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
TLLrclirn, l\lcssìniu. lr.,ll Iiuropr. Airìc:r.

.lbvrrtt. \\e dercribc.r n<s rirerriJ.peties i\itctta botrlli n.
\p.r. I r::r,,rl,,i;c11t n. rp. i\ idcnritird in Lr(c ì\iioc.nc (Ilcssinirn) lo-
crlitics ìn tho circun Mc.lirtrrrne.tn.trcr (lt,rìr rnd Libr,t) rnd in ll.tsr
.\iri..ì íK.nv.rl. liorpho[,:ìc.rlhr thc nes specìcs is chrr.rcrcrized Lrr'.r
rcl.rri. rl! sn,rll \izc .ìnd r lo$ er c.ìrr.ì\sirl s irh shorr t.rl,,nrci.

Àr,,.j//rr., \rlcrr (l.5.rirt,r Lrn.r nuovr specìe divirerridc (\'ircrrl
/-,o:.r'11in.sp..\irt,rrl,o;r11in.sp.ùpresenrrinloc.rlìrìdel \liocenc
tlrnrinrlt {\lL'rini.rnor c,'n unr ili.tribuzn'nc nclì'.rre,r circLrm-l\lctlì-
r.ff.rnca (I!lli,ì. Libir) r,ì in Esr Atri.r (Kenri). \'lorloIrqicrnrent( si
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
t!f,r{, ;rit, i(,f( .on un r.rllonidc rri<'rro.

Introduction

Viierrids rre scrnrilv rcples('nred jn the Mio-
I)liocenc L-uropearr fossiÌ record. Thc earliest knorvn
l.uropean lcpreseDtJtives of rhc sub-lamìlv Vir.errinire
clirte to the bcginnint of thc lVlioccnc, n'hen madc irs
first ,rppc.rr'.rnce So)/gereff.. (Hclbing 192l, 1928; Ensess-
el 1972), i gcnus that survì\'ed untiì the Virllcsian (Pet-
tef 1976; ùlonto\',r ct al. 2001). The earliesr rcpreseDra-
tiles of the genus \.ii.,,-ra occur in thc dcposits of Vicr.rx
Colìongcs (Fr:uce; unit llN.{ in the Europcirn nrammll
bicrchrorolor:r') there are clcscribed as \titerra nodic,t
!ctrlrt, (Virct l95l; Dc Be:runront 1973). This species
is irls,l prcscnt rr irnorhcr Flerrch sirc, at Lr Crive (MN
7/3). rvith .r differe'nt subspccies, n 1\el\,\'ire11'd t tó.lica

t od i ca (yier 1 95 1 ; cìe Be.runront 1 973). Durine Turoli:rn

r ilerrid :rre ven' elusilc in Èurope. A viterrid nrandible
hirs been dcscribed from thc ltrlian,rssemblagc Baccincllo
V3 (Larest Mìocenci MN 13) as )'iterra sp. bv Rook et
il. ( l99l). Thcsc,rurhors,.rlthough recognize rhis fornì
as a possible nerv specics, prefcrrecl not to erect a ne\\r
niìnle for it. MateÌialattributed to a unnamed ncw yi?er,?

spccics hls bcen reco..e.ed irom,rnothc. htest Miocenc
(Mcssinian) locality in the circum-Medjterr:rnean irea,
tlrc locrlitY P2SA of Sahabi, in Libvr (Vico1z n. sp. "A'
in Hoq.cll 1987). A largcr species oi \,'rz'enz is found in
lhc Lower Pìioccrrc (Ruscinian) of sorLthern Francc, rìt
J\'lonrpellicr and Roussillon (MN l.l and MN l5 respec-
tivcllJ, rvhcre \titerra pepratxi rs prcsent (Dcperet 1390).
T)rc same spccies ìs also recorded at "Fornrcc RDB quar-
rr'" (Itah; Larlv Villrrfranchirn, MN l6:r) (Azzaroli er al.
1988). Dr.rring Earìv Pliocenc anotlter rchted genùs is
knon'n to be disrrjbuted in Europe: Megavi."'erra (Krer-
zoi & Feifar 1982). This gcnus hlls been erecred on thc
bLrsis ot nrrterial from lvanovcc 1 (Slovakia; MN 15) ar-
trìbuted to the species l,legz,i;icerra cdrpatorun. The g,e-
nus is also recorded in the Earll Villafranchi:rn of Itrh.
('lriversa, MN 1(ra), *-ith.r cliffercnr species, Megaviaerra
dpcnniDicd (Kretzoi & Fejfrr 1982).

The Baccinello-Cin igiirno brsin (soutì.rern Tus-
cinl', Italy) is knorvn for its Lete Miocene vertebrate
record (Bcnvenuti et al. 20Ol). Latc Miocene (MN l3)
n'ìanlmal remains arc founcl in several sites, from out-
crops of the upper part of tl.re fluvio-l'rcusrrine scdi-
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

1cm

b

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.

porar\'liYerrid from Sahabi (Vicerra n. sp. "A' in Howell
1987) and from Lothagam (Viverrinae sp. ìnder.; \lerdelin
2003) rre re-describcd and attributed to a new species.

Systematic Paleontology

Order Carnivora Boq'dich, 1821

Suborder Ailuroidea Floweq 1869

Famiìv Viverridae Gral', 1821

Subfamil"- Viverrinae Gill, 1872

Gews Vioerra Linné, 1758

Vlverra howelli n. sp.
Iiig l

1987 \'i!,e//, n. sp. 'A , Honell, Fig. 2r
l99l - \'iteza n. sp., Rook et al., Pl. l, iigs.2-3
2oo3 - Viverrinrc sp. indet., Werdelin, Iig. 7.13

Holotype. IGF 2390v (Fig. 1), right mandibulir rrmus bearing
loser c.rnine (;solrred), rr;ors of P,. P.,l' .rnd f, (fourth prcmolat still
presen ing a smrll talonid portion), conrplete iirsr los er molar (c.rrnas
sial),.rnd the rlveolus of M..

Repositon. N.rrural Hìstorr Museum, The Universitv of Flor-
cnce, Irrrll The specimen is kept ìn the collections of the "Ceologv rnd
l'aleontologv Secrìon" s,irh inlentory number "lGF 2890v".

Type locality. C:rprarccce, 100 nr So oi Pogg;o Vrnnini , Bac-
cinello-Cinieiano basin (Crosseto. southern Tusc:ny, Itrly).

Chronologicd distribution. Mcssinirn, lrte Turolian, MN l3
unir in the Europern nrmmal biochronolosy.

blage s,as named V3 (after Lorenz 1968) to distinguish
ìt from older mammrl faunas (knorvn as V0, Vl, and V2)
rhar are present in the sedimentarl, sequence of the b:r-
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-
ii), suggesting that the Tlsco-Sardinian region was an
insular domain characterised by a faunlJ province dis-
tìnct from the provinces of the surrounding coDtinental
masses. This isolation of the Tusco-Sardinian paleobio-
province came to an cnd with the Messinian, when the
region was reached by fornrs rypical of the European
faunal bioprovince (Hurzeler & Engesscr 1976; Rook
l99l; Bernor et al. 2001). The Baccinello - Cinigiano
succession was studied in detail by Lorenz (1968) and,
more reccnrly' bv Benvenuti er al. (1995, 2001) ;rnd
Rook et al. (2000).

The fossiliferous sire of Caprarecce was first discov-
ered by Mr. Malpassi, and a certain number of sp€cinìens
were collectcd either by Mr. Malpassi and b-u-. rescarchcrs
of the Earth Sciences Dept. of the University of Florence.
Some of these sanpÌes have been already describcd (Rook
er al. l99l; Rook Ei Rusrioni 1991; Rook & Torre 1995;
Abbazzi 2001) and rhe folloving taxa havc been at present
recognized in the assemblage for this localitl':

Tapirus c[. araemensis Croizet Er Jobert 1828, Pro-
caPreolus cf. lócryti Pohlig l9l l, Cervidae indet., Kolry2
ocboerus paleochoelrr (Kaup 1833), Vioerra sP., Celaden
sia grossetand Rook & Torrc 1995, Alilepus sp., Castor cl.
praefiber Déperet 189A.

The r.iverrid mandible from this site has been de- Diagnosis. Mindibular ramus f.rirll sturdv rvirh loser profilc
s cribed as Vloerra sp. by Rook er al. (1991 ) . The specimen slightlv .rrched. I-ower c:rrn:rssial stoudy built liigonid makes up rbout
was recognized as b;longing ro a hitherro undescribed spe- r""-thirJs of thc tooth Metrconid n'ell developed' almosr of the samc

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.



'12 00

10.00

800

2.00

!
È

300 '000 1200 .00 600 300 _000
lowsr M1 Le.qtb

l0 00

É
3

4.00

240

2,00 4,00 6.00 Loo ro oo
low€r M1 Talonid Lenglh

000 

-.

Viverra howelli z. sp., a ne.- Latest M;ocene ritetrid 721

Fig2 - Bivarirte plot diagrrm ofM Length vcrsus M, \ridrh (rbovt
and M, Taìonìd Lengrh versus M, Width (belod. Legend:
Vìl:erra nodica, solid circLcs; VìaeÍ't hateUì n. sp.j open
s+ares; Vhetd peptut\1. solid rrìangler yrrerra |eabe1, open
diattlonds; ùeSartrelr, spp., crosscs.

EtvmoÌogy. Thc species ìs namcd afier prof. Clark F. Houell,
for hìs contrìbutìon in the srudy of Mio-Pliocene carnivores rnd for
his r eference in authors' profession:rl c:rrier

Measuremcnts. See Table 1.

Description and comparisons. Vioera hotoelli n. sp.
is morphologically characterized by a quite robusr lower
canine, with a root and crown circular in section. The loe/-
er canine crown has v.eil developed longitudinal furrows,
both on the external and the internal sides. The premolars
are represented only bv their roots in the alveoÌi and small
crown fragmenrs (P., P. and P,). The first lower premolar is
very small in size and its alveolus is located just behind the
posterior malgìn ofthe canine alveolus (with no diastema).
A small diastema occurs berween P, and rhe anterior alveo-
lus o{ the double roored second lower premolar As far as
can be seen, second and third lower premolars are narrow
and slender. The small talonìd fragment preserved seenrs ro
indicate that the fourrh lower premolars q.as cnlarged pos-
teriorly, aìthough the occurrence of accessory cùsps cannot
be determined. The first lower molar (the carnassial) has a
paraconid intermediate in height between protoconid and

metaconid. The M,talonid is relativeÌy short, and charac-
terized by the occurrence of accessory cusplets, either on
the buccal or on rhe lingual side. The second lower molar
is not preserved bur its alveolus indicates rhe occurrence of
an elongated roorh. The mandibular ramus is slender; the
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
level of the second Ìower molar alveolus (Fig. 1).

Viverra hotpelli n. sp. is clearly distìnguished from
thc genus À4egaaicera by irs smaller dimensions (Fig. 2)
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
a right M, and described as Viz,erra n. sp. "A' by Howell
(1987), is fully comparable by size (Fig. 2) with Vioera
bou.'elli n. sp.. In addition, also the fragmentary mandible
from the Lower Nawata Fmt. (Lothagam) recently de-
scribed as Viverrinae sp. indet. (\ll'erdelin 2003) is morpho-
logically comparabl e wrthVit..ena bowel[ n. sp.. The Loth-
agam viverrinae sp. indet. is just larger in carnassial length
in respect to the BaccinelÌo V3 type specìmen and Sahabi
M but shows the same proportions in talonid reduction
(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
spccimens have a relatively short M, talonid in respecr ro V
pepratxi and to the larer (and larger) European Megaùver-
ra. (Fig.2). The Baccinello V3 specimen shows instead a
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
here considered conspecific wìrh Vixetra houelli n. sp.

Discussion. As already poìnted out, the fossil
record of the genus Villera is discontinuous in Europe,
since ìts first occurrence in the early Miocene. Vicer-
ra hoteelli n. sp. represent a latesr Miocene (Messinian)
small sized viverrid with a geographic distriburion from
circum-medìterranean area and Eastern Africa, identi
lied at Baccinello V3 (Itaìy), Sahabi (Libya) and Loth-
agam (Kenya).

Durìng latest Miocene in Africa the genus Viveta
was present, in addition to Sahabi and Lothagam, with a
Jarger species (Viz,ena leabey'1 at Langebaanweg (Messìn-
ian or early Pliocene; Sourh Africa; Hendey 1974, 1981).
Vivena leahey has a longer chronological distribution, be-
ing documented untiÌ the early Pleistocene of the Omo
valley, in the Tanganika area (Petter 1963; HoweÌl 1987).

The first Asian representative of this subfamily ap-
peared during the Astaracian (middle Miocene) ìn Mon-
golia (Tung Gur Fm.). Its extremely primitive character-
istics result in ìts assignment to a separate genus, Tungu-
rucrì (Colbert, 1939). The Siwaliks HiÌls have produced



specès Lenorh wdrh Ia on d Loca 
'ly

Lèn9rh

'+,-oÉr. " ". : : é.0òoF.
560 330 25o LeGtue

Viv.hahaúellt 10.40 530 400 Baccn€ ov3(CapÈrecce)

v)vena peptal, 1250 6 50 600 Rouss on

Vtveúleakey 15.50 730 -610 La^gebaanweg

Vil.ia tèakay 1600 760 Langcbaaiwèg

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

Mèa.vN,èra apènnrnrca 1630 360

722 L. I xtk E; B. .llarthrcz-N,tiarro

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
fcifrr (l9Sl). \\irdeliD (1001).

r$'o small-sized viverrids, one (Viterra cltinjiensis) ironr
the Chin jì Frnr. (nriddlc Mìoce ne), and othcr (.Visbnuic-
ds salnortanus) in rhe Dhok Paran Fmr. (late Miocene
to early Pliocene). Tivo large-sized forns (.Víz,erra bah-
eri and \Iislnùctis durantli) *,cre,rlso recognized in rhis
region, r'hcre thcv werc found in the Pinjor Fmt. (l.rtc
Pliocene) (Pilgrim, 1932). The genus l,'rslrarriclrs rvas cle-
fincd by Pilgrim (1932) on the basis oi thc cxtrenrc height

and rìarrowncss of the skull, chirrircteristics he fecls en-
tirelv preclude its assignrìrent ro ilr'ì\.cxisting vilerrid. Pil-
grinr (1912) ;rlso dcscrìbecl 

'r 
frrsnlent.rrl right mandibu-

larbrrnch bear-ing l.-lv{ (l:rbelled GSI D 135) .rs "LLrtrinee
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
better rrtributable ro 

'r 
vir,clrid .rnd probabll, lcprcsenrs

a nerr' llrge-sizcd gcnus.
\/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.

Conclusive remarks

Thc nervll describecì r'iler-rine spccies \/it'en'a houeLli
hld :r llrest Mioccnc distribution from circunr-Meditcrr.r-
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
l:rrger sized I-ate Miocene rnd Pleistoccnc spc'cics fronr Af-
rìca (Viverra leakel, fronr Lanseeb:rnrve g rnd Omo V.rllc'v)
thrn Europe (\'ii;ena PePt'tttri tt-td ,llegttiten a).

.lckxot- laigtnats. \\'.' rhrnks \lr: \!lc.rio rl.rlp.rssi tiu his .r,
siduous sunel in the (ìinigirno rrer,rnd ptof. I). Tonc fur ùe critìc,rl
rc.rdìng of rhe rll.rnLrscript. Rescrrch supp()ned tlr' IfURST IllUllind
thc Unirersirr oi l:lorcncc (Fondi di Arento) sr.ìnrs (l-R).

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