Ririsr.r lt.rli.rn.r ch l'.rleontoìosir c 5trlisr.rfir rolunrc ll3 PP.7t9-71i D.ccrrbcr l0C.{

SHORT NOTE - \-OTA BREVE

VIVERRA HOWELLI N, SP, A NE\I VIVERRID (CARNIVORA, MAMMALIA) FROM THE BACCINELLO-CINIGIANO BASIN (LATEST MIOCENE, ITALY)

LORL,NZO ROOK & BIENVENIDO MARTINEZ-NAVARRO''

{,.,/:J./.V /'/'. -'CJ .,. ,tf.r/ tr-tl : :A'l

Air ;onlr: ll.rnrnr.rli.r. C.rmiror'.r. \'rcnin.re. \er 5pecies, l.rre r ilerrid :rre ven' elusilc in Èurope. A viterrid nrandible TLLrclirn, l\lcssìniu. lr.,ll Iiuropr. Airìc:r. hirs been dcscribed from thc ltrlian,rssemblagc Baccincllo V3 (Larest Mìocenci MN 13) as )'iterra sp. bv Rook et .lbvrrtt. \\e dercribc.r nspecies oi \,'rz'enz is found in r.ff.rnca (I!lli,ì. Libir) r,ì in Esr Atri.r (Kenri). \'lorloIrqicrnrent( si lhc Lower Pìioccrrc (Ruscinian) of sorLthern Francc, rìt r.rr.rrrcr-ìzz.r pcf r\cr! ul.r r,reli.ì r cl.tii.rm.'rìr c prccol;r c per un dcnre J\'lonrpellicr and Roussillon (MN l.l and MN l5 respec- t!f,r{, ;rit, i(,f( .on un r.rllonidc rri<'rro. tivcllJ, rvhcre \titerra pepratxi rs prcsent (Dcperet 1390). T)rc same spccies ìs also recorded at "Fornrcc RDB quar- Introduction rr'" (Itah; Larlv Villrrfranchirn, MN l6:r) (Azzaroli er al. 1988). Dr.rring Earìv Pliocenc anotlter rchted genùs is Viierrids rre scrnrilv rcples('nred jn the Mio- knon'n to be disrrjbuted in Europe: Megavi."'erra (Krer- I)liocenc L-uropearr fossiÌ record. Thc earliest knorvn zoi & Feifar 1982). This gcnus hlls been erecred on thc l.uropean lcpreseDtJtives of rhc sub-lamìlv Vir.errinire bLrsis ot nrrterial from lvanovcc 1 (Slovakia; MN 15) ar- clirte to the bcginnint of thc lVlioccnc, n'hen madc irs trìbuted to the species l,legz,i;icerra cdrpatorun. The g,e- first ,rppc.rr'.rnce So)/gereff.. (Hclbing 192l, 1928; Ensess- nus is also recorded in the Earll Villafranchi:rn of Itrh. el 1972), i gcnus that survì\'ed untiì the Virllcsian (Pet- ('lriversa, MN 1(ra), *-ith.r cliffercnr species, Megaviaerra tef 1976; ùlonto\',r ct al. 2001). The earliesr rcpreseDra- dpcnniDicd (Kretzoi & Fejfrr 1982). tiles of the genus \.ii.,,-ra occur in thc dcposits of Vicr.rx Colìongcs (Fr:uce; unit llN.{ in the Europcirn nrammll The Baccinello-Cin igiirno brsin (soutì.rern Tus- bicrchrorolor:r') there are clcscribed as \titerra nodic,t cinl', Italy) is knorvn for its Lete Miocene vertebrate !ctrlrt, (Virct l95l; Dc Be:runront 1973). This species record (Bcnvenuti et al. 20Ol). Latc Miocene (MN l3) is irls,l prcscnt rr irnorhcr Flerrch sirc, at Lr Crive (MN n'ìanlmal remains arc founcl in several sites, from out- 7/3). rvith .r differe'nt subspccies, n 1\el\,\'ire11'd t tó.lica crops of the upper part of tl.re fluvio-l'rcusrrine scdi-

t od i ca (yier 1 95 1 ; cìe Be.runront 1 973). Durine Turoli:rn nrcnt:lry succession of thc basin. This nrantmal asscm-

l)ip.rIrinlclltodìSci,:nzedel|:r,lir.rl,Unir'cnitidilljrenzeii.lG'L.rI,ir.r,.{-5012lllIorence,Itrl1:I-m.rìl:ìlllktrrco.uniIi'it ICREA. Crup d'Auroccologi.r Humor. Arer de Prch;sròf; (Unitrt As<,cirdr rl CSIC). Universit;rt Ror ira i Virgìlì - Plaqr Inrperial Trrrrco | +ì005. Trrr:sorr. Sprin. I:nril: bnrrn:rr(rl ptehistorir.uÍr:es 724 L. Rook t< B. llarthtez Natarro

Fig, 1 - Vrsena lnuellì n. sp., holo- rlpe IGI;289C\: il righ! men- dibulrr r,rnus in lingu.rl ricw; b) right nrandibul.rr ranus in occlusal lies: B:rr scrle rcpre- senrs 1cm.

1cm b

blage s,as named V3 (after Lorenz 1968) to distinguish porar\'liYerrid from Sahabi (Vicerra n. sp. "A' in Howell ìt from older mammrl faunas (knorvn as V0, Vl, and V2) 1987) and from Lothagam (Viverrinae sp. ìnder.; \lerdelin rhar are present in the sedimentarl, sequence of the b:r- 2003) rre re-describcd and attributed to a new species. sin. These older faunas, which datc to the Ìate Miocene (Turolian Man.rmal Age), are made up bv cndemic fos- sil taxa (including the fossil ape Oreopitbecùs bambol- Systematic Paleontology ii), suggesting that the Tlsco-Sardinian region was an insular domain characterised by a faunlJ province dis- Order Carnivora Boq'dich, 1821 tìnct from the provinces of the surrounding coDtinental Suborder Ailuroidea Floweq 1869 masses. This isolation of the Tusco-Sardinian paleobio- Famiìv Viverridae Gral', 1821 province came to an cnd with the Messinian, when the Subfamil"- Viverrinae Gill, 1872 region was reached by fornrs rypical of the European faunal bioprovince (Hurzeler & Engesscr 1976; Rook Gews Vioerra Linné, 1758 l99l; Bernor et al. 2001). The Baccinello - Cinigiano succession was studied in detail by Lorenz (1968) and, Vlverra howelli n. sp. more reccnrly' bv Benvenuti er al. (1995, 2001) ;rnd Iiig l Rook et al. (2000). The fossiliferous sire of Caprarecce was first discov- 1987 \'i!,e//, n. sp. 'A , Honell, Fig. 2r ered by Mr. Malpassi, and a certain number of sp€cinìens l99l - \'iteza n. sp., Rook et al., Pl. l, iigs.2-3 2oo3 - Viverrinrc sp. indet., Werdelin, Iig. 7.13 were collectcd either by Mr. Malpassi and b-u-. rescarchcrs of Florence. of the Earth Sciences Dept. of the University Holotype. IGF 2390v (Fig. 1), right mandibulir rrmus bearing Some of these sanpÌes have been already describcd (Rook loser c.rnine (;solrred), rr;ors of P,. P.,l' .rnd f, (fourth prcmolat still er al. l99l; Rook Ei Rusrioni 1991; Rook & Torre 1995; presen ing a smrll talonid portion), conrplete iirsr los er molar (c.rrnas Abbazzi 2001) and rhe folloving taxa havc been at present sial),.rnd the rlveolus of M.. recognized in the assemblage for this localitl': Repositon. N.rrural Hìstorr Museum, The Universitv of Flor- cnce, Irrrll The specimen is kept ìn the collections of the "Ceologv rnd Tapirus c[. araemensis Croizet Er 1828, Pro- Jobert l'aleontologv Secrìon" s,irh inlentory number "lGF 2890v". caPreolus cf. lócryti Pohlig l9l l, Cervidae indet., Kolry2 Type locality. C:rprarccce, 100 nr So oi Pogg;o Vrnnini , Bac- ocboerus paleochoelrr (Kaup 1833), Vioerra sP., Celaden cinello-Cinieiano basin (Crosseto. southern Tusc:ny, Itrly). sia grossetand Rook & Torrc 1995, Alilepus sp., Castor cl. Chronologicd distribution. Mcssinirn, lrte Turolian, MN l3 praefiber Déperet 189A. unir in the Europern nrmmal biochronolosy. The r.iverrid mandible from this site has been de- Diagnosis. Mindibular ramus f.rirll sturdv rvirh loser profilc slightlv .rrched. I-ower c:rrn:rssial stoudy built liigonid makes up rbout s cribed as Vloerra sp. by Rook er al. (1991 ) . The specimen r""-thirJs of thc tooth Metrconid n'ell developed' almosr of the samc was recognized as b;longing ro a hitherro undescribed spe- cie., but the decisìo. *^,ir." "o,ìì1,;:l;;;j.::. ::f::illljì:""T1'jff;;:ru:Xlfi::?:,:,:',r",:$::,:il on it unril wider comparisons would had been made. In in shrpe, and *.;rh: o,ide occlusal basin ringetl by three cusps, an hy- rhis note the Baccinello specimen, as well as thc contem- poconid, a sn.rll hypoconulid,:nd a srouter entoconid. Viverra howelli z. sp., a ne.- Latest M;ocene ritetrid 721

metaconid. '12 00 The M,talonid is relativeÌy short, and charac- terized by the occurrence of accessory cusplets, either on 10.00 the buccal or on rhe lingual side. The second lower molar is not preserved bur its alveolus indicates 800 rhe occurrence of ! an elongated roorh. The mandibular ramus is slender; the È posterior margin of mandibular symphysis is observable on the lingual side of the ramus, reaching the level of P. alveo- li. The ma,,ereric for.a i. riso observable in irs end,.rr rhe 2.00 level of the second Ìower molar alveolus (Fig. 1). Viverra hotpelli n. sp. is clearly distìnguished from

300 '000 1200 .00 600 300 _000 thc genus À4egaaicera by irs smaller dimensions (Fig. 2) lowsr M1 Le.qtb and by its stout trigonid n.rorphologv. In terms of size, Vìoera hotoeLli n. sp. falls between Vrz,era modica ssp, and Viaeta peprani Deperer 1890 (Table 1; Fig. 2). The smalÌ viverrid from Sahabi (Libya; late Tì.rrolian), represented by l0 00 a right M, and described as Viz,erra n. sp. "A' by Howell (1987), is fully comparable by size (Fig. 2) with Vioera É bou.'elli n. sp.. In addition, also the fragmentary mandible 3 from the Lower Nawata Fmt. (Lothagam) recently de- scribed as Viverrinae sp. indet. (\ll'erdelin 2003) is morpho- 4.00 logically comparabl e wrthVit..ena bowel[ n. sp.. The Loth- 240 agam viverrinae sp. indet. is just larger in carnassial length in respect to the BaccinelÌo V3 type specìmen and Sahabi 000 M but shows the same 2,00 4,00 6.00 Loo ro oo proportions in talonid reduction -. low€r M1 Talonid Lenglh (Fig. 2) . As a matter of fact, in addition to the very sin.ri- lar size, it is noticeable ro nore rhar these three Messinian Fig2 - Bivarirte plot diagrrm ofM Length vcrsus (rbovt M, \ridrh spccimens have a relatively short M, talonid in respecr ro V and M, Taìonìd Lengrh versus M, Width (belod. Legend: pepratxi and to the larer (and larger) European Megaùver- Vìl:erra nodica, solid circLcs; VìaeÍ't hateUì n. sp.j open ra. s+ares; Vhetd peptut\1. solid rrìangler yrrerra |eabe1, open (Fig.2). The Baccinello V3 specimen shows instead a diattlonds; ùeSartrelr, spp., crosscs. shorter trigonid like the |arger Vit)eft.:leabey Hendey 1974 from South Africa (Langebaanweg) and Ethiopia (Omo), as does Sahabi and Lothagam M,s (Fig. 2). For the similarity in size and proporti ons,"Virterra n. sp. lf'from Sahabi P28A site (Howell, 1987) and the Viverrinae sp. indet. from Lothagam (Verdelin 2003) are EtvmoÌogy. Thc species ìs namcd afier prof. Clark F. Houell, here considered for hìs contrìbutìon in the srudy of Mio-Pliocene carnivores rnd for conspecific wìrh Vixetra houelli n. sp. his r eference in authors' profession:rl c:rrier Discussion. As already poìnted out, the fossil Measuremcnts. See Table 1. record of the genus Villera is discontinuous in Europe, since ìts first occurrence in the early Miocene. Vicer- Description and comparisons. Vioera hotoelli n. sp. ra hoteelli n. sp. represent a latesr Miocene (Messinian) is morphologically characterized by a quite robusr lower small sized viverrid with a geographic distriburion from canine, with a root and crown circular in section. The loe/- circum-medìterranean area and Eastern Africa, identi er canine crown has v.eil developed longitudinal furrows, lied at Baccinello V3 (Itaìy), Sahabi (Libya) and Loth- both on the external and the internal sides. The premolars agam (Kenya). are represented only bv their roots in the alveoÌi and small Durìng latest Miocene in Africa the genus Viveta crown fragmenrs (P., P. and P,). The first lower premolar is was present, in addition to Sahabi and Lothagam, with a very small in size and its alveolus is located just behind the Jarger species (Viz,ena leabey'1 at Langebaanweg (Messìn- posterior malgìn ofthe canine alveolus (with no diastema). ian or early Pliocene; Sourh Africa; Hendey 1974, 1981). A small diastema occurs berween P, and rhe anterior alveo- Vivena leahey has a longer chronological distribution, be- lus o{ the double roored second lower premolar As far as ing documented untiÌ the early Pleistocene of the Omo can be seen, second and third lower premolars are narrow valley, in the Tanganika area (Petter 1963; HoweÌl 1987). and slender. The small talonìd fragment preserved seenrs ro The first Asian representative of this subfamily ap- indicate that the fourrh lower premolars q.as cnlarged pos- peared during the Astaracian (middle Miocene) ìn Mon- teriorly, aìthough the occurrence of accessory cùsps cannot golia (Tung Gur Fm.). Its extremely primitive character- be determined. The first lower molar (the carnassial) has a istics result in ìts assignment to a separate genus, Tungu- paraconid intermediate in height between protoconid and rucrì (Colbert, 1939). The Siwaliks HiÌls have produced 722 L. I xtk E; B. .llarthrcz-N,tiarro

and rìarrowncss of the skull, chirrircteristics he fecls en- specès Lenorh wdrh Ia on d Loca 'ly tirelv preclude its assignrìrent ro Lèn9rh ilr'ì\.cxisting vilerrid. Pil- grinr (1912) ;rlso dcscrìbecl frrsnlent.rrl right mandibu- '+,-oÉr. " ". : : é.0òoF. 'r 560 330 25o LeGtue larbrrnch bear-ing l.-lv{ (l:rbelled GSI D 135) .rs "LLrtrinee

Viv.hahaúellt 10.40 530 400 Baccn€ ov3(CapÈrecce) gcn et sp. nor: /rasrioti". Thc spccirlen l-,rs collected fronr dcposits con elrted u'ith thc Dhoh Paran Fnr. According to Pilbernr er al. (1979),rnd HoselÌ (1937) rhis fossil is v)vena peptal, 1250 6 50 600 Rouss on better rrtributable ro vir,clrid .rnd probabll, lcprcsenrs 'r Vtveúleakey 15.50 730 -610 La^gebaanweg a nerr' llrge-sizcd gcnus. Vil.ia tèakay 1600 760 Langcbaaiwèg \/it-erra peì rvls crc.cted or'Ì thc llisìs of netcrial lron "Cap" Tr-alertinc ar Zoukoudirn b1' Qiu (19S0), ind sccrlrs to be a forn closc to thc getlt),s llegdti!.-crra. vNù6 teakèy 16.00 3.30 6 to oro vsrey

16 30 3.50 O'no var€y

htègÀ"@ carpatat@ 19 1a 1a aD 7 70 rva.ovce-l Conclusive remarks Mèa.vN,èra apènnrnrca 1630 360 Thc nervll describecì r'iler-rine spccies \/it'en'a houeLli Trb. I - Ve:rsurenents (in nlnt) ol \l ìn so,ne Vircnin.re spccies. D.ua trcu de Bc:run,onr (197.1), Hotrell (l9S/), Krerzoj N hld :r llrest Mioccnc distribution from circunr-Meditcrr.r- fcifrr (l9Sl). \\irdeliD (1001). ne,rn rrer (Brccinello V3 in Itaìv.rnd Sahabi in Lib1.r) and Erstern Alrica (Lothrganr in Kenva). \'item hor;elli n. sp. h,rs gcneral sizc closc to \/i1'errd Peprdtri (Earlv Pliocene of Europe). Gcner,rl proporrions lncl norphologv of los'cr carn:lssiil ho\vcver sugliest rn higher rlfinìrl,degrcc uirh r$'o small-sized viverrids, one (Viterra cltinjiensis) ironr l:rrger sized I-ate Miocene rnd Pleistoccnc spc'cics fronr Af- jì the Chin Frnr. (nriddlc Mìoce ne), and othcr (.Visbnuic- rìca (Viverra leakel, fronr Lanseeb:rnrve g rnd Omo V.rllc'v) ds rhe Paran (late Miocene salnortanus) in Dhok Fmr. thrn Europe (\'ii;ena PePt'tttri tt-td ,llegttiten a). to early Pliocene). Tivo large-sized forns (.Víz,erra bah- eri and \Iislnùctis durantli) *,cre,rlso recognized in rhis .lckxot- laigtnats. \\'.' rhrnks \lr: \!lc.rio rl.rlp.rssi tiu his .r, werc region, r'hcre thcv found in the Pinjor Fmt. (l.rtc siduous sunel in the (ìinigirno rrer,rnd ptof. I). Tonc fur ùe critìc,rl Pliocene) (Pilgrim, 1932). The genus l,'rslrarriclrs rvas cle- rc.rdìng of rhe rll.rnLrscript. Rescrrch supp()ned tlr' IfURST IllUllind fincd by Pilgrim (1932) on the basis oi thc cxtrenrc height thc Unirersirr oi l:lorcncc (Fondi di Arento) sr.ìnrs (l-R).

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