholotype wasarelativelywell-preservedskull(andassociatedosteoderms

andvertebrae)inoriginbutwasdamagedduringWWIIandthen partlylost.TheonlyotherinformativematerialfromSardinia con-sistsofsomeosteodermsandotherpostcranialelementsthatare nowapparentlylost;itcomesfromthelateBurdigalianofLaCrucca and canbetentatively referred toa tomistomine. Sinceextinct tomistomines are generally foundin coastalenvironments and thereforetheywerelikelyabletoeasilydisperse,thevalidityof Tomistomacalaritanumshouldbereassessedin thecontext ofa broadrevisionoftheTethyseantomistomines.

Disclosureofinterest

Theauthorsdeclarethattheyhavenocompetinginterest.

Acknowledgements

EdoardoRazzettihelpedusindisentanglingthenomenclatural issueofTomistomacaralitanus.WearegratefultoDaniele Ormez-zanoforhisassistanceduringouranalysisofthematerialhoused atMGPT-PU.Fig.5Mand Ncourtesyof NellaTuveri, Soprinten-denzaArcheologia,belleartiepaesaggioperleprovincediSassari eNuoro.Figs.2,8and9courtesyofMDLCA.WilmaWesselsand HansdeBruijn(UU)arethankedfortheextendedloanof mate-rialfromOschiri.WegratefullyacknowledgetheSardiniaRegional GovernmentfortheﬁnancialsupporttothePh.DscholarshiptoDZ (P.O.R.SardegnaF.S.E.OperationalProgrammeoftheAutonomous Region of Sardinia, EuropeanSocial Fund 2007–2013– Axis IV HumanResources,Objectivel.3,LineofActivityl.3.1.).Theresearch wassupportedbytheUniversitàdiCagliariCAR Project, “Paleo-biodiversità:strumentodibaseinbiostratigraﬁa,inpaleoecologia enellavalorizzazionedei beniculturali Geo-Paleontologici”led byGLP. ProjectsupportedbyFondi diAteneodell’Universitàdi Torino(2016–2017),GeneralitatdeCatalunya(CERCAProgram), andSpanishAgenciaEstataldeInvestigación(CGL2016-76431-P, AEI/FEDER,EU)toMD.WewouldliketothankreviewersAngela BuscalioniandJérémyMartinforcommentsthatgreatlyimproved thequalityofthemanuscript.

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