NEW DATA ON THE VERTEBRATE ASSEMBLAGE OF

FIUME SANTO (NORTH-WEST SARDINIA, ITALY),

AND OVERVIEW ON THE LATE MIOCENE

TUSCO-SARDINIAN PALAEOBIOPROVINCE

by LAURA ABBAZZI*� , MASSIMO DELFINO* , GIANNI GALLAI* ,

LUCIANO TREBINI� and LORENZO ROOK**Dipartimento di Scienze della Terra, Universita degli studi di Firenze, Via G. La Pira, 4, 50121, Firenze, Italy; e-mails: laura.abbazzi@unifi.it,

massimo.delfino@unifi.it, gianni.gallai@unifi.it, lorenzo.rook@unifi.it (corresponding author)

�Museo di Storia Naturale (Sezione di Geologia e Paleontologia), Via G. La Pira, 4, 50121, Firenze, Italy

�Soprintendenza per i Beni Archeologici per le Province di Sassari e Nuoro, Piazza S. Agostino, 3, 07100 Sassari, Italy

Typescript received 11 May 2006; accepted in revised form 20 March 2007

Abstract: This paper reports the results of a study of more

than 2400 specimens in the collection of fossil vertebrates

from Fiume Santo in north-west Sardinia. This locality rep-

resents the westernmost documentation of the Tusco-Sardi-

nian palaeobioprovince, which was in existence during the

late Tortonian in the North Tyrrhenian area. During the

Tortonian, the region that presently corresponds to southern

Tuscany and the Sardinian Massif (Central Italy) was occu-

pied by a complex of large islands characterised by endemic

vertebrate populations. Morphological features, adaptations

and trophic structure (e.g. low diversity and scarce carnivore

taxa), mainly in the mammal faunas, attest to a long period

of isolation from the continental palaeobioprovinces of the

Mediterranean and Central Europe. The greatest numbers of

fossil remains of the well-known endemic Oreopithecus fauna

of the so called V0–V2 assemblages, are found at several sites

in southern Tuscany (e.g. Montebamboli, Casteani, Bacci-

nello). In Sardinia this endemic fauna occurs only at the

Fiume Santo locality, from which 11 taxa have been recog-

nized: Crocodylia indet., Chelonii indet., Oreopithecus bam-

bolii, Mustelidae indet., Eumaiochoerus cf. E. etruscus,

Umbrotherium azzarolii gen. et sp. nov., Tyrrhenotragus grac-

illimus, Bovidae gen. et sp. indet. (?Neotragini), Maremmia

cf. M. lorenzi, Etruria viallii gen. et sp. nov., Turritragus cas-

teanensis gen. et sp. nov. Analysis of these fossils and com-

parison with material from localities in Tuscany has led to a

re-evaluation of the latter and to the description of three

new endemic taxa among the ruminants.

Key words: herpetofauna, mammal faunas, late Miocene,

Tortonian, Tusco-Sardinian palaeobioprovince, Italy, evolu-

tion, insular endemism.

The Fiume Santo site in north-west Sardinia (Sassari

Province; Text-fig. 1) yields a rich assemblage of endemic

fossil vertebrates. The fauna includes Oreopithecus, a

highly derived extinct ape, and the bovids Maremmia and

Tyrrhenotragus. The occurrence of these taxa characterises

the faunal assemblages V0–V2 of the well-known Torto-

nian faunas (Turolian Mammal Age, Mammalian Neo-

gene (MN) zones 11–12, ‘mammal zone’; Text-fig. 2) of

the Maremma region in southern Tuscany (Hurzeler and

Engesser 1976), where the most complete faunal succes-

sion crops out in the Baccinello-Cinigiano Basin (Lorenz

1968; Engesser 1989; Rook et al. 1996, 1999a; Bernor

et al. 2001). The Sardinian site represents the westernmost

documentation of the extinct insular fauna of the so-

called Tusco-Sardinian palaeobioprovince (Oreopithecus

faunal assemblages, or ‘OZF’ Oreopithecus Zone faunas, as

defined in Bernor et al. 2001). This province was in exis-

tence until the latest Tortonian in the North Tyrrhenian

region, when this insular region began to be connected to

the European mainland.

The Fiume Santo site was discovered in the early 1990s

(Cordy and Ginesu 1994; Cordy et al. 1995) during deep

excavations carried out for the construction of a parking

area within the thermo-electric power station of the Ital-

ian State Electricity Company (now ENDESA Italia). The

site has been declared protected by the local office of the

Italian Ministry for Cultural Heritage and Archaeology,

the ‘Soprintendenza per i Beni Archeologici per le prov-

ince di Sassari e Nuoro’. The fossiliferous area is relatively

large and within the industrial complex of the electricity

company. In September 2001, the ‘Soprintendenza per i

Beni Archeologici per le province di Sassari e Nuoro’ put

[Palaeontology, Vol. 51, Part 2, 2008, pp. 425–451]

ª The Palaeontological Association doi: 10.1111/j.1475-4983.2008.00758.x 425

the University of Florence in charge of the excavation and

study of the site and its fauna. The present contribution

presents the first results of an analytical study of the

1994–95 fossil sampling, a small portion of the entire

Fiume Santo assemblage, which currently consists of more

than 13,000 specimens.

At present, the geological context of the site is not sat-

isfactorily known (Cordy et al. 1995). An extensive geo-

logical survey is being carried out (Benvenuti and Papini

in progress) within the framework of the project.

Institutional abbreviations. Bac, Baccinello collection housed in

the Naturhistorisches Museum, Basel; FS, field inventory register

of Fiume Santo material; IGF, Natural History Museum, Geol-

ogy and Palaeontology section, University of Florence; IR, formal

inventory register of Fiume Santo material according to the ‘In-

ventario Regionale della Soprintendenza’.

SYSTEMATIC PALAEONTOLOGY

The faunal assemblage

The revision of the Fiume Santo fossil assemblage, the

object of the present paper, has been based on the

remains collected during the first excavations (1994–95),

as noted above. It was carried out under the direction of

one of us (LT) and stems from a study of more than

2400 specimens (of which at least one-quarter have not

yet been identified), representing 11 vertebrate taxa: Croc-

odylia indet., Chelonii indet., Oreopithecus bambolii,

Mustelidae indet., Eumaiochoerus cf. E. etruscus, Umbro-

therium azzarolii gen. et sp. nov. (nomen nudum in Hurz-

eler and Engesser 1976), Tyrrhenotragus gracillimus,

?Neotragini gen. et sp. indet., Maremmia cf. M. lorenzi,

Etruria viallii gen. et sp. nov. (nomen nudum in Hurzeler

and Engesser 1976), and Turritragus casteanensis gen. et

sp. nov. (for a complete list of the material analysed

herein, see Supplementary Data file on the website of

the Palaeontological Association: http://www.palass.org).

The available material did not allow us to identify some

of the taxa listed by Cordy et al. (1995), which includes

two rodents (the murid Valerymys aff. V. turoliensis and a

large Gliridae indet.). No micro-mammal remains were

recovered from the material at our disposal. The same

holds true for carnivores. Cordy et al. (1995) mentioned

the occurrence of the endemic Ursidae ‘Hyaenictis’

anthracitis, but our study revealed the occurrence of

only two fragmentary remains attributable to the

Mustelidae.

Although the fossil assemblage analysed represents only

a small part of the whole Fiume Santo collection, our tax-

onomic study and general conclusions are significant

enough to be presented here and will provide a basis for

future palaeontological investigations at the site.

The fossils are not often well preserved. The enamel of

teeth is perfectly preserved but dentine and roots are

often chemically eroded or sometimes absent altogether.

Bones generally appear ‘decorticated’ with the innermost

part of cortical bone or even the trabecular bone usually

being exposed (Text-fig. 3). These states of preservation

suggest that the teeth and bones were in highly alkaline

environments both before and after burial, which strongly

altered the most organic components of skeletal elements

(Fernandez-Jalvo et al. 2002).

The identifications of mammals are based mainly on

tooth morphology and, in the case of bovid remains, on

Ser

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TEXT -F IG . 2 . Geochronological framework.

Fiume Santo

Sassari

Roma

0 200 km

N

Oristano

Cagliari

Nuoro

TEXT -F IG . 1 . Map showing the location of Fiume Santo site.

426 P A L A E O N T O L O G Y , V O L U M E 5 1

horns. Postcranials are mostly badly fragmented, prevent-

ing their certain identification. A first estimate of the per-

centage occurrences of each taxon is provided in Table 1

and Text-figure 4. In Text-figure 4A the percentages have

taken into account the entire set of identified fossils

(horns, teeth and bones of mammals, teeth of crocodil-

ians and shell fragments of chelonians). In Text-figure 4B

only mammalian dentition and isolated teeth have been

considered in order to avoid over-representation of the

most common and easily identifiable taxon Maremmia.

From these diagrams it is apparent that bovids are the

best documented group, representing more than 75 per

cent of the whole sample of mammal dentitions, whereas

carnivores and suids are represented by only 2–4 frag-

ments per taxon.

The taxonomic composition and diversity of the Fiume

Santo fauna is fully comparable with that of southern

Tuscany (Baccinello V0–V2, Montebamboli, Casteani, Ri-

bolla, Montemassi, Serrazzano), and all occurrences of

mammal taxa at Fiume Santo are the only records of their

presence outside Tuscany (see data in Rook et al. 2000).

Although both faunas are characterised by the dominance

of the bovid Maremmia, some differences in the relative

abundance of taxa are evident: for example, the giraffid

Umbrotherium azzarolii is the second most abundant

form at Fiume Santo, whereas in southern Tuscany it is

represented by only one specimen (the type specimen

from Casteani). Another difference concerns the charac-

teristics of the dentition of Maremmia, which suggest dif-

ferent ecological conditions between the regions (see

discussion of bovid taxonomy below).

Class REPTILIA Laurenti, 1768

Remarks. The herpetofauna is represented by remains of

crocodylians and chelonians. No amphibians or small

reptiles have been identified in the 1994–95 samples,

although Cordy et al. (1995) listed four unidentified

forms: one chelonian, one anuran and two crocodylians.

B

A

C

TEXT -F IG . 3 . A, example of a fossiliferous assemblage from Fiume Santo; in the foreground is a right mandible of Umbrotherium

azzarolii; · 0.5. B, Maremmia cf. M. lorenzi M3 with absent dentine; · 2. C, Maremmia cf. M. lorenzi humerus; arrow indicates an area

where trabecular bone is exposed; · 0.5.

A B B A Z Z I E T A L . : M I O C E N E V E R T E B R A T E A S S E M B L A G E F R O M S A R D I N I A 427

Order CROCODYLIA Gmelin 1789

Crocodylia indet.

Text-figure 5A–B

Description. The crocodylians are represented exclusively by iso-

lated teeth (49 recovered) of medium size (crown height c. 7–

33 mm) and rather massive form (Text-fig. 5A–B). Differences in

elongation and size indicate that they come from different parts of

the teeth-bearing bones. The teeth are variably fractured and

incomplete. The enamel is usually translucent and when the surface

is well preserved, it is usually characterized by fine longitudinal

ridges and irregularly scattered small pustules that may anastomose

in the distal sector where they diverge in laterally, reaching the

mesiodistal keels (without producing denticles or serrations). In

specimens having a transverse constriction on the crown, the ridges

are weakly expressed and the small pustules more in evidence.

Comparisons and discussion. Despite the poor phyloge-

netic value of isolated teeth (Brochu 2000), some taxo-

nomic relevance cannot be denied in particular contexts

of known age and geographic provenance, as in the case

of the Fiume Santo assemblage. The crocodylians of the

late Miocene Mediterranean islands are represented exclu-

sively by members of the family Crocodylidae since the

range of the alligatoroid Diplocynodon was restricted to

the European mainland, possibly because some sort of

intolerance to salt water limited its dispersal (see discus-

sion in Delfino in press). The isolated teeth from Fiume

Santo do not belong to this genus, whose Miocene repre-

sentatives have teeth that are more acute with smaller

crowns and are usually not ornamented by any ridges. It

is, therefore, possible to doubt the presence of this genus

at Fiume Santo. Cordy et al. (1995) listed two crocodylian

taxa from this locality but the available isolated teeth sug-

gest derivation from different parts of the teeth-bearing

bones of a single taxon. According to a recent review of

the Italian Cenozoic crocodylians (Kotsakis et al. 2004),

Sardinia was inhabited by at least one long-snouted croco-

dylid of the Tomistominae during the late Miocene. Short-

snouted crocodylids surely inhabited southern Tuscany

(‘Crocodylus bambolii’ Ristori, 1890 is a crocodylid of

uncertain affinity; Delfino and Rook 2008), and possibly

the Apulo-Abruzzi Province, where non-Diplocynodon-like

isolated teeth have been found at the locality of Scontrone

(Rustioni et al. 1992), whereas Crocodylus cranial remains

have been identified recently from the Gargano palaeo-

archipelago (Delfino et al. 2007). Well-grounded evidence

for Crocodylus in Sardinia is not yet available, even if the

morphology of the isolated teeth reported here does not

differ from that of a member of this genus. Further

remains (cranial elements) are needed to support such a

determination; hence the material from Fiume Santo is at

present conservatively identified only at order level.

Order CHELONII Brongniart, 1800

Chelonii indet.

Text-figure 5C–D

Description. Chelonians are represented by a few small carapace

or plastron fragments. The most interesting specimen is frag-

ment FS1995#0579, which is tentatively considered to be the

external portion of a right xiphiplastron (Text-fig. 5C–D).

Sutures are not preserved but a transverse ‘step’ on the ventral

(external) surface may represent the boundary between the fem-

oral and anal shields. Such a boundary is not visible on the dor-

sal surface of the element (in the lateral area covered by the

shield), probably because of inadequate preservation. The surface

immediately caudal to the ‘step’ is evidently slightly depressed in

a way similar to that seen in modern Testudo graeca from Tur-

key (in the female the depression is also developed anterior to

the furrow between the femoral and anal shields). Unlike the

shape of shells of modern Mediterranean testudinids, the pos-

TABLE 1 . Absolute numbers and proportional distribution of

vertebrate taxa at Fiume Santo: A, total remains; B, mammalian

dental elements only.

Taxa No. %

A

Indet. 661 26.72

Chelonii indet. 22 0.89

Crocodylia indet. 49 1.98

Mustelide indet. 2 0.081

O. bambolii 25 1.01

E. cf. etruscus 4 0.16

Bovidae indet. 429 17.33

T. gracillimus 45 1.77

?Neotragini indet. 36 1.45

M. cf. lorenzi 853 34.46

Etruria viallii 35 1.41

T. casteanensis 14 0.56

U. azzarolii 301 12.17

Total 2475

B

Indet. 86 5.03

Mustelide indet. 2 0.12

O. bambolii 25 1.46

E. cf. etruscus 4 0.23

Bovidae indet. 352 20.62

T. gracillimus 44 2.57

?Neotragini indet. 33 1.93

M. cf. lorenzi 824 48.21

Etruria viallii 35 2.05

T. casteanensis 14 0.82

U. azzarolii 290 16.97

Total 1709

Bovidae indet. includes fragments (e.g. of enamel walls and

bones) that cannot be referred with certainty to any of the small

to middle-sized species.

428 P A L A E O N T O L O G Y , V O L U M E 5 1

terolateral edge of the xiphiplastron is not approximately

rounded but straight. The maximum thickness of 12.5 mm is

reached at the level of the proximal (broken) tip.

Comparisons and discussion. The chelonian fragments

available are of little diagnostic value. Although their size

rules out the presence of marine turtles and the proximal

thickness of specimen FS1995#0579 could suggest a tor-

toise, all the remains are simply referred to Chelonii indet.

Class MAMMALIA Linnaeus, 1758

Order PRIMATES Linnaeus, 1758

Family HOMINIDAE Gray, 1825

Genus OREOPITHECUS Gervais, 1872

Type species. Oreopithecus bambolii Gervais, 1872, from Monte-

bamboli (Grosseto, Central Italy), late Tortonian, late Miocene.

Oreopithecus bambolii Gervais, 1872

Plate 1; Table 2

Description

Twenty isolated teeth and one right fragmentary mandible bear-

ing P3–P4. No postcranial elements have been identified. Apart

from a short note on the fragmentary mandible IR#63906 by

Cordy and Ginesu (1994, Text-figs 3–4), none of these speci-

mens has been described previously. A selection of the material

is illustrated in Plate 1 and measurements of the specimens are

given in Table 2.

Upper dentition. Upper premolars are represented by one right

P3 (IR#63902) and one right P4 (IR#63904); they are typically

bicuspid, with subequally developed cusps (the buccal one being

slightly more robust). The P3 (IR#63902; Pl. 1, figs 1–4) is a vir-

tually unworn tooth; a strong cingulum encircles its base, and is

more developed on the lingual aspect. The P4 (IR#63904) is a

A

T. gracillimus

T. casteanensis

O. bambolii

?Neotragini indet.

Mustelide indet.

?Neotragini indet.

Mustelide indet.

M. cf. lorenzi

E. cf. etruscus

M. cf. lorenzi

total remains

Bovidae indet.

Chelonii indet.

Crocodylia indet.

Etruria viallii

E. cf. etruscus

Indet.

Indet.

dentitions

T. gracillimusU. azzarolii

Bovidae indet.

Etruria viallii

T. casteanensis

O. bambolii

U. azzarolii

B

TEXT -F IG . 4 . Proportional

occurrences of taxa from Fiume Santo

based on A, total remains and B,

mammalian dental remains.

A B B A Z Z I E T A L . : M I O C E N E V E R T E B R A T E A S S E M B L A G E F R O M S A R D I N I A 429

deeply worn tooth, with the two cusps worn down to be con-

nected distally around a dentine lake. A marked cingulum runs

on the lingual side. The first upper molar crowns are longer

than broad, with strongly rounded corners. Of the two M1s in

the sample, one is nearly unworn (FS1995#0005), while the other

(FS1995#0011; Pl. 1, figs 5–7) is in an advanced stage of wear,

having dentine exposure on three main cusps (protocone, para-

cone and hypocone). The protocone (the largest of the four

cusps) is relatively low and conical in shape. No metaconule is

present along the crista obliqua. Mesial to the protocone there is

a cingular shelf, which ends on the mesial face of the paracone.

The mesial fovea is variable, being easily identifiable in

FS1995#0011 and very small in FS1995#0005. In both M1s, distal

to the metacone there is a relatively broad distal shelf, which in

FS1995#0005 bears a distinct distal conule. The second molars

are wider than the first. Of the four specimens, three are unworn

or slightly worn (IR#63903, FS1995#0001, FS1995#0002) while

one is at a relatively advanced stage of wear (IR#63900; Pl. 1,

figs 8–10), with all four main cusps exposing a dentine tip. The

mesial margin is characterized by the occurrence of a wide

mesial cingulum, delimiting a variably sized mesial fovea. The

metaconule is absent along the crista obliqua in all four M2s.

Distal to the hypocone and metacone is a robust, broad distal

shelf, which in IR#63903, FS1995#0001 and FS1995#0002 is

characterised by the presence of a well-defined distal conule. The

third upper molars are characterised by the narrowing of the

distal portion of the tooth. All four M3s present in the sample

are at a very low stage of wear. Two are complete (IR#63901,

FS1995#0010; Pl. 1, figs 11–13), while the other two

(FS1995#0006, FS1995#0012) are damaged in the distolabial part.

All are characterised by a wide, robust mesial cingulum running

mesial to the protocone and paracone, and by a distal shelf that,

starting at the disto-internal margin of the metacone, goes

around the hypocone base as far as the labial side of the tooth.

The distal shelf is characterised by the occurrence of small cus-

plets on its ridge. A peculiarity of IR#63901 is the reduction of

the hypocone that gives a ‘triangular’ shape to the tooth.

Lower dentition. The lower deciduous dentition of Orepithecus is

known thanks to the left D4 of the type specimen (Gervais 1872;

Hurzeler 1951). The right D4 from Fiume Santo (FS1995#0009;

Pl. 1, figs 14–16) is virtually identical in its dimensions and

morphology to the Monte Bamboli D4 illustrated and described

by Hurzeler (1951, 1958). In respect to the type D4 (which is

partially damaged with the hypoconid area missing), the Fiume

Santo specimen is complete and shows a slightly more advanced

stage of wear.

A crown fragment of a left I1 (FS1995#0013) is the only inci-

sor present in our collection. The specimen lacks its base (cer-

vix) and the buccal margin is more intact. The occlusal margin

is unworn and characterised by the occurrence of a larger (dis-

tal) and a smaller (mesial) mammelon. The margin ridges on

the lingual side are almost undetectable, the mesial one being

more marked.

Lower premolars are represented by a right fragmentary man-

dible with P3–P4 (IR#63906), a left isolated P3 (FS1995#0003)

and a left isolated P4 (IR#64428; Pl. 1, figs 17–19). Both P3s rep-

resent a bicuspid tooth, with a large protoconid and a smaller,

but prominent metaconid located transversely opposite the pro-

toconid. There is no evidence of wear produced by the honing

action of the upper canine. The lingual aspect of the metaconid

A B

C D

TEXT -F IG . 5 . Herpetofauna from Fiume Santo. A–B,

Crocodylia indet., isolated tooth IR#63925 in A, lingual and B,

mesial views; · 1. C–D, Chelonii indet., right xiphiplastron

FS1995#0579 in C, ventral (the dark area is concretion filling a

shallow depression) and D, dorsal views; · 0.8.

TABLE 2 . Measurements (in mm) of Oreopithecus bambolii

from Fiume Santo. L, total length; B, breadth; Ba, anterior

breadth; Ba, posterior breadth; *, inferred measurement on

slightly broken or damaged elements.

Taxa L B Ba Bp

IR#63902 P3 7.1 9.2

IR#63904 P4 6.8 9.2

FS1995#0005 M1 8.5 8.0 7.3

FS1995#0011 M1 8.7 8.6 8.5

FS1995#0012 M1 10.5

IR#63900 M2 9.4 9.3 9.1

IR#63903 M2 9.9 9.5 7.7

FS1995#0001 M2 8.9 8.6 7.9

FS1995#0002 M2 10.1 8.8 8.1

IR#63901 M3 9.7 9.2 7.5

FS1995#0006 M3 *11.4 10.4

FS1995#0010 M3 9.6 9.1 8.0

FS1995#0012 M3 10.5

FS1995#0009 D4 7.4 5.4 4.6

FS1995#0003 P3 5.8 7.3

IR#63906 P3 6.5 8.5

IR#63906 P4 7.0 7.7

IR#64428 P4 7.8

IR#63905 M1 9.4 7.7 7.2

IR#64427 M1 7.3

FS1995#0007 M2 10.5

FS1995#0004 M3 11.4 9.2 8.8

430 P A L A E O N T O L O G Y , V O L U M E 5 1

differs between IR#63906 and FS1995#0003, the latter one show-

ing a concave profile. The mesial fovea is made by an extension

of the lingual cingulum, which in Oreopithecus is expanded. It is

worth mentioning that P3 in Oreopithecus is extremely variable,

ranging from a relatively narrow, single-cusped sectorial tooth to

a broad, bicuspid molariform tooth (Rook et al. 1996). In this

respect the notation of Cordy and Ginesu (1994), which stresses

the differences between IR#63906 and lower premolars from the

Tuscan samples, especially for the P3, seems inappropriate.

Regarding the P4s (IR#63906 and IR#64428), both are at a

comparable stage of wear (exposure of dentine tip on metaco-

nid) and well preserved, although in IR#64428 a flake of

enamel is missing on the distal wall of the paraconid. As for

the P3s, the tooth is broad and elliptical in shape. The talonid

basin forms a broad, but shallow triangular platform distal to

the main cusps. The distal marginal ridge bears cusplets along

the crest. In both, P4s have a continuous cingulum (stronger

in IR#63906) that encircles the buccal aspect of the protoconid.

In common with the upper molars, the lower molars are char-

acterized by four main cusps alternately arranged. Two M1s are

present in the sample, one complete (IR#63905; Pl. 1, figs 20–

22) and one represented by the distal part of the trigonid

(IR#64427), both at a similar stage of wear, with dentine

exposed on tips of the main cusps. The protoconid and metaco-

nid are large and transversely aligned. A centroconid is situated

in the midline of the tooth. The distal shelf is wide and runs

around the metaconid forming a lingual cingulum. The only M3

(FS1995#0004; Pl. 1, figs 23–25) consists of the crown of a well-

preserved, unworn tooth. It is relatively long but not as in other

Oreopithecus specimens. The protoconid and metaconid are the

larger and most elevated cusps. These are transversely aligned, a

condition typical for Oreopithecus. From the apex of the pro-

toconid, a short preprotocristid connects it to a relatively small

protoconid. A centroconid is clearly positioned in the midline of

the tooth. The talonid area is unusual because a distal fovea is

not clearly delimitated. The hypoconid and entoconid are rela-

tively large. Such a talonid morphotype falls within the range of

the observed morphology of Oreopithecus.

The mandibular ramus fragment IR#63906 is poorly preserved

with no inner or outer surface of the bone so that the thickness

of the ramus cannot be estimated; the only feature that can be

seen is the height of the ramus below the premolars (23.5 mm

below P4), which is within the range observed in the Tuscan

sample (contra Cordy and Ginesu 1994).

Comparisons and discussion. Oreopithecus was a large-

bodied hominoid with some features that are typical of

living apes, reflecting significant adaptations to vertical

climbing (Harrison 1991), but also other characters in

several parts of its skeleton that are more likely to be

linked to bipedality (Kohler and Moya-Sola 1997, 2003;

Moya-Sola et al. 1999, 2005; Rook et al. 1999b, 2004).

The site of Fiume Santo, with its enormous record of

undistorted bone remains, has the potential to yield un-

distorted post-cranial remains of Oreopithecus, which will

eventually provide a critical contribution to the contin-

uing debate about its locomotor behaviour (Moya-Sola

et al. 2005; Rook et al. 2004). At present, it is repre-

sented at Fiume Santo only by dental remains and the

sample available for this study shows that no appreciable

differences exist between it and that from southern Tus-

cany in either morphology or dimensions (Text-fig. 6).

A more comprehensive study, including the material col-

lected during site exploitation from 1996 until the pres-

ent, will allow for more extensive comparisons and a

better definition of the Fiume Santo Oreopithecus in due

course.

Order CARNIVORA Bowdich, 1821

Remarks. Predators are usually absent from insular mam-

malian assemblages or limited to forms with peculiar spe-

cializations in behavioural ⁄ feeding strategies (MacArthur

and Wilson 1963; Sondaar 1977, 1987). The mammal

assemblage of the so-called Oreopithecus faunal assem-

blages (OZF sensu Bernor et al. 2001) is no exception in

this regard. Those in southern Tuscany have yielded some

carnivores, all represented by very limited remains. An

isolated mandible from Montebamboli represents a bear

(Indarctos anthracitis) showing unusual dental adaptation

to an omnivorous diet (Weithofer 1888), and a maxillary

fragment represents the mustelid Mustela majori (Ficcar-

elli and Torre 1967). In addition, and better documented,

are the lutrine carnivores: these are known to be relatively

diverse, being represented by three species of two endemic

genera: Tyrrhenolutra helbingi (Baccinello V1), Paludolutra

maremmana (Montebamboli), and Paludolutra campanii

(Montebamboli, Baccinello V2) (Hurzeler 1987).

Among carnivores, Cordy et al. (1995) recorded only

the bear ‘Hyaenarctos’ anthracitis from Fiume Santo.

Within the material studied, however, identified carnivore

remains are limited to two very fragmentary specimens

that are not identifiable beyond family level.

Suborder CANIFORMIA Kretzoi, 1943

Family MUSTELIDAE Fischer de Waldheim, 1817

Mustelidae indet.

Description. Material attributable to Mustelidae indet. consists of

two small dental fragments, one representing part of the anterior

portion of possibly a left P4 paracone, the other a portion of the

talon (with protocone, hypocone and lingual cingulum) of a left

M1.

Comparisons and discussion. The fragmentary status and

the limited sample do not allow much scope for compari-

son and discussion. We are confident that analyses of

A B B A Z Z I E T A L . : M I O C E N E V E R T E B R A T E A S S E M B L A G E F R O M S A R D I N I A 431

post-1995 recoveries and new site excavations at Fiume

Santo will yield more evidence of the carnivores that cha-

racterise the faunal assemblage.

Order ARTIODACTYLA Owen, 1848

Family SUIDAE Gray, 1821

Genus EUMAIOCHOERUS Hurzeler, 1982

Type species. Eumaiochoerus etruscus (Michelotti), 1861; from

Montebamboli (Grosseto, Central Italy), late Tortonian, late

Miocene.

Eumaiochoerus cf. E. etruscus

Text-figure 7

Description. The Suidae are represented by only two complete

teeth: a right I3 and a right M3 (Text-fig. 7), the latter without a

root but the occlusal surface is well preserved, without wear and

with no eroded enamel. This specimen is subtriangular in shape

with four main cusps: paracone (anterobuccal), metacone (pos-

terobuccal), protocone (anterolingual), and hypocone (postero-

lingual). These are not opposed but alternating. The anterior

margin has a strong cingulum with a central cusp (protoprecon-

ule). There is an accessory cusp (tetrapreconule) positioned cen-

trally between the anterior and posterior pairs of the main

cusps. Two small additional cusplets are present labially at the

tetrapreconule. A second accessory cusp (pentapreconule) occurs

between the posterior pair of main cusps and the talon. This last

one is simple with a large central cusp divided by a small fur-

row, and three small pillars on the labial side.

Based on its dimensions (length, 31.53 mm; width,

20.77 mm) and morphology, this specimen can be referred to

Eumaiochoerus etruscus following the redescription by Mazza and

Rustioni (1997) of the Montebamboli specimens, the only signif-

icant difference being that the three grooves (‘furchen’) in the

Fiume Santo specimen, normally present on the main cusps of

Suidae (Hunermann 1968), are shallow in the paracone and

metacone and completely lost in the protocone and hypocone.

No conclusions are possible from the other fragmentary remains

from Fiume Santo.

Comparisons and discussion. Van der Made (1999) consid-

ered the simplification of molars with the loss of ‘furchen’

to be a result of increasing enamel thickness, a derived

feature occurring in suids under insular conditions. The

endemic Sus sondaari from the Plio-Pleistocene of Sardinia

lacks them. The fact that grooves are lacking in the Fiume

Santo suid, unlike that from Montebamboli-Baccinello V2,

suggests a more pronounced degree of endemic evolution

in the former. The limited sample does not allow much

scope for further comparison, so despite the close resem-

blance, we prefer to take a cautious approach and refer our

material to Eumaiochoerus cf. E. etruscus.

Family GIRAFFIDAE Gray, 1821

Genus UMBROTHERIUM gen. nov.

1976 Umbrotherium Hurzeler and Engesser, p. 334 (nomen

nudum)

Derivation of name. After one of the oldest peoples of Central

Italy (Umbri; First Millennium AD) and Greek, therion, wild

beast, a common reference to mammals.

Type and only species. Umbrotherium azzarolii sp. nov.

Diagnosis. Middle sized ruminant with fairly brachyodont

dentition. Markedly rugose enamel wall. P3 and P4 not

molarised, rectangular in outline, wider than long. Upper

molars with a weak entostyle and cingulum. Mesostyle

joins metastyle via a poorly developed cingulum; anterior

lobes longer than posterior lobes. Cranial features, in par-

ticular whether or not ossicones occur, currently

unknown.

Remarks. The remains originally referred to Umbrother-

ium by Hurzeler and Engesser (1976), and analogously to

Etruria viallii (see below), were not described, rendering

it a nomen nudum according to the ICZN (1999). In

order to make the name available for use, we provide an

extensive description and formal definition here.

Umbrotherium azzarolii Hurzeler and Engesser, 1976

was referred to the Giraffidae family by these authors.

This attribution is followed here although it is not firmly

grounded because the main diagnostic characters, such as

EXPLANATION OF PLATE 1

Figs 1–25. Oreopithecus bambolii from Fiume Santo. 1–4, IR#63902, right P3 in 1, distal, 2, occlusal, 3 proximal and 4, palatal views.

5–7, FS1995#0011, left M1 in 5, occlusal, 6, palatal and 7, buccal views. 8–10, IR#63900, left M2 in 8, occlusal, 9, palatal and 10,

buccal views. 11–13, FS1995#0010, right M3 in 11, occlusal, 12, palatal and 13, buccal views. 14–16, FS1995#0009, right D4 in 14,

occlusal, 15, lingual and 16, buccal views. 17–19, IR#64428, left P4 in 17, occlusal, 18, lingual and 19, buccal views. 20–22,

IR#63905, left M1 in 20, occlusal, 21, lingual and 22, buccal views. 23–25, FS1995#0004, right M3 in 23, occlusal, 24, lingual and

25, buccal views. All · 3.

432 P A L A E O N T O L O G Y , V O L U M E 5 1

PLATE 1

ABBAZZI et al., Oreopithecus

1 2 3 4

11

12765

89

10

13

23

1415 16

191817

20

21 2225

24

bilobate lower canines and ossicones, are not documented

from either southern Tuscany or Fiume Santo. Neverthe-

less, some features present in the dentition, such as the

markedly rugose enamel, highly molarised P4 and fusion

of enamel folds between cones ⁄ conids, which occurs in

very worn teeth, are typical of giraffids.

Umbrotherium azzarolii sp. nov.

Text-figure 8; Table 3

1888 Antilope (Palaeoryx?) sp. Weithofer, p. 365.

1889 Antilope (Palaeoryx?) sp. Weithofer, pp. 57, 62.

1912 Antilope (Palaeoryx?) sp. n. Del Campana, p. 212,

pl. 18.

1976 Umbrotherium azzarolii Hurzeler and Engesser,

p. 334 (nomen nudum).

Derivation of name. After Prof. Augusto Azzaroli, palaeontologist

and Emeritus Professor at the University of Florence.

Holotype. IGF14615, an upper left series with P3–M2, housed in

the Natural History Museum, Geology and Palaeontology sec-

tion, University of Florence (Text-fig. 8A–C).

Locality, horizon and age. Casteani (southern Tuscany), faunal

assemblage V1; late Miocene, late Tortonian, late Turolian.

Other referred material. Fiume Santo, north-western Sardinia;

see below.

Diagnosis. As for the genus.

Description

Upper dentition. There are no more features to report in addi-

tion to those already listed in the diagnosis apart from a strong

concavity in the lingual walls, which is more evident in less

worn teeth.

A

B

10.0

9.0

8.0

7.0

6.06.0 7.0 8.0 9.0

length (mm)

10.0 11.0 12.0

10.0

11.0

9.0

8.0

post

erio

r w

idth

(m

m)

post

erio

r w

idth

(m

m)

7.0

6.0

5.06.0 7.0 8.0 9.0

length (mm)

10.0 11.0 12.0

TEXT -F IG . 6 . Scatter diagrams comparing total length and

posterior breadth of A, M2 and B, M3 of Oreopithecus bambolii

samples from Fiume Santo (solid diamonds) and sites from

southern Tuscany (open diamonds).

A

B

C

TEXT -F IG . 7 . Eumaiochoerus cf. E. etruscus from Fiume Santo.

A–C, FS1995#0196, right M3 in A, labial, B, lingual and C,

occlusal views; · 2.

434 P A L A E O N T O L O G Y , V O L U M E 5 1

Lower dentition. Incisors show an asymmetric outline (e.g.

1995#0195). D4 has a protruding metasylid and two well-

developed interlobal columns. P2 simple, short and large, with

a triangular outline. P3 is not molarised and lacks a metaco-

nid, fusion between hypoconid and entoconid occurs late,

entoconid joins protoconid, paraconid and parastylid well

developed (Text-fig. 8). P4 is molarised, with metaconid in

the form of an anterior–posterior wall, which joins the parac-

onid and lacks the transverse connection to the protoconid.

Entoconid and hypoconid are two obliquely orientated crests.

A deep groove in labial side divides P4 into two unequal

parts, anterior and posterior lobes, with the posterior one

being strongly reduced. On lingual side, a less deep groove

occurs (Text-fig. 8). P4 and P3 are both characterised by

having a labially sharp hypoconid. Lingual walls of molars

obliquely orientated, in particular the anterior one. In M3 the

A

C

G

H

I

B

D E F

J

K

L

TEXT -F IG . 8 . Umbrotherium azzarolii gen. et sp. nov. from southern Tuscany and Fiume Santo. A–C, IGF 14615, holotype, left

maxillary fragment with M2–P3 from Casteani in A, occlusal, B, lingual and C, labial views; · 1. D–F, FS1995#0340, right P3 from

Fiume Santo in D, occlusal, E, lingual and F, labial views; · 1. G–I, FS1995#0342, left partial mandible with M3–P3 from Fiume Santo

in G, occlusal, H, lingual and I, labial views. J–L, FS1995#0343, right partial mandible with M3–P4 from Fiume Santo in J, occlusal, K,

lingual and L, labial views; · 0.5.

A B B A Z Z I E T A L . : M I O C E N E V E R T E B R A T E A S S E M B L A G E F R O M S A R D I N I A 435

second and third lobes are separated by a clear step; the hyp-

oconulid is high but narrow (Text-fig. 8). Ectostylids are

absent; a feeble cingulum occurs only on anterolabial side of

lower molars. The mandible is characterised by a fairly thick

horizontal ramus (Text-fig. 8).

Comparisons and discussion. A substantial part of mam-

malian dental remains at Fiume Santo belongs to this

puzzling form. In the preliminary notes of Cordy and

Ginesu (1994) and Cordy et al. (1995) the occurrence of

two giraffid species (differing in size) was reported. How-

ever, morphometric analysis of the material indicates the

presence of only one form (Table 3; Text-fig. 9); some

minor differences in size between dental elements may

reflect sexual dimorphism.

The advanced morphology of P4 is comparable to that

which characterized giraffid species of late Miocene age

referred to the subfamilies Giraffinae and Sivatheriinae,

while that of P3 is very peculiar. The molarised P3, with a

highly developed metaconid joined to the paraconid,

seems to be an apomorphy in the Giraffinae (as in Paleo-

tragus and Samotherium; Geraads 1978, 1979, 1986), while

it is not recorded in Sivatherinae. Umbrotherium has a

non-molarised lower third premolar (Text-fig. 8) lacking

a metaconid; this could preclude a link to Paleotraginae,

because it is untenable in the context of an evolutionary

simplification process in the Tusco-Sardinian palaeobio-

province. On the other hand, a primitive P3 is present in

Decennatherium pachecoi (Crusafont Pairo 1952; Morales

and Soria 1981; pers. obs.) from the Vallesian site Los

Valles de Fuentiduena (MN9) in Spain, and in Hellado-

therium from Greece.

Biometric comparison of Umbrotherium azzarolii shows

that it is comparable in size to the medium-sized giraff-

ids, such as the late Miocene Paleotragus (Text-fig. 9).

However, this does not help to clarify phyletic relation-

ships because the reduction in size may be a result of the

evolution of Umbrotherium within the insular domain of

the Tusco-Sardinian palaeobioprovince.

Family BOVIDAE Gray, 1821

Remarks. As reported above, the taxonomic diversity of

bovids from Fiume Santo is fully comparable with that of

assemblages V1 and V2 from continental Italy. Despite

the fact that bovids are well represented and diverse at

sites in southern Tuscany, only two genera have been

described in detail in the literature: Maremmia and Tyr-

rhenotragus (Hurzeler 1983; Thomas 1984). Another

endemic species was named Etruria viallii by Hurzeler

and Engesser (1976) but was not accompanied by a for-

mal description (and hence has been a nomen nudum

TABLE 3 . Measurements (in mm) of upper and lower teeth of

Umbrotherium azzaroli gen. et sp. nov. from Casteani (holotype)

and Fiume Santo.

Specimen Measurements ol ln mw h

IGF 14615 M2–P3 (type)

M2 23.0 21.3 21.3 14.2

M1 19.8 16.2 19.4 10.9

P4 13.6 11.3 17.4 14.4

P3 13.4 12.4 17.2 12.3

FS1994#0461 M3 24.25 22.0 34.0 21.64

IR#63976 M3 22.7 23.0 20.7 24.3

IR#63971 M3 23.5 24.9 24.8 28.3

IR#63975 M3 21.8 20.0 25.0 26.0

SN M3 *22 *22 24.4

SN M2 *23 *23 27.0

FS1995#0312 M2 22.6 23.0 23.5 25.6

FS1994#0437 M2 25.8 23.0 24.0 24.2

IR#63972 M2 24.0 23.2 25.6 26.5

FS1995#0311 M1 21.2 20.4 21.3 17.6

FS1995#0313 M1 22.9 20.5 22.0 8.6

FS1995#0310 P2 12.0 12.7 14.4 22.3

FS1995#0056 M3 33.0 32.0 15.0 22.0

FS1995#0140 M3 31.3 31.6 15.0 16.3

FS1995#0126 M3 28.5 28.0 13.4 26.4

FS1995#0210 M3 29.2 29.5 15.6 11.1

FS1994#0342 M3–P2 *106

M3–M1 62.6 66.5

P4–P3 37.9

M3 28.0 28.7 14.7 11.9

M2 19.8 19.4 14.2 8.4

M1 16.2 16.2 13.2 3.2

P4 15.0 13.6 12.0 10.4

P3 11.2

FS1994#0343 M3–P4

M3–M1 66.2 71.8

M3 28.5 28.0 13.4 26.4

M2 22.2 21.1 14.6 16.0

M1 17.7 17.7 14.8 12.7

P4 15.2 14.4 11.4 16.9

IR#63967 M3 *26 *28 27.3

M2 22.3 *23

IR#63968 M3 27.0 28.0 14.5 21.7

M2 22.6 19.6 21.5

IR#63969 M3 28:4 29.4 15.3 30.1

IR#63974 M2 26.1 21.8 16.0 28.0

IR#64468 M2 22.0 22.0 14.1 15.0

FS1995#0256 M2 25.0 23.0 15.6 20.6

FS1995#0449 M2 23.9 21.3 12.9 22.6

FS1995#0448 M2 or M1 21.3 20.6 15.4 11.5

FS1995#0208 M1 *22 *20 *14 *11

FS1995#0618 M1 19.6 18.7 15.5 12.0

FS1994#0126 P2 8.4 9.8 6.0 9.3

IR64472 P2 10.1 10.9 8.4 12.1

ol, occlusal length; ln, length at the neck; mw, maximum width;

h, height at metastylid ⁄ mesostyle; *, inferred measurement on

slightly broken or damaged elements.

436 P A L A E O N T O L O G Y , V O L U M E 5 1

hitherto), and other forms have been reported in faunal

lists as undetermined Bovidae. We describe these rumi-

nants here for the first time, taking into account both

Tuscan and Sardinian material.

The reconstruction of morphologies of bovid taxa has

been strongly limited by the fact that no dentitions have

been found in direct association with skulls and horns,

and the isolated horn remains of middle size are difficult

to refer to Etruria, which is formally diagnosed below.

Indeed, only for Maremmia and Neotragini, the largest

and smallest bovids, is the association between dentitions

and horn cores possible. The dentitions differ in size,

morphology and height of dental crown (hypsodonty).

On the whole, the species have hypsodont dentition, with

basal pillar and goat-folds usually absent. Maremmia

shows the greatest relative values of dental height (see

below). Description of the dental features follows the ter-

minology of Gentry (1992).

Tribe NEOTRAGINI (Sclater and Thomas, 1894)

Genus TYRRHENOTRAGUS Thomas, 1984

Type species. Tyrrhenotragus gracillimus (Weithofer, 1888) from

Montebamboli, late Miocene, late Tortonian.

Diagnosis (after Thomas 1984). Small neotragini charac-

terized by short and posteriorly orientated horn cores. Very

hypsodont dentitions. Upper premolar series reduced, but

with all dental elements. In upper molars styles are well

developed, in lingual walls lobi are carenated; interlobal

columns are lacking. M3 presents a strongly developed pos-

terior lobe. Lower premolar series reduced, but complete.

In P4 the metaconid is joined to the entoconid and entosty-

lid. Lingual wall of lower molars slightly undulating, labial

lobi carenated, in particular in upper part of wall. The third

lobe of M3 is fairly developed.

Tyrrhenotragus gracillimus (Weithofer, 1888)

Text-figure 10A–I; Table 4

Description. Upper molars are high-crowned. Lower molars have

an undulating lingual wall, with well-developed parastylid. M3

has a variably developed third lobe, which frequently has an

accessory posterior stylid (Text-fig. 10F).

Comparisons and discussion. Tyrrhenotragus gracillimus is

present in both V1 and V2 assemblages in southern Tus-

cany. It is the smallest bovid species. According to Tho-

mas (1984), dental and horn morphologies suggest that it

is referable to the Neotragini.

?Neotragini gen. and sp. indet.

Text-figure 10J–O; Table 4

Description. The Fiume Santo assemblage includes some specimens

that are similar to Tyrrhenotragus, but larger (Text-fig. 11). They are

provisionally referred to an undetermined taxon of Neotragini.

Some fragmentary horn cores are small enough to belong to a neo-

35.0

30.0

25.0

max

imum

wid

h (m

m)

20.0

15.0

10.0

5.020.0 30.0 40.0 50.0 60.0

U. azzarolii

D. pachecoi

B. schaubi

I. arabicum

P. rouenii

S. boissieri

D. cf. pachecoi

occlusal length (mm)

TEXT -F IG . 9 . Plot of length at the neck and maximum width of M3s of Umbrotherium azzaroli from Fiume Santo and some

Miocene giraffid samples. Original data for Decennatherium pachecoi and Bigerbohlinia schaubi from Los Valles de Fuentiduena and

Piera respectively, housed in the Palaeontological Institute ‘M. Crusafont’ in Sabadell; data for Decennatherium cf. D. pachecoi from

Ravin de la Pluie and Nombrevilla are from Geraads (1979); data for Injanatherium arabicum from Al Jadidah are from Morales et al.

(1987); data for Samotherium boissieri and Paleotragus rouenii are from Geraads (1978).

A B B A Z Z I E T A L . : M I O C E N E V E R T E B R A T E A S S E M B L A G E F R O M S A R D I N I A 437

tragin representative, but larger than those of Tyrrhenotragus; they

are short, laterally inserted on the skull and slightly concave.

Comparisons and discussion. The possibility that the

remains here referred to Neotragini indet. belong to Tyr-

rhenotragus males has been considered. Thomas (1981)

reported an important size dimorphism for the presumed

neotragin species Homoidorcas tugenium from the Ngorora

Formation. A size difference of even 30 per cent among the

dental remains from Fiume Santo appears too large, how-

ever, for this to be attributed to sexual dimorphism in Tyr-

rhenotragus (Text-fig. 11); these remains are, therefore,

tentatively attributed to a new taxon. Analyses of sexual

dimorphism in different groups of Recent African ante-

lopes (Jarman 1974; Perez-Barberıa et al. 2002) indicate

that in small species (such as those of the Neotragini) sex-

ual dimorphism is reflected by reduced body size.

Material from the Baccinello-Cinigiano Basin stored in

the Basel Natural History Museum includes some frag-

ments of isolated teeth and horns that are also attribut-

able to this form.

Tribe ?ALCELAPHINI (Rochebrune, 1883)

Genus MAREMMIA Hurzeler, 1983

Type species. Maremmia haupti (Weithofer, 1888); from Casteani

(Grosseto, Central Italy), late Miocene, late Tortonian.

Diagnosis (emended after Hurzeler 1983 and Thomas

1984). Alcelaphini ‘aberrant’, with posteriorly orientated

horn cores. Horn cores subcircular to oval in cross sec-

tion with transverse ridges on anterior surface. Dentitions

very hypsodont with poorly developed roots. Upper pre-

molar series reduced, lacking P2. Upper molars with

enamel islet on occlusal surface. Very reduced M1and very

large M3. Overgrown lower incisors and canine, in the

form of enamel lamellas. Lower molars with undulating

lingual walls and an occlusal surface with an 8-shaped

profile; internal cavities with mediolateral constriction.

Lower premolar series strongly reduced, lacking P2 and

P3. Among lower molars, M1 relatively small, M3 very

large. Posterior part of horizontal ramus of mandible very

high; the ascending ramus is vertically orientated.

Maremmia cf. M. lorenzi Hurzeler, 1983

Text-figures 12–13; Table 5

Description

The most abundant taxon at Fiume Santo, with 852 identified

remains (horns, teeth and postcranial elements).

Horn cores. Horns present an anti-clockwise torsion on the right

side, and a kind of hollowing inside the pedicles. This feature,

which was judged to be absent from the highly deformed mate-

rial from the lignitiferous level of southern Tuscany localities

A B C

DE

F

GH

I J K

L M N

O

TEXT -F IG . 10 . A–I, Tyrrhenotragus gracillimus from Fiume Santo. A–C, FS1995#0039, right M2 in A, labial, B, lingual and C,

occlusal views. D–F, FS1995#0022, right M3 in D, lingual, E, labial and F, occlusal views. G–I, FS1995#0035, right M3 in G, lingual,

H, labial and I, occlusal views. J–O, Neotragini gen. et sp. indet. from Fiume Santo. J–L, FS1995#0071, right M1 in J, occlusal, K,

lingual and L, labial views. M–O, FS1995#0018, left M3 in M, lingual, N, labial and O, occlusal views. All · 2.

438 P A L A E O N T O L O G Y , V O L U M E 5 1

(Thomas 1984; Vrba 1997) is clearly present in the Fiume Santo

assemblage (Text-fig. 12A–B) and in recently discovered remains

from the Baccinello-Cinigiano area (Benvenuti et al. 2001; Text-

fig. 12C–D). In some well-preserved specimens transverse ridges

on the anterior surface of horn cores are evident. The cores are

lateromedially compressed.

Upper dentition. Upper molars with strong styles. M3 is signifi-

cantly more developed than M2 and M1, and shows a character-

istically enlarged metasyle (Text-fig. 13A–C). The premolar

series is reduced, the second premolar being absent.

Lower dentition. Molars have an occlusal surface with the highly

distinctive 8-shaped profile and a stretched connection between

lobes (Text-fig. 13). Lobes have cavities that are constricted cen-

trally. M3 is significantly larger than M2 and M1. The lower pre-

molar row is highly reduced with even the P3 absent. P4 is

molarised with a fused paraconid and metaconid, and short

hypoconid (Text-fig. 13D–E). The overgrown lower incisors are

characterised by the presence of enamel only on the ventral side

of the tooth. The horizontal ramus of the mandible progressively

increases in height from anterior to posterior, under the M3

(Text-fig. 13J–K); the angular tuberosity for the insertion of the

masseter muscle is prominent.

Postcranial bones. The largest bovid remains at Fiume Santo are

attributable to this species. The humerus is characterized by a

medial condyle with an expanded top and, on the ventral side, a

deep bulge.

Comparisons and discussion. The Maremmia material

from Fiume Santo closely resembles M. lorenzi from

the V2 level at Baccinello-Cinigiano. According to

Hurzeler and Engesser (1976) and Hurzeler (1983),

Maremmia is represented in southern Tuscany by two

species belonging to the same phyletic lineage, Marem-

mia haupti and M. lorenzi, occurring in V1 and V2

assemblages respectively. They differ in size and in the

degree of dental series reduction, with a noticeable

increase in upper and lower M3 in M. lorenzi. Thomas

(1984) did not agree with the distinction between M.

haupti and M. lorenzi, only recognizing M. haupti as a

valid taxon. However, biometric comparison of material

from Baccinello-V1, Fiume Santo and Baccinello-V2

underlines, in addition to the larger size of the latter

two samples, the relatively more developed M3 in the

V2 assemblages, which, in our view, justifies their sepa-

ration at the species level. Moreover, Maremmia from

Fiume Santo appears to be slightly larger when com-

pared to the material from the V2 level of Baccinello-

Cinigiano (Text-fig. 14). It is therefore referred to

Maremmia cf. M. lorenzi.

The origin and evolutionary relationships of Maremmia

are controversial. To Hurzeler (1983), the morphology of

the horn cores and dentition (e.g. the 8-shaped profile of

the occlusal surface and P4 morphology) suggested a link

with the Alcelaphini. Thomas (1984) considered the ori-

gin of Maremmia to be closely associated with the middle

Miocene Afro-Eurasian Caprotragoides, and Vrba (1997)

TABLE 4 . Measurements (in mm) of upper and lower teeth of

Tyrrhenotragus gracillimus and Neotragini gen. et sp. indet. from

Fiume Santo; abbreviations as for Table 3.

Taxa ol ln mw h

T. gracillimus

FS1995#0039 M2 6.3 5.6 6.3 7.5

FS1995#0330 M2 8.0 7.2 7.3 9.1

FS1995#0255 M3 6.9 7.4 6.3 9.5

FS1995#0333 M3 6.7 8.1 6.5 10.1

FS1995#0067 M2 6.7 6.3 4.3 5.5

FS1994#0537 M2 8.7 6.9 4.7 11.3

FS1995#0275 M2 8.0 6.5 4.6 9.3

FS1994#0621 M3 13.0 13.0 5.0 12.0

FS1995#0035 M3 12.1 11.9 4.7 9.2

FS1995#0022 M3 10.1 9.6 4.2 10.4

IR#63964 M3 12.5 13.9 4.9 8.4

Neotragini sp. indet.

FS1995#0216 P4 7.2 6.2 8.8 12.6

FS1995#0332 M1 9.3 8.6 8.7 11.0

FS1995#0071 M1 9.7 8.5 8.0 12.2

IR#63959 M1 or M2 10.6 8.3 9.0 14.2

FS1995#0168 P3 7.0 6.8 3.8 3.8

FS1995#0028 M1 or M2 9.2 8.0 5.3 6.0

FS1995#0053 M1 or M2 8.2 8.2 5.3 7.6

IR#63961 M2 9.3 7.8 4.9 12.0

FS1995#0029 M2 10.5 8.2 5.6 4.3

FS1995#0018 M3 15.0 14.7 6.0 7.7

FS1995#0020 M3 13.4 13.5 5.4 9.9

FS1994#0373 M3 14.6 14.0 5.7 11.3

FS1995#0577 M3 13.6 14.7 5.7 13.0

FS1995#0692 M3 12.9 *13 5.5 10.4

Neotragini indet.

7.0

6.0

5.0

4.0

max

imum

wid

th (

mm

)

3.0

2.04.0 9.0 14.0

occlusal length (mm)19.0

Tyrrh. V2 Tyrrh. FSTyrrh. V1

TEXT -F IG . 11 . Plot of occlusal length and maximum width

of M3 of Tyrrhenotragus from Fiume Santo and Baccinello-

Cinigiano, and Neotragini gen. et sp. indet. from Fiume Santo.

A B B A Z Z I E T A L . : M I O C E N E V E R T E B R A T E A S S E M B L A G E F R O M S A R D I N I A 439

believed that Maremmia is the sister taxon of Caprotrago-

ides. Gentry (1997) considered the ‘puzzling’ European

bovid Maremmia haupti to lie close to the ancestry

of African Alcelaphini. Although the similarity with

Alcelaphini is striking, the oldest known records of these

bovids are latest Miocene–Early Pliocene in Africa

(Gentry 1980) and, therefore, clearly post-date the exis-

tence of the Tusco-Sardinian palaeobioprovince. Hence,

the evolutionary relationships of Maremmia remain un-

resolved.

C

DB

A

TEXT -F IG . 12 . Maremmia cf. M. lorenzi from Fiume Santo. A, FS#63942, right and FS#63943, left horn cores in frontal view. B,

FS#63942 in medial view. C–D, IGF 8186, right horn core of Maremmia sp. from Podere Passonaio (Baccinello-Cinigiano Basin) in C,

frontal and D, lateral views. Arrows indicate the occurrence of sinuses inside frontals and pedicles. All · 0.5.

440 P A L A E O N T O L O G Y , V O L U M E 5 1

Subfamily undetermined (?ANTILOPINAE)

Tribe undetermined

Genus ETRURIA gen. nov.

1976 Etruria Hurzeler and Engesser, p. 333 (nomen

nudum).

Derivation of name. From Etruria, Latin name of the region in

Central Italy where the Etruscan civilization developed and the

Oreopithecus fauna existed.

Type and only species. Etruria viallii sp. nov.

Diagnosis. Bovid similar to a gazelle in size. Lower molars

lack basal pillars, with protruding metastylid, lobi tri-

angular in shape with flat labial wall. Third lobe of M3

TABLE 5 . Summary statistics of dentition of Maremmia cf. M. lorenzi from Fiume Santo; abbreviations as for Table 3.

ol ln mw

n min mean max s.d. n min mean max s.d N min mean max s.d.

M3 37 15.7 21.4 26.7 3.14 38 16.1 23.1 29.8 2.87 35 11.6 14.1 18.4 1.61

M2 20 15.9 18.8 27.6 2.40 20 13.5 16.9 27.5 2.97 19 10.5 12.9 17.3 1.60

M1 9 14.0 16.9 19.8 1.75 9 11.0 13.1 15.5 1.63 8 9.8 11.3 13.0 1.05

P4 2 10.2 10.2 10.2 – 2 7.5 8.5 9.5 1.41 2 7.5 7.6 7.7 0.14

P3 1 – 8.0 – – – – – – – 1 – 5.0 – –

M3 29 21.6 25.5 29.3 1.62 27 25.5 25.5 30.6 1.54 25 8.0 9 10.1 0.55

M2 19 14.1 18.8 21.5 2.2 18 13.0 15.7 18.0 1.57 15 7.1 7.8 9.3 2.25

M1 13 11.3 16.5 19.3 2.1 13 12.8 12.8 15.5 1.43 13 6.5 7.5 8.8 0.66

P4 5 10.9 11.8 12.9 0.75 5 8.1 9.4 10.2 0.99 4 5.3 5.6 6.0 0.31

A B

C

D E F G

HN

I K L M

J

TEXT -F IG . 13 . Photographs and drawings of Maremmia cf. M. lorenzi from Fiume Santo. A–C, FS1995#0488, left M3 in A, labial,

B, lingual and C, occlusal views; · 0.5. D, FS1995#0692, left P4 in occlusal view; · 1. E, FS1995#0692, left unworn P4 in occlusal

view; · 1.5. F–H, FS1995#0300, left M3 in F, occlusal, G, labial and H, lingual views; · 1. I–K, FS1994#0344, left mandible with M3–

M1 in I, occlusal, J, labial and K, lingual views; · 0.5. L–N, FS1995#0247, right M2 in L, lingual, M, labial and N, occlusal views; · 1.

10.5M. hauptiVI

M. lorenziFS

M. lorenziV2

10.09.59.08.58.0

wid

th (

mm

)

7.57.06.56.0

15.0 20.0 25.0Length at neck (mm)

30.0

Type

35.0

TEXT -F IG . 14 . Plot of length at the neck and maximum

width of M3 in Maremmia haupti, M. lorenzi and Maremmia cf.

M. lorenzi from Baccinello-Cinigiano Basin levels V1 and V2,

and Fiume Santo. The holotype of M. lorenzi n. Bac93 from

Trasubbie Creek, stored in Basel Naturhistorisches Museum, is

indicated by the arrow.

A B B A Z Z I E T A L . : M I O C E N E V E R T E B R A T E A S S E M B L A G E F R O M S A R D I N I A 441

fairly well developed and with a central cavity; metastylid

disappears with advanced wear stages. Lower premolars

short, in particular P2, whose length represents about 18

per cent of length of premolar row; P3 triangular in shape

with well-developed parastylid and paraconid; P4 some-

what primitive with an anterior valley between paraconid

and metaconid, while metaconid, entoconid and endo-

stylid tend to be fused lingually (Text-fig. 15).

Etruria viallii sp. nov.

Text-figure 15; Table 6

1976 Etruria viallii Hurzeler and Engesser, p. 333 (nomen

nudum).

Derivation of name. In honour of Prof. Vittorio Vialli (1914–

1983), eminent palaeontologist at Bologna University.

Holotype. Bac1002, a right fragmented mandible with complete

tooth series P2–M3, housed in the Basel Natural History Museum

(Text-fig. 15A–C); for measurements of specimen, see Table 6.

Locality, horizon and age. Baccinello (Grosseto district, southern

Tuscany), lignite level, faunal assemblage V1; late Miocene, late

Tortonian, late Turolian.

Other referred material. Fiume Santo, north-west Sardinia: see

Supplementary Data file (http://www.palass.org).

Diagnosis. As for the genus.

Description. Upper dentition: P4 rectangular in outline (long and

narrow); the anterior crista of the hypocone does not join the

parastyle; the fragmentary upper molars have poorly developed

styles. Lower dentition: the Fiume Santo material adds nothing to

the features previously recorded from the type material.

Comparisons and discussion. Etruria viallii has the lowest

degree of dental reduction and hypsodonty among the

bovids of the Tusco-Sardinian palaeobioprovince. Unlike

Maremmia spp. and Bovidae gen. et sp. nov. (see below),

the dental roots are well developed. The dental characters

described for E. viallii were reported by Gentry (1992) to

be distinctive of the Antilopini tribe (e.g. Gazella).

Subfamily undetermined

Tribe undetermined

Genus TURRITRAGUS gen. nov.

1918 Gazella haupti Del Campana, p. 170, pl. 16,

figs 4A–C, 6A–C.

1984 Maremmia haupti Thomas, p. 86.

Derivation of name. After the town of Porto Torres in north-

west Sardinia, whose surroundings include the industrial

complex in which the Fiume Santo site is located.

Type and only species. Turritragus casteanensis sp. nov.

Diagnosis. Bovid of small size, similar to Etruria vialli;

with hypsodont dentition, roots very short. Highly

reduced dental series, lacking P2 and P2–P3. Upper molars

have a well-developed mesostyle, basal pillars are absent.

Upper premolars rectangular in outline. Inner islets of

upper and lower molars show an evident anterior–poster-

ior compression. Lower molars have an undulating lin-

gual wall and bear a metastylid only at a low level of

wear. M3 bears a narrow hypoconulid. P4 not molarised

and lacks transverse metaconid crest. Hypoconid sepa-

rated from metaconid by a groove (Text-fig. 16).

Remarks. The revision of the material from Casteani and

other sites in southern Tuscany housed in the Natural

History Museum of the University of Florence, and analy-

sis of the Fiume Santo fossils allows us to recognise

TABLE 6 . Measurements (in mm) of upper and lower teeth of

Etruria viallii gen. et sp. nov. from Casteani (including the holo-

type) and Fiume Santo; abbreviations as for Table 3.

Taxa ol ln mw h

Casteani

Bac 1002 M3–P2 (type) 67.2 66.4

M3–M1 43.0 40.1

P4–P2 24.1 23.1

M3 15.8 16.5 7.3 15.6

M2 13.0 10.0 7.0 12.1

M1 11.0 9.0 7.0 9.0

P4 9.2 7.6 6.1 9.3

P3 8.0 6.1 6.0 8.0

P2 4.4 4.8 4.2 5.0

Bac 1003 M3–M1 39.4 39.3

M3 18.0 18.7 9.1 5.4

M2 11.5 10.2 8.7 4.4

M1 9.7 8.2 7.3 1.6

P4 7.2 6.3 6.1 2.5

P2 6.8 6.1 4.8 3.5

Fiume Santo

FS1995#0145 M2 12.9 *11 *11 14.9

FS1995#0136 M1 12.7

FS1995#0027 M2 13.1 11.6 6.5 14.7

IR#64440 M2 15.2 13.7 16.1 7.5

IR#64452 M3 17.3 18.3 5.1

IR#64452 M2 10.3 10.3 3.4

FS1995#0620 M3 18.5 17.4 7.9 12.7

FS1995#0088 P3 7.5 6.6 4.5 9.0

FS1995#0014 M3 19.1 17.5 7.6 6.6

FS1995#0240 M3 18.0 17.6 11.5

FS1994#0520 P4 11.7 9.2 5.8 14.9

442 P A L A E O N T O L O G Y , V O L U M E 5 1

another new middle-sized to small bovid species. The

remains from Casteani were ascribed to M. haupti by

previous authors.

Turritragus casteanensis sp. nov.

Text-figure 16; Table 7

Derivation of name. After Casteani, type locality of the species.

Holotype. IGF 14631, a left mandible with complete tooth series

P4–M3, housed in the Natural History Museum, Geology and

Palaeontology section, University of Florence (Text-fig. 16A–C);

for measurements, see Table 7.

Referred material from type locality. IGF 11746, left mandible

with P4–M3 (Text-fig. 16D–F); IGF 14633, upper series P3–M2

(Text-fig. 16G–I) (Table 7); for material from the Fiume Santo

site, see Supplementary Data file (http://www.palass.org).

Locality, horizon and age. Casteani (Grosseto district, southern

Tuscany), Faunal assemblage V1; late Miocene, late Tortonian,

late Turolian

Diagnosis. As for the genus.

Comparisons and discussion. The material from Casteani

indicates another bovid species which, like Maremmia,

has a markedly reduced dental series. The shortening of

the dental series and the highly hypsodont teeth make

Turritragus casteanensis, by comparison with Maremmia,

very derived for its chronological position. A reduction of

the premolar series, although with all elements present, is

recorded in Aragoral mudejar from the Upper Vallesian of

Spain, which is considered to be a primitive representa-

tive of the Caprinae. Very hypsodont bovid species were

described by Robinson (1986) from late Middle Miocene

deposits in Tunisia, and regarded as primitive representa-

tives of the Rupicaprini. The absence of cranial parts and

horn cores in T. casteanensis prevents precise determina-

tion of its systematic affiliation.

Very few isolated highly hypsodont teeth referable to T.

casteanensis are present in the Fiume Santo collection

(Text-fig. 16J–L). They do not add anything new to the

diagnostic features recorded from the type material from

Casteani.

Remarks on bovid species from the Tusco-Sardinian

palaeobioprovince

As a result of our analysis it is possible to compile a list

of dental differences, other than size, for distinguishing

between the various species (Table 8; Text-fig. 17). The

most significant features and profound changes in the

A

B

C

D

E

F

TEXT -F IG . 15 . Photographs and drawings of Etruria viallii gen. et sp. nov. from southern Tuscany and Fiume Santo. A–C,

Bac1002, holotype, right fragmentary mandible with M3–P2 in A, occlusal, B, lingual and C, labial views; · 1. D–F, FS1995#0027,

right M2 from Fiume Santo in D, lingual, E, occlusal and F, labial views, · 1.5.

A B B A Z Z I E T A L . : M I O C E N E V E R T E B R A T E A S S E M B L A G E F R O M S A R D I N I A 443

masticator apparatus are found in Maremmia: a strong

reduction of the premolar series and enlargement of lower

and upper third molars, which indicates a tendency to

chew posteriorly closer to the fulcrum of the mandible;

and a shortening and deepening of the horizontal

mandibular ramus. The diagnostic dental features of

Maremmia also include a high degree of hypsodonty.

Hypsodonty has been widely used as an indicator of feed-

ing preferences in many mammal categories and is con-

sidered to be a functional adaptation to an abrasive diet

(Janis 1995; Fortelius et al. 2002) and climatic parameters,

in particular aridity (Damuth and Fortelius, 2001; Forte-

lius 2003). The calculated values of this index for the bo-

vids of the Tusco-Sardinian palaeobioprovince (Table 8;

h = height of unworn M2 divided by the length according

to Fortelius et al. 2002; Fortelius 2003) indicate that it

A

D E

F

G

H I

J

K

L

B C

TEXT -F IG . 16 . Photographs and drawings of Turritragus casteanensis gen. et sp. nov. from Southern Tuscany and Fiume Santo. A–

C, IGF 14631, holotype, left mandible with M3–P4 from Casteani in A, labial, B, lingual and C, occlusal views. D–F, IGF 11746, left

mandible with M3–P4 from Casteani in D, labial, E, occlusal and F, lingual views. G–I, IGF 14633, left maxillary with M2–P3 from

Casteani in G, occlusal, H, lingual and I, labial views. J–L, FS1995#0091, left M2 from Fiume Santo in J, labial, K, lingual and L,

occlusal views. All · 1.

444 P A L A E O N T O L O G Y , V O L U M E 5 1

can be classified as a hypsodont species (h > 1.2) and sug-

gests the occurrence of open, dry environments (for the

inverse connection between hypsodonty and humidity,

see Damuth and Fortelius 2001; Fortelius et al. 2002,

2003; Fortelius 2003; Eronen and Rook 2004; Eronen

2006).

With regard to the postcranium, 83 bovid remains

have been identified in the Fiume Santo collection. The

most common and best preserved skeletal element is the

astragalus.

Specific characters, e.g. articular facets or areas for ten-

don ⁄ ligament insertion, are difficult to discern on the

bones because the material is highly weathered; however,

after a preliminary analysis, some differences in the shape

of the tuber calcanei of the calcaneus and in the proximal

articulation of the metatarsals suggest differences in loco-

motor behaviour among the various bovid taxa, and con-

firms the occurrence of more than one middle-sized

species (Etruria and Turritragus). Two metacarpals (both

recorded as Bac1013 in the collection of the Basel Natural

History Museum; Text-fig. 18) from the Baccinello-Cini-

giano Basin are characterised by a strong shortening (total

length, c. 60 mm; proximal width, c. 14 mm; distal width,

c. 16 mm). These dimensions and proportions allow their

exclusion from Neotragini and Maremmia; it is possible

that they are referable to Etruria or Turritragus casteanen-

sis. The shortening of metapodials, which reflects an

adaptation to rough, rocky ground, is well documented

among bovids and frequently found in Caprinae and

more primitive forms (Alcala and Morales 1997). The

strong correlation between size and morphology of distal

limb bones and habitat is well established in Recent bovid

communities. Limb morphology reflects speed, jumping

capacity and ground conditions. However, in the case of

the Fiume Santo bovids, more comprehensive ecological

characterisation will only be possible following analysis of

more abundant postcranial remains.

THE FAUNAL ASSEMBLAGES OF THELATE MIOCENE TUSCO-SARDINIANPALAEOBIOPROVINCE

The faunal succession V0–V1–V2 within the sedimentary

succession across the various southern Tuscan basins (OZF

Oreopithecus Zone faunas in Bernor et al. 2001) is a classic

TABLE 7 . Measurements (in mm) of upper and lower teeth of

Turritragus casteanensis gen. et sp. nov. from Casteani (including

the holotype) and Fiume Santo; abbreviations as for Table 3.

Taxa ol ln mw h

Casteani

IGF 14633 M2–P3

M2 12.0

M1 10.8 *8 10.4 15.0

P4 *8 6.3 8.6 15.7

P3 8.0 7.0 6.4 10.5

IGF14631 M3–M1 46.1 47.4

M3 19.2 20.0 8.0 25.0

M2 13.7 12.8 7.3 10.6

M1 10.8 7.9 6.7 11.3

P4 9.6 8.3 4.3 12.3

IGF11746 M3–M1 (type) 44.0 48.0

M3 17.8 19.6 6.7 30.3

M2 13.7 12.5 7.3 8.4

M1 10.3 9.0 7.0 7.0

P4 8.8 8.3 4.4 7.0

Fiume Santo

IR#63950 M3 15.0 16.3 12.7 28.4

FS1995#0091 M2 14.4 13.0 7.2 22.0

IR#63937 M2 14.5 14.1 7.5 22.6

TABLE 8 . Distinctive features in bovid dentitions from Fiume Santo and Southern Tuscany localities.

Taxa T. gracillimus Etruria viallii T. casteanensis M. cf. lorenzi

Characters

Deepening of horizontal ramus,

beneath molars

moderate moderate intermediate strong

Premolar reduction moderate1 moderate1 yes2 yes2

Hypsodonty high (1.6) moderate (1.3) high (>1.7*) very high (2.1)

Molarization of P4, with

paraconid-metaconid fusion

no no no yes

Stretched connection between

lobes of lower molars

no no no yes

Metasylid development early intermediate very early very early

Development of posterior lobe of M3 intermediate high low low

Compression of central cavities

of molars

constricted

anteroposteriorly

constricted

anteroposteriorly

constricted

anteroposteriorly

constricted centrally

1, P2 is short but always present; 2, P2 is lacking; *, only one slightly worn M2 was available.

A B B A Z Z I E T A L . : M I O C E N E V E R T E B R A T E A S S E M B L A G E F R O M S A R D I N I A 445

example of a chronofauna according to the concept of

Olson (1952), which aims to indicate the persistence and

continuity through time of evolving animal assemblages

under changing environmental conditions. The Fiume

Santo vertebrate assemblage offers the opportunity to com-

pare local populations belonging to the same time span as

the V2 assemblage, within the Tusco-Sardinian chrono-

fauna.

Overall, the Fiume Santo fauna is very similar to the V2

assemblage (Montebamboli and Baccinello V2 faunas).

Nevertheless, there are some differences, such as the slightly

derived features in the suid Eumaiochoerus cf. E. etruscus

and in the antelope Maremmia cf. M. lorenzi, which could

be a result of environmental differences, and perhaps also

related to slight chronological differences. The dominance

of bovids with a high degree of hypsodonty in the mammal

assemblage from Fiume Santo suggests wide open spaces;

forest-adapted forms are under-represented. The occur-

rence and ⁄ or abundance of taxa generally in the so-called

Oreopithecus assemblages seem to have been strongly influ-

enced by environmental conditions. The Fiume Santo

time-equivalent (or evolutionary-equivalent) V2 assem-

blages in Tuscany vary in composition. While the suid

Eumaiochoerus and the primate Oreopithecus are abundant

at Montebamboli, both are extremely rare (Oreopithecus)

or absent (Eumaiochoerus) in assemblages from the Bacci-

nello-Cinigiano Basin (Rook et al. 1999a). Such differences

are easily explained if the sedimentary and palaeoenviron-

mental context of the sites is taken into account: swampy

and marshy-margined lake at Montebamboli versus flood-

dominated fluvio-lacustrine sediments near an uplifted

delta plain in the Baccinello-Cinigiano Basin (Benvenuti

et al. 1999, 2001).

ZOOGEOGRAPHY OF TUSCO-SARDINIAN LATE MIOCENEBIOPROVINCE

A zoogeographic study of the Fiume Santo faunal assem-

blage was carried out by means of the Genus Faunal

Resemblance Index (GFRI) between the Tuscan OZF

(Oreopithecus faunal assemblages: Baccinello V0–V2,

Montebamboli, Casteani, Montemassi, Ribolla, Serrazz-

ano, Baccinello V0–V2) and Fiume Santo together with

some European localities across the MN10–13 time inter-

val. For this purpose, we based our analyses on the Neo-

gene Old World (NOW) Database. This database was

released to the public in 1996 and since then the number

and geographic range of localities have been increased

substantially. The latest public NOW dataset can be freely

downloaded from the website http://www.helsinki.fi/

science/now/.

12.0

10.0

8.0

wid

th (

mm

)

6.0

4.0

2.05.0 10.0 15.0 20.0

occlusal length (mm)

25.0 30.0 35.0

E. vialli

T. casteanensis

M. cf. lorenzi

Neotragini indet.

T. gracillimus

TEXT -F IG . 17 . Plot of occlusal length

and maximum width of bovid M3s from

Fiume Santo.

TEXT -F IG . 18 . Bovidae indet., Bac1013, right metacarpal

from Baccinello, anterior view; · 1.

446 P A L A E O N T O L O G Y , V O L U M E 5 1

According to the procedure followed by Bernor et al.

(2001, 2004) we calculated both Dice and Simpson

GFRIs. Dice’s index is the most commonly used faunal

resemblance index (especially by zoologists), while Simp-

son’s index has long been used by palaeontologists and

additionally adjusts for differences in sample size, which

is a problem in the present analysis (see Bernor et al.

2001 and 2004 for a discussion of the method). The GFRI

graph (Text-fig. 19) shows, as expected, very high values

of resemblance (100 per cent) for the Fiume Santo and

southern Tuscany assemblages, while the GFRI is very low

for the rest of the faunas. Apart from the obvious conclu-

sions regarding the endemic character of the Tusco-Sardi-

nian palaeobioprovince, it is interesting to note that the

GFRI index is 0 for the late Miocene faunal assemblages

of Greece (Ouranopithecus Faunal Zone), meaning that no

taxa (at generic level) are in common between the two

faunas. Very low values characterise comparisons between

the Oreopithecus Faunal Zone and other, older (Can

Llobateres) and younger (Pikermi; Baccinello V3; Mar-

amena) assemblages, owing to the common occurrence of

a few genera of micro-mammals and carnivores (Mustela,

Huerzelerimys, Kowalskia, Parapodemus).

It has been suggested that the survival in the latest

Miocene (MN11–12) Tyrrhenian area of the extremely

derived ape Oreopithecus bambolii (in contrast to the

extinction pattern of European mainland hominoids since

MN10), was linked to a particular combination of palaeo-

geographic and palaeoclimatic conditions. The terminal

Miocene palaeogeographic changes in the Tyrrhenian

area, a consequence of the intensive Messinian tectonism

which produced the Apennine mountain chain, led to the

end of the Tusco-Sardinian palaeobioprovince and the

extinction of the Oreopithecus bambolii insular fauna

(Rook et al. 2000).

The origin of the endemic vertebrates in the Tusco-Sar-

dinian province has been discussed in detail in the litera-

ture. On the basis of the presence of the non-endemic

murid Huerzelerimys vireti (Engesser 1989) in the older

levels of the Baccinello-Cinigiano Basin (V0 assemblage),

it appears that the entry of the ancestors to the endemic

taxa may be dated close to the beginning of the Turolian

(MN 11 unit; Mein 1999). However, because of the lim-

ited similarities, the correlation of endemic faunal succes-

sions with the late middle ⁄ early late Miocene continental

European and African faunal successions is problematic.

Undeniably, the systematics and, especially, estimates of

phylogenetic relationships of insular taxa are affected by

endemic evolutionary modification. In the case of long-

lasting geographic isolation, the endemic changes could

be so great as to render the identification of ancestral taxa

very difficult, as it becomes increasingly hard to establish

which characters are inherited from ancestral taxa and

which are derived as a result of endemic evolution. The

phylogenetic affinities of some of the mammals, such as

Oreopithecus and the suid Eumaiochoerus, lie mainly with

species from the European continent (Moya-Sola and

Kohler 1997; Bernor et al. 2001); however, those of other

taxa, in particular the bovids, are more contentious and,

as yet, poorly understood, although it has been suggested

previously that Maremmia and Tyrrhenotragus were Afri-

can in origin (Hurzeler and Engesser 1976; Hurzeler

1983; Thomas 1984; Abbazzi in press).

The Tusco-Sardinian palaeobioprovince chronofauna

became extinct at the end of latest Miocene when the

youngest endemic faunal assemblages were replaced by

the dispersal of a completely renovated mammal fauna

(the so called Bacinello V3) made up of continental taxa

with fully continental European affinities (Hurzeler and

Engesser 1976; Bernor et al. 2001) and different environ-

mental tracking (Eronen and Rook, 2004), pointing to a

renewed and definite biogeographical connection with

Europe. The ultimate cause of this faunal turnover is

attributed to the intense tectonic activity that affected the

area during latest Messinian times and led to the uplift of

the Apennine mountain chain (Boccaletti et al. 1990); the

slopes of the newly emerged Apennines constituted a wide

pathway for the dispersal of mammal communities. This

faunal change marks the time when the Corso-Sardinian

Massif was definitely isolated from southern Tuscany by

the opening of the Tyrrhenian Sea and southern Tuscany

became fully connected to the Apennine chain (Benvenuti

et al. 2001; Rook et al. 2006).

CONCLUSIONS

The rock succession at Fiume Santo contains the richest

and best fossil record of the youngest stage in the evo-

1.00

0.80

0.60

0.40

0.20

0.00A B C D E F

TEXT -F IG . 19 . Genus-level Faunal Resemblance Index

(GFRI): pair-wise comparison between the set of localities under

consideration and the Tuscany ‘Oreopithecus FZ’: A, Can

Llobateres; B, Ouranopithecus FZ; C, Fiume Santo; D, Pikermi;

E, Baccinello V3; F, Maramena; black bars, Dice index; white

bars, Simpson index. Comparative data updated from Fortelius

(2005).

A B B A Z Z I E T A L . : M I O C E N E V E R T E B R A T E A S S E M B L A G E F R O M S A R D I N I A 447

lution of the chronofauna that in latest Miocene times

(late Tortonian) characterised the North Tyrrhenian

area, the so-called Tusco-Sardinian palaeobioprovince.

During the latest Miocene, the North Tyrrhenian area,

a region presently corresponding to southern Tuscany

and the Sardinian Massif, was occupied by a complex

of large islands characterised by endemic vertebrate

populations. Morphological features, adaptations and

trophic structure (e.g. low diversity and scarce carnivore

taxa), mainly in the mammal faunas, attest to a long

period of isolation from the continental palaeobiopro-

vinces of the Mediterranean and Central Europe. The

analysis of the Sardinian fossils and their comparison

with material previously recovered from various locali-

ties in Tuscany, allowed us to re-evaluate the latter and

to describe three new endemic taxa among the rumi-

nants. Tusco-Sardinian vertebrate assemblages are char-

acterized through time and across geographically

separated areas (Tuscany vs. Sardinia at the level of V2

faunas), and consideration is given to zoogeographic

patterns and evidence of evolution of an endemic chro-

nofauna in insular conditions, and its extinction during

the latest Miocene times.

Acknowledgements. This work was carried out under an agree-

ment between the ‘Soprintendente ai Beni Archeologici per le

Province di Sassari e Nuoro’ and the Vertebrate Palaeontology

Group of the University of Florence. We are indebted to the Sas-

sari Soprintendente, Dr Francesco Nicosia for the support. We

thank all the personnel of the ‘Fiumesanto’ thermoelectric power

station for their help during all phases of field work, in particu-

lar the former and present directors of the power station, Drs

Francesco Capriotti and Marco Bertolino respectively, and the

person responsible for external relations, Mr Mario Abeltino.

For facilitating access to fossil collections in their care, we thank

Burkart Engesser and Christian A. Meyer (Naturhistorisches

Museum, Basel), Elisabetta Cioppi and Menotti Mazzini (Natu-

ral History Museum, Geology and Palaeontology Section, Uni-

versity of Florence), Walter Landini and Chiara Sorbini (Museo

di Storia Naturale e del Territorio di Calci, Pisa), Salvador

Moya-Sola and Meike Kohler (Barcelona), Bernard Battail and

France de Lapparent de Broin (Musee national d’Histoire natu-

relle, Paris). Among numerous people who have helped greatly

with suggestions and discussion we mention Marisa Arca and

Caterinella Tuveri (Nuoro), Raymond L. Bernor (Washington,

DC), F. Clark Howell (Berkeley), Jorge Morales and Jan van der

Made (Madrid), Salvador Moya-Sola and Meike Kohler (Saba-

dell) and Maria Rita Palombo (Rome). The editorial work of

David J. Batten (Manchester) is appreciated. Our research at Fi-

ume Santo is part of a wider research program on the evolution

of late Neogene mammal faunas co-ordinated by LR at the Uni-

versity of Florence. Field work at Fiume Santo was made possi-

ble thanks to the support of the National Geographic Society

(grant #7484-03 to LR), the RHOI program at University of

Berkeley (project NSF-BCS-0321893), and the logistic and eco-

nomic support of ENDESA Italia. For the purposes of this

paper, LA dealt with Bovidae and Giraffidae, MD with Herpe-

tofauna, GG with Suidae, and LR with Primates, Carnivora and

zoogeography. All contributed equally to the general discussion

and concluding sections.

REFERENCES

A B B A Z ZI , L. in press. Late Miocene endemic bovids in the

Tyrrhenian palaeobioprovince: from Africa or Europe? Pro-

ceedings of the Third Symposium on Sedimentary Basins of

Libya, Geology of East Libya. Benghazi, 21–23 November 2004.

A L C A L A , L. and M OR A L E S , J. 1997. A primitive caprine

from the Upper Valesian of La Roma 2 (Alfambra, Teruel,

Aragon, Spain). Comptes Rendus de l’Academie des Sciences de

Paris, Serie 2, 324, 947–953.

B E N V E N U T I , M., PA P I N I , M. and T E S T A , G. 1999.

Sedimentary facies analysis in paleoclimatic reconstructions.

Examples from the Upper Miocene–Pliocene successions of

the south-central Tuscany (Italy). 355–377. In A G U S T I , J.,

R OO K, L. and A N D R E W S , P. (eds). Hominoid evolution

and climatic change in Europe. Volume 1. The evolution of Neo-

gene terrestrial ecosystems in Europe. Cambridge University

Press, Cambridge, 512 pp.

—— —— and RO O K, L. 2001. Mammal biochronology,

UBSU and paleoenvironment evolution in a post-collisional

basin: evidence from the Late Miocene Baccinello-Cinigiano

Basin in southern Tuscany, Italy. Bollettino della Societa Geo-

logica Italiana, 120, 97–118.

B E R N O R, R. L., FO R T E L I U S , M. and R O OK , L. 2001.

Evolutionary biogeography and paleoecology of the ‘‘Oreopi-

thecus bambolii Faunal Zone’’ (late Miocene, Tusco-Sardinian

Province). Bollettino della Societa Paleontologica Italiana, 40,

139–148.

—— K OR D O S , L., R O O K, L., A G US T I , J., A N DR E W S ,

P., A R M O U R - CH E L U, M., B E G UN , D. R., C A M E R -

ON , D. W., D A M U TH , J., DA X N E R - HO C K, G., DE

BO N I S , L., F E J F A R , O., F E S S A H A , N., FO R T E L I U S ,

M., F R A N ZE N , J., G A S P A R I K , M., G E N T R Y , A.,

HE I S S I G, K., HE R N Y A K, N., K A I S E R , T., KO UF O S ,

G. D., K R OL O P P, E., J A N OS S Y , D., L L E N A S , M.,

M E S Z A R O S , L., M U L L E R , P., R E N N E , P., R O CE K , Z.,

S E N , S., S C OT T , R., S ZY N DL A R , Z., TO P A L , G. Y.,

UN G A R , P. S., UT E S C H E R , T., V A N D A M , J. A.,

W E R DE L I N , L. and Z I E G L E R , R. 2004. Recent advances

on multidisciplinary research at Rudabanya, Late Miocene

(MN9), Hungary: a compendium. Palaeontographia Italica, 89

(for 2002), 3–36.

B OC C A L E T TI , M., C I A R A N F I , N., C OS E N TI N O, D.,

DE I A N A , G., G E L A TI , R., L E N T I N I , F., M A S S A R I , F.,

M O R A T TI , G., P E S C A TO R E , T., R I CC I L U CC H I , F.

and T OR T O R I C I , L. 1990. Palinspastic restoration and

paleogeographic reconstruction of the peri-Tyrrhenian area

during the Neogene. Palaeogeography, Palaeoclimatology,

Palaeoecology, 77, 41–50.

B OW DI CH , T. E. 1821. An analysis of the natural classifications

of Mammalia, for the use of students and travellers. J. Smith,

Paris, iv +115 pp.

448 P A L A E O N T O L O G Y , V O L U M E 5 1

B R O CH U, C. A. 2000. Phylogenetic relationships and diver-

gence timing of Crocodylus based on morphology and the fos-

sil record. Copeia, 3, 657–673.

B R O N GN I A R T , A. 1800. Essai d’une classification naturelle

des reptiles. Bulletin de la Societe Philomatique, 3, 2, 11 (=35),

81–82; 12 (=36), 88–91.

C OR D Y , J. M. and G I N E S U , S. 1994. Fiume Santo (Sassari,

Sardaigne, Italie): un nouveau gisement a Oreopitheque (Or-

eopithecidae, Primates, Mammalia). Comptes Rendus de

l’Academie des Sciences de Paris, Serie 2, 318, 679–704.

—— —— O ZE R, A. and S I A S , S. 1995. Geomorphological and

palaeoecological characteristics of the Oreopithecus site of

Fiume Santo (Sassari, northern Sardinia, Italy). Geografia Fisi-

ca e Dinamica Quaternaria, 18, 7–16.

C R US A FO N T P A I R O , M. 1952. Los jirafidos fosiles de

Espana. Diputacion provincial de Barcelona, Memoria y

Comunicaciones del Instituto Geologico, Consejo Superior de

Investigaciones Cientificas, Barcelona, 239 pp.

D A M U T H, D. J. and F OR T E L I US , M. 2001. Reconstructing

mean annual precipitation, based on mammalian dental mor-

phology and local species richness. 23–24. In A G US T I , J.

and O M S , O. (eds). EEDEN plenary workshop on Late Mio-

cene to Early Pliocene environments and ecosystems. ESF, Saba-

dell (Spain), 78 pp.

D E L CA M PA N A , D. 1918. Considerazioni sulle Antilopi

terziarie della Toscana. Palaeontographia Italica, 24, 147–

234.

D E L F I N O , M. 2008. Late Neogene crocodylian faunas from

Libya and the Mediterranean area. Proceedings of the Third

Symposium on Sedimentary Basins of Libya, Geology of East

Libya. Benghazi, 21–23 November 2004.

—— and R O OK , L. 2008. African crocodylians in the Late

Neogene of Europe: a revision of Crocodylus bambolii Ristori,

1890. Journal of Paleontology, 82, 336–343.

—— B OH M E , M. and R O OK , L. 2007. First European

evidence for transcontinental dispersal of Crocodylus (late

Neogene of southern Italy). Zoological Journal of the Linnean

Society of London, 149, 293–307.

E N G E S S E R , B. 1989. The Late Tertiary small mammals of

the Maremma region (Tuscany, Italy). II Part: Muridae and

Cricetidae (Rodentia, Mammalia). Bollettino della Societa

Paleontologica Italiana, 29, 227–252.

E R O N E N , J. 2006. Eurasian Neogene large herbivorous

mammals and climate. Acta Zoologica Fennica, 216, 1–71.

—— and R OO K , L. 2004. The Mio-Pliocene European primate

fossil record: dynamics and habitat tracking. Journal of Human

Evolution, 4, 323–341.

F E R N A N D E Z - J A L V O , Y., S A N C H E Z- C H I L L O N , B.,

A N DR E W S , P., F E R N A N D E Z - L OP E Z , S. and A L -

CA LA M A R T I N E Z , L. 2002. Morphological taphonomic

transformations of fossil bones in continental environments,

and repercussions on their chemical composition. Archaeo-

metry, 44, 353–361.

F I C C A R E L L I , G. and T O RR E , D. 1967. Il mustelide Enhy-

drictis galictoides del Pleistocene della Sardegna. Palaeonto-

graphia Italica, 63, 139–160.

F I S CH E R D E W A L DH E I M , G. 1817. Adversaria zoologica.

Memoires de la Societe Imperiale Naturelle (Moscow), 5, 368–482.

F O R TE L I US , M. 2003. Evolution of dental capability in

western European large mammal plant eaters 22–2 million

years ago: a case for environmental forcing mediated by

biological processes. 61–67. In L E G A KI S , A., S F E N T H O-

UR A K I S , S., PL Y M E N I , R. and T H E S S A L O U - L E G A -

KI , M. (eds). The new panorama of animal evolution. 18th

International Congress of Zoology, Pensoft, Sofia and Mos-

cow, 264 pp.

—— (co-ord.) 2005. Neogene of the Old World Database of

Fossil Mammals (NOW). University of Helsinki. http://

www.helsinki.fi/science/now/. Public release 030717.

—— E R O N E N , J., J E R N V A L L , J., L I U , L., PU S H KI N A ,

D., R I N N E , J., T E S A K OV , A., V I S L O BO K OV A , I.,

Z HA N G , Z. and Z H OU , Z. 2002. Fossil mammals resolve

regional patterns of Eurasian climate change over 20 million

years. Evolutionary Ecology Research, 4, 1005–1016.

—— —— L I U, L., TE S A KO V , A., V I S L O B OK O V A , I. and

Z HA N G , Z. 2003. Continental-scale hypsodonty patterns, cli-

matic paleobiogeography and dispersal of Eurasian Neogene

large mammal herbivores. Deinsea, 10, 1–11.

G E N T R Y , A. W. 1980. Fossil Bovidae (Mammalia) from

Langebaanweg, South Africa. Annals of the South African

Museum, 79, 213–337.

—— 1992. The subfamilies and tribes of the family Bovidae.

Mammal Review, 22, 1–32.

—— 1997. Fossil ruminants (Mammalia) from the Manonga

Valley, Tanzania. 107–135. In H A R R I S O N , T. (ed.). Neogene

paleontology of the Manonga Valley. Topics in Geobiology.

Plenum Press, New York, NY, 418 pp.

G E R A A D S , D. 1978. Les Palaeotraginae (Giraffidae, Mamma-

lia) du Miocene superieur de la region de Thessalonique

(Grece). Geologie Mediterraneenne, 5, 269–276.

—— 1979. Les Giraffinae (Artiodactyla, Mammalia) du Miocene

superieur de la region de Thessalonique (Grece). Bulletin du

Museum National d’Histoire Naturelle, Paris, Serie 4, C, 4,

377–389.

—— 1986. Remarques sur la systematique et la phylogenie des

Giraffidae (Artiodactyla, Mammalia). Geobios, 19, 465–478.

G E R V A I S , P. 1872. Sur un singe fossile d’un espece non

ancore decrite, qui a ete decouverte au Monte Bamboli.

Comptes Rendus Hebdomadaires des Seances de l’Academie des

Sciences de Paris, 74, 1217–1223.

G M E LI N , J. F. 1789. Regnum animalia. 1033–1516. In B E E R ,

G. E. (ed.). Caroli a linne systema naturae per regna tri natu-

rae, secundum classes, ordines, genera, species, cum characteri-

bus, differentiis, synonymis, locis, 1. Leipzig.

G R A Y , J. E. 1821. On the natural arrangement of vertebrose

animals. London Medical Repository, 15, 296–310.

—— 1825. Outline of an attempt at the disposition of the Mam-

malia into tribes and families with a list of the genera appar-

ently appertaining to each tribe. Annales Philosophiae, New

Series, 10, 337–344.

H A R R I S O N , T. 1991. The implications of Oreopithecus bam-

bolii for the origin of bipedalism. 235–244. In C OP P E N S , Y.

and S E N UT , B. (eds). Origine(s) de la bipedie chez les homi-

nides. CNRS, Paris, 301 pp.

H UN E R M A N N , K. A. 1968. Die Suidae (Mammalia, Artio-

dactyla) aus den Dinotheriensanden (Unterpliozan + Pont)

A B B A Z Z I E T A L . : M I O C E N E V E R T E B R A T E A S S E M B L A G E F R O M S A R D I N I A 449

Rheinhessens (Sudwestdeutschland). Schweizerische Palaonto-

logische Abhandlungen, 86, 1–96.

H U R Z E L E R , J. 1951. Contribution a l’etude de la dentition de

lait d’Oreopithecus bambolii Gervais 1972. Eclogae Geologicae

Helvetiae, 44, 404–411.

—— 1958. Oreopithecus bambolii Gervais: a preliminary report.

Verhandlungen der Naturforschenden Gesellschaft in Basel, 69,

1–47.

—— 1982. Sur le suide du lignite de Montebamboli (prov. Gros-

seto, Italie). Comptes Rendus de l’Academie des Sciences de

Paris, Serie 2, 295, 697–701.

—— 1983. Un alcelaphine aberrant (Bovide, Mammalia) des lig-

nites de Grosseto en Toscane. Comptes Rendus de l’Academie

des Sciences de Paris, Serie 2, 296, 497–503.

—— 1987. Die Lutrinen (Carnivora, Mammalia) aus dem ‘‘Gros-

seto Lignit’’ der Toscana. Schweizerische Palaontologische

Abhandlungen, 110, 27–48.

—— and E N G E S S E R , B. 1976. Les faunes des mammiferes

neogenes du Bassin de Baccinello (Grosseto, Italie). Comptes

Rendus de l’Academie des Sciences de Paris, Serie 2, D, 283,

333–336.

ICZN 1999. International Code of Zoological Nomenclature.

Fourth edition. International Commission on Zoological

Nomenclature, International Trust for Zoological Nomencla-

ture, c ⁄ o The Natural History Museum, London, xxix + 306

pp.

J A N I S , C. 1995. Correlations between craniodental morphology

and feeding behaviour in ungulates: reciprocal illumination

between living and fossil taxa. 76–98. In T HO M A S ON , J.

(ed.). Functional morphology in vertebrate paleontology. Cam-

bridge University Press, Cambridge, 277 pp.

J A R M A N , P. J. 1974. The social organisation of antelope in

relation to their ecology. Behaviour, 48, 215–166.

K OT S A K I S , T., D E L F I N O , M. and P I R A S , P. 2004. Italian

Cenozoic crocodilians: taxa, timing and biogeographic impli-

cations. Palaeogeography, Palaeoclimatology, Palaeoecology, 210,

67–87.

K OH L E R , M. and M OY A - S OL A , S. 1997. Ape-like or homi-

nid-like? The positional behaviour of Oreopithecus bambolii

reconsidered. Proceedings of the National Academy of Sciences,

USA, 94, 11,747–11,750.

—— —— 2003. Understanding the enigmatic ape Oreopithecus

bambolii. Courier Forschungsinstitut Senckenberg, 243, 11–123.

K R E T ZO I , M. 1943. Kochictis centennii n. g. n. sp., ein alter-

tumlicher Creodonte aus dem beroligozan Siebenburgens.

Foldtani Kozlony, 73, 10–17, 190–195.

L I N N A E US , C. 1758. Systema Naturae per regna tria naturae

secundum classis, ordines, genera, species cum characteribus, dif-

ferentiis, synonymis, loci. Laurentii Salvii, Stockholm, 823 pp.

L O R E N Z , H. G. 1968. Stratigraphisches und mikropalaonto-

logische Untersuchungen des Braunkohlengebietes von Bacci-

nello (Grosseto, Italien). Rivista Italiana di Paleontologia e

Stratigrafia, 74, 147–170.

M A C A R TH U R , R. H. and W I L S ON , E. O. 1963. An equilib-

rium theory of insular zoogeography. Evolution, 17, 373–387.

M A DE , J. VAN DER 1999. Biogeography and stratigraphy of the

Mio-Pleistocene mammals of Sardinia and the description of

some fossils. 337–360. In R E U M E R , J. W. F. and D E V O S ,

J. (eds). Elephants have a snorkel! Papers in honour of Paul Y.

Sondaar. Deinsea, 7, 422 pp.

M A ZZ A , P. and R US T I ON I , M. 1997. Neotype and phylog-

eny of the suid Eumaiochoerus etruscus (Michelotti) from

Montebamboli (Grosseto, southern Tuscany). Paleontologia i

Evolucio, 30–31, 5–18.

M E I N , P. 1999. The Late Miocene small mammal succession

from France, with emphasis on the Rhone Valley localities.

140–164. In A G US T I , J., R O OK , L. and A N D R E W S , P.

(eds). Hominoid evolution and climatic change in Europe.

Volume 1. The evolution of Neogene terrestrial ecosystems in

Europe. Cambridge University Press, Cambridge, 512 pp.

M OR A L E S , J. and S O R I A , D. 1981. Los artiodactilos de los

Valles de Fuentiduena. Estudios Geologicos, 37, 477–501.

—— —— and T H O M A S , H. 1987. Les Giraffidae (Artiodac-

tyla, Mammalia) d’Al Jadidah du Miocene moyen de la For-

mation Hofuf (province du Hasa, Arabie Saoudite). Geobios,

20, 441–467.

M OY A - S OL A , S. and K O HL E R , M. 1997. The phylogenetic

relationships of Oreopithecus bambolii Gervais, 1872. Comptes

Rendus de l’Academie des Sciences de Paris, Serie 2, 324, 141–

148.

—— —— and R O O K, L. 1999. Evidence of hominid-like preci-

sion grip capabilities in the hand of the European Miocene

ape Oreopithecus. Proceedings of the National Academy of

Sciences, USA, 96, 313–317.

—— —— —— 2005. The Oreopithecus’ thumb: a strange case in

hominoid evolution. Journal of Human Evolution, 49, 395–

404.

O L S ON , E. C. 1952. The evolution of Permian vertebrate chro-

nofauna. Evolution, 6, 181–196.

O W E N , R. 1848. Contributions to the history of British fossil

mammals. Taylor, London, 71 pp.

P E R E Z - B A R B E R I A , F. J., G O R DO N , I. J. and P A GE L , M.

2002. The origins of sexual dimorphism in body size in ungu-

lates. Evolution, 56, 1276–1285.

R O C H E B R U N E , T. DE A. 1883. Faune de la Senegambie.

Mammiferes. Actes de la Societe Linneenne de Bordeaux, Serie

4, 37, 7, 49–203.

R O BI N S O N , P. 1986. Very hypsodont antelopes from the Be-

glia Formation (central Tunisia), with a discussion of the

Rupicaprini. Contributions to Geology, University of Wyoming,

Special Paper, 3, 305–315.

R O OK , L., A BB A Z Z I , L. and E N G E S S E R , B. 1999a. An

overview on the Italian Miocene land Mammal faunas. 191–

204. In A G U S T I , J., R OO K , L. and A N D R E W S , P. (eds).

Hominoid evolution and climatic change in Europe. Volume 1.

The evolution of Neogene terrestrial ecosystems in Europe. Cam-

bridge University Press, Cambridge, 512 pp.

—— B ON D I O L I , L., C A S A L I , F., R O S S I , M., KO H L E R ,

M., M O Y A - S O L A , S. and M A C CH I A R E L L I , R. 2004.

The bony labyrinth of Oreopithecus bambolii. Journal of

Human Evolution, 46, 347–354.

—— —— KO H L E R , M., M OY A - S O L A , S. and M A C CH -

I A R E L L I , R. 1999b. Oreopithecus was a bipedal ape after

all: evidence from the iliac cancellous architecture. Proceed-

ings of the National Academy of Sciences, USA, 96, 8795–

8799.

450 P A L A E O N T O L O G Y , V O L U M E 5 1

—— G A L LA I , G. and T O R R E , D. 2006. Lands and endemic

mammals in the Late Miocene of Italy: constraints for

paleogeographic outlines of Tyrrhenian area. Palaeogeography

Palaeoclimatology, Palaeoecology, 238, 263–269.

—— HA RR I S O N , T. and E N G E S S E R , B. 1996. The taxo-

nomic status and the biochronological implications of new

finds of Oreopithecus from Baccinello (Tuscany, Italy). Journal

of Human Evolution, 30, 3–27.

—— R E N N E , P., BE N V E N UT I , M. and PA P I N I , M.

2000. Geochronology of Oreopithecus-bearing succession at

Baccinello (Italy) and the extinction pattern of European

Miocene hominoids. Journal of Human Evolution, 39, 577–

582.

R U S T I O N I , M., M A Z ZA , P., A Z ZA RO L I , A., B O S CA -

G L I , G., CO Z Z I N I , F., D I V I T O, E., M A S S E T I , M. and

P I S A N O , A. 1992. Miocene vertebrate remains from Scont-

rone, National Park of Abruzzi, Central Italy. Rendiconti della

Accademia Nazionale dei Lincei, Classe di Scienze Fisiche Mate-

matiche e Naturali, Serie 9, 3, 227, 237.

S C L A T E R , P. L. and T H OM A S , O. 1894. The book of the

antelopes. Volume 1. R. H. Porter, London, ix + 216 pp.

S O N D A A R , P. Y. 1977. Insularity and its effects on mammal

evolution. 671–707. In H E CT , M. N., GO O DY , P. L. and

H E CT , B. M. (eds). Major patterns in vertebrate evolution.

Plenum Press, New York, NY, 422 pp.

—— 1987. Pleistocene mammals and extinctions of island en-

demics. Memoires de la Societe Geologique de France, Nouvelle

Serie, 150, 159–165.

T H OM A S , H. 1981. Les Bovides miocenes de la formation de

Ngorora du Bassin de Baringo (Rift Valley, Kenya). Proceedings

of the Koninklijke Nederlandse Akademie van Wetenschappen,

Series B, 84, 335–410.

—— 1984. Les origines africaines des Bovidae (Artiodactyla,

Mammalia) miocenes des lignites de Grosseto (Toscane, Italie).

Bulletin du Museum National d’Histoire Naturelle, Paris, Serie

4, C, 6, 81–101.

V R B A , E. S. 1997. New fossils of Alcelaphini and Caprinae

(Bovidae: Mammalia) from Awash, Ethiopia and phyloge-

netic analysis of Alcelaphini. Palaeontologia Africana, 34,

127–198.

W E I T HO F E R , K. A. 1888. Alcune osservazioni sulla fauna

delle ligniti di Casteani e di Montebamboli (Toscana). Bolletti-

no del Reale Comitato Geologico d’Italia, 19, 363–368.

—— 1889. Ueber die tertiaren Landsaugethiere Italiens. Jahrbuch

der Kaiserlich-Koniglichen Geologischen Reichsanstalt, 39,

55–82.

A B B A Z Z I E T A L . : M I O C E N E V E R T E B R A T E A S S E M B L A G E F R O M S A R D I N I A 451