Quaternary Science Reviews 87 (2014) 82e97

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Quaternary Science Reviews

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Acinonyx pardinensis (Carnivora, Felidae) from the Early Pleistocene of

Pantalla (Italy): predatory behavior and ecological role of the giant
PlioePleistocene cheetah
Marco Cherin a, *, Dawid Adam Iurino b, Raffaele Sardella b, Lorenzo Rook c
a
b
c

Dipartimento di Scienze della Terra, Perugia University, Piazza Universit, 06123 Perugia, Italy
Dipartimento di Scienze della Terra, Sapienza University of Rome, Piazzale A. Moro 5, 00185 Rome, Italy
Dipartimento di Scienze della Terra, Florence University, Via G. La Pira 4, 50121 Florence, Italy

a r t i c l e i n f o

a b s t r a c t

Article history:
Received 25 November 2013
Received in revised form
3 January 2014
Accepted 4 January 2014
Available online

The site of Pantalla (central Italy) yielded a rich late Villafranchian (Early Pleistocene) faunal assemblage,
which includes some well-preserved large mammal skulls. We describe here two nearly complete crania
and a left hemimandible of Acinonyx pardinensis from this locality, representing the most complete
cranial material of this species in Europe. These nds allowed us to dene more clearly the craniodental
morphology of A. pardinensis. Similarly to the forms from North Africa and China, the giant cheetah from
Pantalla has a more generalized skull than the living Acinonyx jubatus, showing some primitive,
pantherine-like features such as the less domed dorsal outline of the cranium, the more developed
sagittal and nuchal crests and the less bowed zygomatic arches. High-resolution CT scans of the specimens were used to obtain the rst 3D model of a cranium with articulated mandible of A. pardinensis.
Starting from the insertion areas on this model we reconstructed the jaw muscles of the Pantalla felid,
conrming its afnities with pantherine felines. In the light of the musculoskeletal skull anatomy and the
average body mass (about 80 kg), it is likely that A. pardinensis could kill large prey through a hunting
strategy more similar to pantherine cats than to the living cheetah.
2014 Elsevier Ltd. All rights reserved.

Keywords:
Acinonyx pardinensis
Giant cheetah
Hunting behavior
Jaw muscles
Pantalla
Villafranchian

1. Introduction: what we know about cheetah-like cats

The extant cheetah Acinonyx jubatus is the fastest creature in
terrestrial ecosystems, reaching 103 km/h in running (Sharp, 1997).
The cheetahs killing strategy is based on high speed: after a
noiseless and stealthy approach to the potential prey, the chase
normally begins with a gentle trot followed by an extraordinary
acceleration; the capture occurs at high speed, thanks to a precise
action of clawing at one side of the preys rear, causing its fatal fall
(Turner and Antn, 1997).
The cheetahs skeletal anatomy is highly specialized for this
peculiar hunting behavior (Hildebrand, 1985; Van Valkenburgh
et al., 1990; Turner and Antn, 1997): the body is very slender on
the whole, with a long lumbar region relative to the thoracic one,
thus increasing spinal dorsiexion during running; pelvic limb
bones are elongated, reecting increasing cursoriality; the long tail
is used as a rudder to counterbalance body weight during the
running; the skull is short and small, and teeth are reduced in size,

* Corresponding author. Tel.: 39 340 5123518.

E-mail addresses: marco.cherin@unipg.it, marco.cherin@gmail.com (M. Cherin).
0277-3791/$ e see front matter 2014 Elsevier Ltd. All rights reserved.
http://dx.doi.org/10.1016/j.quascirev.2014.01.004

determining a reduction of the head mass; respiratory passages are

enlarged thanks to an expansion of the posterior portion of the
frontals and an enlargement of nares, nasal passages and sinuses,
thus assuring a good oxygenation.
A similar body plan (with probably comparable hunting strategy) is displayed in at least two PlioePleistocene cheetah-like felids: the North American cheetah Miracinonyx, with the two species
Miracinonyx inexpectatus and Miracinonyx trumani (Van
Valkenburgh et al., 1990), and the Old World giant cheetah Acinonyx pardinensis (Ficcarelli, 1984).
The fossil record suggests that cheetahs originated in Africa
(Fig. 1). Werdelin and Dehghani (2011) reported some remains of
Acinonyx sp. from Laetoli (Tanzania), which are very similar in size
with the modern A. jubatus. These remains are 3.85e3.60 Ma in age,
representing the rst occurrence of the genus, together with those
from Sterkfontein Member 2 (South Africa), although the age of the
latter site is disputed (cfr. Werdelin, 2010, p. 31). Other SubSaharian African sites with cheetah fossils are reported in Table 1.
Since the second half of the XX Century, remains of cheetah-like
cats were described in a number of Eurasian and North African sites
(Fig. 1, Table 1), but their taxonomic attribution is debated.
Considering the scarcity of the fossil record, Hemmer et al. (2008,

M. Cherin et al. / Quaternary Science Reviews 87 (2014) 82e97

83

Fig. 1. Map of the PlioePleistocene records of cheetah-like cats in the Old World. 1, Villafranca dAsti IT; 2, Les touaires FR; 3, Villarroya ES; 4, Las Higueruelas ES; 5, Shamar MN; 6,
Beregovaya RU; 7, Yushe CN; 8, Tuozidong Cave CN; 9, Pantalla IT; 10, Casa Frata IT; 11, Montopoli IT; 12, Olivola IT; 13, Pirro Nord IT; 14, Saint Vallier FR; 15, La Cte dArd FR; 16,
Senze FR; 17, Fonelas P-1 ES; 18, La Puebla de Valverde ES; 19, Ahl Al Oughlam MA; 20, Varshets BG; 21, Khapry RU; 22, Dmanisi, GE; 23, Kuruksay TJ; 24, Siwalik Hills IND; 25,
Longdan CN; 26, Nihewan CN; 27, Untermassfeld DE; 28, Le Vallonnet FR; 29, Yuanqu Loc. 105 CN; 30, Gongwangling CN; 31, Longgudong Cave CN; 32, Liucheng Gigantopithecus
Cave CN; 33, Hundsheim AT; 34, Mosbach DE; 35, Saint Estve FR; 36, Zhoukoudian Locs. 1, 13 CN. The unpublished material of Sivapanthera sp. from Yushe (7) was preliminarily
referred to the Middle Pliocene (Qiu, 2006), allowing a possible late Ruscinian rather than early Villafranchian biochronological attribution of this locality. The site of Tuozidong Cave
(8) is actually late Villafranchian in age (w2.0 Ma), but it bears remains referred to A. p. arvernensis (Dong et al., 2013), suggesting that this primitive form probably survived in the
Far East longer than in Europe (Hemmer et al., 2011). The position of the marker no. 24 has to be considered approximate, as we do not know the position of the exact locality of
discovery of the cheetah-like remains within the Siwalik Hills. (Online version in color.)

2011) propose to consider A. pardinensis as a broad macrospecies,

represented by several subspecies in Eurasia and North Africa
during the PlioePleistocene (Table 1): A. p. arvernensis, for the remains from touaires dated to the Pliocene to Pleistocene transition
(Felis arvernensis sensu Schaub, 1949), and from the younger site of
Tuozidong (China; Dong et al., 2013); A. p. aicha, for the earliest
Pleistocene cheetah from North Africa (Acinonyx aicha sensu
Geraads, 1997); A. p. pardinensis, for a number of Early Pleistocene
sites from Europe to Central Asia; A. p. linxiaensis, for the northern
Chinese forms previously referred to Sivapanthera linxiaensis (Qiu
et al., 2004); A. p. pleistocaenicus, for the late Early Pleistocene
(latest Villafranchian, 1.05 Ma) large-sized cheetah from Untermassfeld and the older Chinese Sivapanthera pleistocaenica (Cynaiulurus pleistocaenicus sensu Zdansky, 1925); A. p. intermedius, for
the Galerian (i.e., Middle Pleistocene) specimens from Hundsheim
and Mosbach (Acinonyx intermedius sensu Thenius, 1954).
In addition, Hemmer et al. (2011) claried that there was a longlasting mistake in the identication of the type locality of
A. pardinensis in the last 50 years. In fact, the type mandible was
attributed to the French site of touaires, together with the type
mandible of the larger Felis arvernesis (Schaub, 1949), which is an
older synonym of A. pardinensis according to Viret (1954). However,
the former mandible was found at La Cte dArd, which is not far
from the touaires classic site, but stratigraphically higher. As a
consequence, A. pardinensis was found in two different sites of the
Puy de Dme, La Cte dArd and touaires, the former being the
type locality.

Evolutionary relationships among all these forms, as well as

between Old and New World forms, are still cloudy. At least two
taxa were suggested to be the possible ancestor of cheetahs: the
puma-like cat Puma pardoides from Eurasia (Kurtn, 1976) and
Panthera crassidens from Africa (Petter and Howell, 1976). The latter
however should not be considered because the holotype was
demonstrated to be a composite specimen (Turner, 1990). The
American cheetah M. inexpectatus probably reached North America
in the Late Pliocene and could be ancestral to both M. trumani and
the extant cougar Puma concolor (Van Valkenburgh et al., 1990).
Miracinonyx inexpectatus and A. pardinensis seem to appear
approximately at the same age. Although the origin of A. pardinensis
is still unclear, Hemmer et al. (2011) supposed that European and
Asian forms followed a separate evolutionary history during the
Early Pleistocene: while in the West A. p. pardinensis was probably a
chronological successor of A. p. arvernensis during the early to
middle Villafranchian transition, in the East the primitive A. p. linxiaensis was substituted by A. p. pleistocaenicus, which invaded
Europe in conjunction with the climatic aridication at the beginning of the latest Villafranchian (Spassov, 2011).
All the above taxonomic, phylogenetic and paleobiogeographic
points about cheetah-like fossil cats, and A. pardinensis in particular, are still unsolved due to the scantiness of the fossil record.
Most of the European sites yielded fragmentary specimens, mainly
represented by mandibular and postcranial material. Sufciently
complete A. pardinensis crania are known from only three sites:
Montopoli (Ficcarelli, 1984), Untermassfeld (Hemmer, 2001) and

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M. Cherin et al. / Quaternary Science Reviews 87 (2014) 82e97

Table 1
List of the Old World localities bearing PlioePleistocene remains of cheetah-like cats. Numbers in the third column refer to Fig. 1.
Continent

State

No. Locality

Original determination

Revised determination

Age

References

Africa

S Africa

Acinonyx sp.

Tanzania

Acinonyx sp.

Ethiopia

cf. Acinonyx

S Africa

Acinonyx sp.

Morocco

19

Sterkfontein
Member 2
Laetoli Beds
Upper Unit
Omo Shungura
C-Usno
Makapansgat
Member 3
Ahl Al Oughlam

Acinonyx aicha

A. pardinensis aicha (type)

Werdelin and
Peign (2010)
Werdelin and
Dehghani (2011)
Werdelin and
Peign (2010)
Werdelin and
Peign (2010)
Geraads (1997)

Kenya

Acinonyx sp.

S Africa

Acinonyx jubatus

S Africa

Acinonyx jubatus

Tanzania

Koobi Fora Upper

Burgi-KBS-Okote
Swartkrans
Member 1
Swartkrans
Members 2e3
Olduvai Bed 2

Acinonyx cf. jubatus

Ethiopia

Awash 7

Acinonyx sp.

Middle Pliocene
(ca 4.0e3.0 Ma)a
Middle Pliocene
(3.85e3.60 Ma)
Late Pliocene
(ca 3.3e3.0 Ma)
Late Pliocene
(3.2e2.5 Ma)
early Early Pleistocene
(ca 2.5 Ma)
Early Pleistocene
(2.0e1.4 Ma)
Early Pleistocene
(ca 1.9e1.8 Ma)
late Early Pleistocene
(?ca 1.6 Ma)
late Early Pleistocene
(ca 1.4 Ma)
Early Pleistocene

Morocco

Doukkala 1e2

Acinonyx jubatus

MiddleeLate Pleistocene
?Middle Pliocene
Late Plioceneeearliest
Pleistocene
Late Plioceneeearliest
Pleistocene
early Early Pleistocene
(2.58e2.25 Ma)
Early Pleistocene
(2.5e1.7 Ma)
early Early Pleistocene
(MN17)
early Early Pleistocene
(ca 2.0 Ma)
Early Pleistocene
(1.9e1.6 Ma)

Asia

Europe

Europe

China
Mongolia

7
5

Yushe
Shamar

Sivapanthera sp.
Acinonyx sp.

Sivapanthera sp.
e

E Russia

Beregovaya

Acinonyx sp.

China

25

Longdan

Sivapanthera linxiaensis

India

24

Felis brachygnathus

Tajikistan

23

Siwalik Hills,
Pinjor Stage
Kuruksay

A. pardinensis linxiaensis
(type)
Acinonyx pardinensisb

Acinonyx pamiroalayensis

A. pardinensis pardinensis

Tuozidong Cave

Acinonyx arvernensis

A. pardinensis arvernensis

Werdelin and
Peign (2010)
de Ruiter (2003)
de Ruiter (2003)
Turner (1990)
Werdelin and
Peign (2010)
Werdelin and
Peign (2010)
Qiu (2006)
Sotnikova (1978)
Sotnikova (1978)
Qiu et al. (2004)
Lydekker (1884);
Spassov (2011)
Sharapov (1986);
Hemmer et al. (2011)
Hemmer et al. (2011);
Dong et al. (2013)
Teilhard de Chardin
and Piveteau (1930);
Qiu et al. (2004)
Zdansky (1925);
Qiu (2006); Spassov
(2011)
Qiu (2006)

China

China

26

Nihewan

Cynailurus pleistocaenica

Acinonyx sp. (archaic form)

China

29

Yuanqu Loc. 105

Cynailurus pleistocaenica

A. pardinensis pleistocaenicus
(type)

late Early Pleistocene

(1.6e1.2 Ma)

China

30

Gongwangling

Sivapanthera pleistocaenica

A. pardinensis pleistocaenicus

China
China

31
32

Sivapanthera pleistocaenica
Sivapanthera pleistocaenica

A. pardinensis pleistocaenicus
A. pardinensis pleistocaenicus

China

36

Longgudong Cave
Liucheng
Gigantopithecus Cave
Zhoukoudian
Locs. 1, 13
Villafranca dAsti

late Early Pleistocene

(1.6e1.2 Ma)
Early Pleistocene
Early Pleistocene

Acinonyx sp.

early Middle Pleistocene

Acinonyx pardinensis

Late Pliocene (Triversa FU) Kurtn and Crusafont

Pair (1977)
Late Plioceneeearliest
Schaub (1949);
Pleistocene (MN16)
Viret (1954);
Hemmer et al. (2011)
Late Plioceneeearliest
Villalta (1952)
Pleistocene (MN16)
Late Plioceneeearliest
Villalta (1952)
Pleistocene (MN16)
Late Plioceneeearliest
Ficcarelli (1984)
Pleistocene
(Montopoli FU)
early Early Pleistocene
Schaub (1949);
Viret (1954); Hemmer
et al. (2011)
early Early Pleistocene
Viret (1954);
(MN17)
Argant (2004);
Spassov (2011)
early Early Pleistocene
Spassov (2011)
(MN17)
early Early Pleistocene
Sotnikova et al. (2002);
(MN17)
Hemmer et al. (2011)
early Early Pleistocene
Schaub (1942);
(MN18)
Delson et al. (2006)
early Early Pleistocene
Garrido (2008)
(MN18)

Italy

France

Les touaires,
Puy de Dme

Acinonyx pardinensis

A. pardinensis arvernensis
(type)

Spain

Villarroya

Acinonyx pardinensis

Spain

Las Higuerelas

Acinonyx pardinensis

Italy

11

Montopoli,
Lower Valdarno

Acinonyx pardinensis

France

15

La Cte dArd,
Puy de Dme

Acinonyx pardinensis

A. pardinensis pardinensis
(type)

France

14

Saint Vallier

Acinonyx pardinensis

A. pardinensis pardinensis

Bulgaria

20

Varshets

Acinonyx pardinensis

A. pardinensis pardinensis

W Russia

21

Khapry

Acinonyx pardinensis

A. pardinensis pardinensis

France

16

Senze

Acinonyx pardinensis

Spain

17

Fonelas P-1

Acinonyx pardinensis

Spain

18

Acinonyx pardinensis

Hou and Zhao (2010)

Tong and Gurin (2009)
Pei (1934)

M. Cherin et al. / Quaternary Science Reviews 87 (2014) 82e97

85

Table 1 (continued )
Continent

State

No. Locality

Original determination

Revised determination

La Puebla de
Valverde
Italy

10

Georgia

22

Casa Frata,
Upper Valdarno
Dmanisi

Acinonyx pardinensis

A. pardinensis pardinensis

A. pardinensis ?pardinensis

Italy

Pantalla

Acinonyx pardinensis

Italy

12

Olivola

Acinonyx pardinensis

A. pardinensis pardinensis

Italy

13

Pirro Nord

Acinonyx pardinensis

Germany

27

Untermassfeld

A. pardinensis pleistocaenicus e

France

28

Le Vallonnet

Acinonyx pardinensis

Austria

33

Hundsheim

Acinonyx intermedius

Germany

34

Mosbach

Acinonyx intermedius

A. pardinensis intermedius
(type)
A. pardinensis intermedius

France

35

Saint Estve

Acinonyx pardinensis

Age

References

early Early Pleistocene

(ca 2.04 Ma)

Kurtn and
Crusafont Pair (1977);
Alcal et al. (1989e1990)
Ficcarelli (1984);
Spassov (2011)
Hemmer et al. (2011)

late Early Pleistocene

(Tasso FU)
early Early Pleistocene
(ca 1.8 Ma)
late Early Pleistocene
(OlivolaeTasso FU)
late Early Pleistocene
(Olivola FU)
late Early Pleistocene
(Pirro FU)
late Early Pleistocene
(ca 1.05 Ma)
late Early Pleistocene
(0.98e0.91 Ma)
Middle Pleistocene
Middle Pleistocene
Middle Pleistocene
(MN22)

This paper
Ficcarelli (1984);
Spassov (2011)
Petrucci et al. (2013)
Hemmer (2001)
de Lumley et al. (1988)
Thenius (1954);
Hemmer et al. (2008)
Schtt (1970);
Hemmer et al. (2008)
Bonifay and
Bonifay (1963)

The age of the Sterkfontein site is a topic of debate. See Werdelin (2010) for a synthesis of the different hypotheses.
In the light of our re-analysis of the Siwalik material (hemimandibles NHM 16537 and NHM 16576), we agree with Spassov (2011) in attributing the specimens to Acinonyx
pardinensis.
b

Saint Vallier (Viret, 1954). Nevertheless, the cranium IGF 12477

from Montopoli is so heavily deformed and fragmented that only
the upper teeth can be fully described. The specimen IQW 1980/
16350 from Untermassfeld is well preserved only in the ventral
area. The complete skull with associated mandible MHNL QSV.112
from Saint Vallier gured and described by Viret (1954) was heavily
reconstructed, especially in the posterior portion, while a second
cranium (MHNL QSV.115) is crushed and unreconstructed (Geraads,
2008).
In this scenario the new sample from the site of Pantalla
(Perugia, Italy; Gentili et al., 1997) is of crucial importance because
it is the most complete available cranial material of A. pardinensis in
Europe and offers a unique opportunity to deepen our knowledge
on this carnivores cranial anatomy and function. The rst detailed
description of A. pardinensis from Pantalla is offered here. In addition, some inferences on the predatory behavior of this species are
discussed starting from the analysis of the masticatory system.

concentrated in a very small area (about 2 m2); the upper level was
composed by silty clays interpreted as a drained paleosol, where
fragmented postcranial bones of herbivores and few micromammal
teeth were found.

2. Geological framework and biochronology of the Pantalla

assemblage
The Early Pleistocene locality of Pantalla (about 30 km S to
Perugia, Italy; 42 520 46.7900 N, 12 240 23.2600 E) yielded abundant
remains of late Villafranchian continental mammals, mostly represented by well-preserved skulls.
The site is located near the western margin of the Apennine
Chain, in the southwestern branch of the Tiber Basin, a wide
extensional continental basin with a characteristic upside down Y
shape (Fig. 2). This basin was lled by clastic (lacustrine, palustrine
and uvial) and, secondly, carbonate deposits during a time span
ranging from the early Late Pliocene until the early Late Pleistocene
(Basilici, 1997).
The Pantalla mammal fauna (Gentili et al., 1997) was recovered
in 1995 from a 15 m-thick stratigraphic succession referred to the
Early Pleistocene Santa Maria di Ciciliano Unit. In particular, fossil
bones were discovered within two levels: the lower level was
composed by uvial silty sands interpreted as crevasse-splay deposits and yielded abundant herbivore and carnivore remains

Fig. 2. Location of the paleontological site of Pantalla (Italy). The Tiber Basin is highlighted in light grey along the middle part of the region Umbria.

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M. Cherin et al. / Quaternary Science Reviews 87 (2014) 82e97

The mammal fauna is represented by the following taxa:

Apodemus cf. A. dominans, Canis etruscus, Vulpes sp., Lynx issiodorensis valdarnensis, Acinonyx pardinensis, Lutra sp., Sus cf.
S. strozzii, Pseudodama nestii, Leptobos aff. L. furtivus, Equus sp.,
Mammuthus cf. M. meridionalis (Gentili et al., 1997; Cherin et al.,
2013a,b).
From a biochronological point of view, the assemblage from
Pantalla can be referred to the Olivola/Tasso Faunal Unit (Gentili
et al., 1997), in a time span approximately ranging 1.8e1.7 Ma
(Rook and Martnez-Navarro, 2010).

3. Materials and methods

The sample of A. pardinensis from Pantalla consists of two
mostly complete crania and a left hemimandible (Fig. 3).
For the purpose of comparison we studied also the A. pardinensis
sample from Montopoli (Italy) and we referred to bibliographic
data about A. pardinensis from the Eurasian and African sites listed
in Table 1. Cranial and mandibular remains of Panthera gombaszoegensis (P. toscana) from the Upper Valdarno (Italy) and the type
locality of Gombaszog (Slovakia) were also analyzed. Additional

Fig. 3. Acinonyx pardinensis from Pantalla (Italy). aed) Cranium SBAU 337624 in dorsal (a), ventral (b), left lateral (c) and rostral (d) view. eei) Cranium SBAU 337648 in dorsal (e),
ventral (f), left lateral (g) with close-up of the upper cheek teeth (h) and rostral (i) view. jem) Left hemimandible SBAU 337627 in lingual (j), labial (k) and occlusal (l) view with
close-up of the lower teeth (m). Scale bars 5 cm.

M. Cherin et al. / Quaternary Science Reviews 87 (2014) 82e97

data on P. gombaszoegensis were taken from Koufos (1992) and

Hank (2007). Morphometric data about A. p. aicha are from
Geraads (1997), while those about Puma pardoides are from Argant
(2004), Hemmer et al. (2004) and Cherin et al. (2013c). Our
comparative dataset was completed by morphometric data
collected on the following extant felids stored in various European
museum and institutions (the number of analyzed specimens is in
brackets): A. jubatus (14), Panthera onca (21), Panthera pardus (17),
Panthera uncia (9), Panthera leo (14), Panthera tigris (18), Puma
concolor (3). Additional morphometric data were taken from the
literature also for extant species (references given in the text and
tables). Cranial, mandibular and dental measurements were
recorded to the nearest 0.1 mm with a digital caliper.
The Pantalla specimens were also scanned through a Philips
Brilliance CT 64-channel scanner in order to analyze the internal
anatomy, to virtually correct the diagenetic deformation and to
model virtual 3D reconstructions. Tomographic images were made
at the M.G. Vannini Hospital of Rome (Italy). The specimens were
scanned in the coronal (i.e., transverse) plane from rostral to caudal.
The scanning resulted in 436 slices (i.e., images) for SBAU 337624,
297 slices for SBAU 337648, 226 slices for SBAU 337627, 218 slices
for MCZR 367 (skull of the extant A. jubatus), 279 slices for MCZR
258 (skull of the extant P. onca) and 308 slices for IGF 13577 (cast of
the complete A. pardinensis skull from the French locality of Saint
Vallier), with images size of 512  512 pixels. The slices were
0.8 mm thick with an interslice space (i.e., the space between
consecutive slices) of 0.4 mm. Segmentation, 3D rendering and
elaboration of the fossil and modern materials were computed
using Osirix 3.9.4 32-bits for Mac, an Open-Source image processing software dedicated to DICOM les, and the 3D Open-Source
software Blender 2.63a Intel 32 bits for Mac.
Blender was also used to create the jaw muscles of SBAU 337624,
MCZR 367 and MCZR 258 in order to perform a comparative analysis of
the masticatory system. The 3D images of the specimens were
exported from OsiriX as .obj les and successively imported in
Blender. The reconstruction process of the masticatory muscles
included two different phases: (1) creation and modeling of the 3D
objects; (2) enrichment of surface details and insertion of color effects.
Each muscle was rst created as a Blender new mesh (3D object) with
a low number of polygons in order to improve the computer performance. Afterward we started the modeling process following the
muscle insertion areas clearly evident on the skulls (Antn et al.,
2009). The volume of the muscle mass was reconstructed taking
into account the spatial distance between the temporal surface and
the zygomatic arch for the temporal muscle and the distance between
the external surface of the zygomatic arch and the horizontal branch of
the mandible for the masseter muscle. In order to confer a more
realistic aspect to the 3D models, the number of polygons was then
increased to improve the graphic details of the muscles, and additional
artistic works, as color, surface and light effects were performed.
Institutional abbreviations e HMV, Hezheng Paleozoological
Museum (China); HNHM, Hungarian Natural History Museum and
Geological Institute of Hungary, Budapest (Hungary); ICP, Museo
del Institut Catal de Paleontologia Miquel Crusafont, Sabadell
(Spain); IGF, Museo di Storia Naturale, Sezione di Geologia e Paleontologia, Florence University (Italy); IGME, Museo Geominero,
Instituto Geolgico y Minero de Espaa, Madrid (Spain); INSAP,
Institut National des Sciences de lArchologie et du Patrimoine du
Royaume du Maroc, Casablanca (Morocco); IQW, Senckenberg
Research Institute, Quaternary Large Mammal Section, Weimar
(Germany); IVPP, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Nanjing (China);
MCZR, Museo Civico di Zoologia, Rome (Italy); MHNB, Musum
dHistoire Naturelle, Basel (Switzerland); MHNL, Musum dHistoire Naturelle, Lyon (France); MHNP, Musum national dHistoire

87

naturelle, Paris (France); SMF, Senckenberg Naturmuseum Frankfurt (Germany); NHM, Natural History Museum, London (UK);
SBAU, Soprintendenza per i Beni Archeologici dellUmbria, Perugia
(Italy); SPE, Museo di Storia Naturale, Sezione di Zoologia La Specola, Florence University (Italy).
4. Analytical study
4.1. CT scanning and 3D virtual modeling
The good state of preservation of the fossils and the features of
the sediment in which they laid (uvial silty sands) allowed to
obtain high quality two- and three-dimensional tomographic images, favoring an easy discrimination of materials with different
density. The boundaries among fossilized bone, tooth enamel and
sandy sediment are therefore visible in the virtual slices of the
specimens (Fig. 4).
In the case of SBAU 337624 this type of investigation allowed to
determine the non-pathological nature of the swelling on the right
maxilla (Fig. 4). CT images clearly show a line of discontinuity between the wall of the maxillary bone and the swelling, which has
different density (Fig. 4a). Therefore, what from the outside
appeared to be a bony excrescence, proved to be a splinter of bone,
connected to the cranium by a piece of unremoved sediment. Using
the density lters available on Osirix, it was possible to verify the
excellent preservation of the right inner ear of SBAU 337624 (Fig. 5).
At present the study of this important anatomical structure is in
progress and may help to explore interesting aspects of the hearing
capability of these felids. Furthermore, within both the crania very
dense mineral veins are evident, developing from rostral to caudal
(Fig. 5). These structures arose during the fossilization process, with
precipitation of minerals due to water percolation.
Inner CT sections of the two crania and the hemimandible SBAU
337627 allowed us to have a look at the root morphology (Fig. 5).
The analysis of the whole tooth structures made possible to exclude
any dental anomaly, presence of supernumerary roots, intra-vitam
fractures or diseases (Iurino et al., 2013b).
3D computer graphics was used with the aim of minimizing the
deformation of the two crania that took place during the fossilization process (the hemimandible is virtually undeformed) (Iurino
et al., 2013a). The process of remodeling of the fossils required as
rst step to identify and virtually track the sagittal plane of the
crania (Fig. 6b and k). In this way the symmetry plane of each
cranium was created. Secondly, the mesh (3D object) was locked
along the whole palatal area, which is the least affected by the
deformation. The term locked in this case means that each option
was disabled by virtual movements of the highlighted object. It was
thus possible to shift the remaining portion of the cranium
following the symmetry plane (on the right in the case of SBAU
337624 and on the left in the case of SBAU 337648) to compensate
for the diagenetic deformation. During the operations of remodeling of SBAU 337624 the foreign swelling on the right maxilla was
also deleted. This series of operations allowed obtaining reliable
models of the crania, free from deformations and other anomalies
due to fossilization (Fig. 6k).
The computer graphic technology was also used for reconstructing the whole jaw starting from the left hemimandible SBAU
337627. Firstly the left hemimandible was cloned and mirrored,
obtaining the right hemimandible. The two hemimandibles were
then hinged together along the mandibular symphysis and in order
to improve the angle between the two portions, they were articulated to SBAU 337624, which is the cranium with almost complete
dentition and complete zygomatic arch. Simulations of the closing
and opening of the jaw were nally performed, verifying the correct dental occlusion (Fig. 6men).

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M. Cherin et al. / Quaternary Science Reviews 87 (2014) 82e97

Fig. 5. Acinonyx pardinensis from Pantalla (Italy). 3D models of the cranium SBAU
337624 and some of its inner structures. a) External surface of the cranium in right
lateral view. bec) View in transparency (b) and 3D models (c) of the inner structures:
teeth, inner ear (ie) and mineral veins (mv) developed during the diagenesis. Scale bar
3 cm. (Online version in color.)

Fig. 4. Acinonyx pardinensis from Pantalla (Italy). The coronal section (a) of the cranium
SBAU 337624 allows observing the inner structure of the specimen, in particular the
boundaries among fossilized bone (fb), tooth enamel (te), sandy sediment (se) and
dense mineral veins (mv). The black arrows indicate the line of discontinuity between
the wall of the maxillary bone and the swelling. b) The external look of the swelling on
the right maxilla on the 3D model. c) Close-up of the swelling consisting of a splinter of
bone (sb) connected to the cranium by a piece of unremoved sediment (se). Scale bars
3 cm (a) and 1 cm (bec). (Online version in color.)

The 3D visualization of the scanned specimens (MCZR 367, IGF

13577, SBAU 337648 and MCZR 258) accelerated and facilitated the
comparative analysis (Fig. 7). The possibility to turn the digitalized
skulls in the three directions of the space proved to be a very useful
tool to detect a wide range of morphological features, much more
than a classic comparison of 2D images (Iurino et al., 2013a).
4.2. Anatomical description
4.2.1. Cranium SBAU 337624
The cranium (Figs. 3aed and 6aef, Table 2) shows a very good
state of preservation, although it was partially deformed during the

diagenesis. In rostral view, it is evident a plastic translation on the

right side, especially in the upper portion. As a consequence, the
middle and rostral parts of the nasals were slightly crushed into the
nasal cavities. Besides this distortion, the cranium is almost complete, although the right zygomatic arch and the jugular and mastoid processes on both sides are fragmented. Unfortunately, sutures
are not more visible on the bone surfaces. Moreover, the inlling
sandy sediment covering the body of the basisphenoid and the
choanae has not been mechanically removed for not endangering
the specimens integrity.
In lateral view, the cranium appears quite domed and rostrocaudally compressed. The orbit is oriented rostrally and quite
large. Well-developed zygomatic process of the frontal and frontal
process of the zygomatic delimit the orbit respectively caudodorsally and caudoventrally; a little process is also present on the
rostral margin of the orbit, just above the lacrimal pit. A single, oval
and quite small infraorbital foramen opens rostroventrally to the
orbit. The zygomatic arch is quite slender and dorsoventrally
arched. The braincase is wide respect to total length, mostly thanks
to the enlargement of the caudal portion of frontal bones. A quite
tall sagittal crest crosses all the neurocranium till the akrokranion,
where it reaches a well-developed nuchal crest.
In ventral view, the palate appears large and at; two elongated
palatine ssures are visible rostrally to the incisors. The nasal spine
and the pterygoid hamuli are broken. The tympanic bullae are quite
large and elliptical in shape; they are separated by a wide

M. Cherin et al. / Quaternary Science Reviews 87 (2014) 82e97

89

Fig. 6. Acinonyx pardinensis from Pantalla (Italy). aef) 3D models of the cranium SBAU 337624 in caudal (a), rostral (b), left lateral (c), right lateral (d), dorsal (e) and ventral (f) view.
gej) 3D models of the left hemimandible SBAU 337627 in labial (g), lingual (h), occlusal (i) and ventral (j) view. k) Cranium SBAU 337624 after the retrodeformation process. The
dotted line indicates the sagittal symmetry plane. The swelling on the right maxilla (sw) was digitally removed. l) Virtual reconstruction of the whole skull of A. pardinensis from
Pantalla, realized by articulating the cranium SBAU 337624, the left hemimandible SBAU 337627 and the cloned and mirrored right hemimandible (in brown). men) Reconstructed
skull in left lateral view with the jaw open (m) and closed (n). Scale bars 5 cm. (Online version in color.)

basioccipital, which is characterized by the presence of a sharp

crest in the middle. Unfortunately, no foramina are clearly visible in
the temporal region because of their obliteration by the inlling
sediment.
Even if the nasal cavities were slightly deformed and narrowed
during the diagenesis, they look large in rostral view.
All the upper teeth are present except for both M1. Incisors are
robust and progressively larger from I1 to I3. A short diastema
separates the incisors from the canines, which appear small and

slender. Cheek teeth are very close one another. The P2 is very small
and circular in occlusal view. It is characterized by a thin cutting
ridge running from the mesiolingual to the distolabial margin. The
P3 is quite narrow and high-crowned. The protocone is very small
and located in lingual position, while the paracone is high and
directed vertically; the hypocone is well developed and the distal
cingulum is as well isolated as appearing an accessory cusp. The P4
is a robust tooth characterized by the strong reduction of the protocone, which appears as a simple swelling of the mesiolingual part

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M. Cherin et al. / Quaternary Science Reviews 87 (2014) 82e97

Fig. 7. Comparative cranial morphology of Acinonyx jubatus MCZR 367 (a1ea4), Acinonyx pardinensis IGF 13577 from Saint Vallier (b1eb4), Acinonyx pardinensis SBAU 337624 from
Pantalla (c1ec4) and Panthera onca MCZR 258 (d1ed4). The 3D models are shown in left lateral (a1ed1), dorsal (a2ed2), ventral (a3ed3) and rostral (a4ed4) view. The images are
normalized. (Online version in color.)

of the crown base, between the parastyle and the paracone. The
parastyle is strong and mesiolabially enlarged at the base. The
protocone and metacone are similar in size, with the former slightly
higher than the latter; in labial view, the protocone is directed
vertically as the paracone of P3.
4.2.2. Cranium SBAU 337648
The specimen is fully similar to SBAU 337624, both morphologically and morphometrically (Fig. 3eei, Table 2). The cranium
shows a smaller degree of diagenetic distortion but is more
incomplete: both zygomatic arches are broken and most of the
occipital region and the left tympanic bulla are missing. Thanks to
the lower dorsoventral deformation, the cranium is more domed
than SBAU 337624. The upper dentition lacks the right I3 and both
the P2, although the root of the left P2 is visible in the CT scans.
Unlike SBAU 337624, the right M1 is still present. In occlusal view, it
is elliptical in shape, with the main axis perpendicular to the tooth
row; a thin cutting ridge runs through the crown.
4.2.3. Hemimandible SBAU 337627
The left hemimandible (Figs. 3jem and 6gej, Table 3) is practically complete, with the exception of all the incisors. It is short
and slender on the whole. The corpus is straight both in lateral and
dorsal view. The coronoid process is quite low and vertical and the
angular process is very short, as it does not reach the caudal
margin of the condylar process. In occlusal view, the long axis of
the condyle is inclined respect to the corpus with an angle of about
45 . The symphysis is leaf-shaped and very short. The rostroventral margin of the hemimandible is quite squared.

Unfortunately, no mental foramina are clearly visible on the lateral

surface of the corpus.
The lower canine is short and slender. The crown of the P3 is
almost as tall as the one of P4. The diastema is short. In labial view,
the premolar teeth show a eur-de-lis morphology, with the
paraconid and the hypoconid diverging respectively mesially and
distally from the straight and high protoconid. Both the P3 and P4
have a well-developed cingulum distally to the hypoconid. The
relative length of the P3 is quite small if compared to the M1. In
occlusal view, postcanine teeth are so close one another to appear
partially overlapped.
4.3. Comparisons and discussion
From an anatomical point of view, modern cheetahs are clearly
different from other large felids, especially considering the skull
morphology, which retains a number of features that normally
distinguish small cats (ORegan, 2002).
The felid from Pantalla shows a large set of characters that
allowed us to refer it to the cheetah group. At the same time
through our comparisons we recognized some important differences between the Pantalla cat and the extant A. jubatus, which are
discussed in the Section 5 from a functional and ecological point of
view. In order to highlight the main anatomical similarities and
differences between our sample and some extinct and extant taxa
we produced Fig. 7, where the 3D model of the cranium SBAU
337624 is compared to those of A. pardinensis from Saint Vallier and
the modern A. jubatus and P. onca. The latter species was selected
within the pantherine clade (including the genera Panthera and

M. Cherin et al. / Quaternary Science Reviews 87 (2014) 82e97

Table 2
Measurements (mm) of the cranium and upper teeth of Acinonyx pardinensis from
Pantalla (Italy).
Cranium
Condylobasal length
Basal length
Viscerocranium length (NasioneProsthion)
Basicranial axis (BasioneSynsphenion)
Basifacial axis (SynsphenioneProsthion)
Zygomatic breadth
Neurocranium breadth
Breadth at the canine alveoli
Max palatal breadth (at the P4 alveoli)
Mastoid breadth
Breadth of the occipital condyles
Breadth of the foramen magnum
Max diameter of the tympanic bulla
Min diameter of the tympanic bulla
I1
Length
Breadth
2
I
Length
Breadth
I3
Length
Breadth
max
C
Length
Breadth
Crown height
P2
Length
Breadth
3
P
Length
Max breadth
Mesial breadth
Paracone length
Paracone height
P4
Length
Length at the protocone
Max breadth
Breadth behind the protocone
Max distal breadth
Paracone length
Metacone length
Paracone Metacone length
Paracone height
1
M
Length
Breadth

SBAU
337624

SBAU
337648

190.0
175.5
108.3
ca 48
ca 132
ca 150
67.7
56.4
84.2
77.4
44.6
ca 20
31.0
21.0
r
4.3
3.4
r
5.6
4.5
r
7.3
5.2
r
15.1
11.4
28.1
r
6.1
4.6
r
17.9
8.9
7.6
8.8
11.4
r
29.2
26.9
12.9
10.5
10.7
11.2
12.8
22.9
14.4
r
e
e

e
e
117.4
ca 50
ca 135
e
64.3
51.7
82.1
ca 73
e
e
31.3
14.4
r
4.4
3.2
r
5.3
3.8
r
e
e
r
15.5
11.5
e
r
e
e
r
18.1
8.5
7.7
7.5
12.0
r
28.5
26.3
14.4
9.7
9.7
11.9
12.0
22.7
15.7
r
4.4
10.0

l
4.3
3.2
l
5.4
4.1
l
7.3
5.8
l
15.0
11.4
e
l
5.8
4.7
l
19.0
9.0
8.1
8.8
12.3
l
29.9
27.2
13.1
10.6
10.8
12.2
12.7
24.2
14.6
l
e
e

l
4.4
3.3
l
5.2
4.2
l
6.9
5.1
l
14.8
11.9
23.7
l
e
e
l
17.2
8.6
7.6
7.5
12.2
l
28.3
25.8
13.0
9.7
9.8
11.4
12.0
22.8
10.0
l
e
e

Neofelis; Johnson et al., 2006) being characterized by a quite

generalized cranium and a mean body size comparable to that of
the extinct A. pardinensis.
From a lateral view the cranium from Pantalla is quite domed and
rostrocaudally compressed (Fig. 7c1). This character normally occurs
in the extant and extinct cheetah-like cats (Spassov, 2011), and
differs from the Panthera lineage, whose members show more
elongated crania. The only exception is represented by the snow
leopard P. uncia, which is the only member of its clade having
a rostrocaudally shortened cranium, although the dorsal outline is
much less domed than in A. jubatus. Recent genetic studies (Johnson
et al., 2006) exclude any phylogenetic relation between P. uncia and
the other short-cranium large cats (AcinonyxePuma lineage), making this anatomical character a morphologic convergence among
large felids. The cranial shortening in P. uncia could be related to the
enlargement of the nasal cavities and sinuses, which could be linked
to life in cold environment (Turner and Antn, 1997).

91

Table 3
Measurements (mm) of the mandible and lower teeth of Acinonyx pardinensis
from Pantalla (Italy).
Mandible

SBAU 337627

Length at the condylar process

Length at the coronoid process
Max height
Height behind the canine
Height behind M1
Breadth at P4
Length of the diastema
Postcanine tooth length (alveolar)
Cmand
Length
Breadth
Crown height
P3
Length
Max breadth
Mesial breadth
Paraconid length
Paraconid height
P4
Length
Max breadth
Mesial breadth
Paraconid length
Paraconid height
M1
Length
Max breadth
Protoconid length
Protoconid height
Paraconid length
Paraconid height
Height at the central notch

154.0
ca 150
74.0
29.5
29.2
15.6
12.2
52.0
13.5
10.3
22.3
14.7
8.1
6.4
8.4
10.4
18.4
8.5
6.9
9.1
11.8
21.3
10.4
10.5
12.7
11.9
14
7.5

The Pantalla felid also shows a lateral enlargement of the frontals caudally to the zygomatic processes (Fig. 7c3). This character
was pointed out as a synapomorphy of the closely related genera
Acinonyx and Puma (Spassov, 2011), but is observable to a lesser
degree also in P. uncia.
The Pantalla specimens share with A. pardinensis from Saint
Vallier the widening of the nares and orbits, with the latter oriented
frontward (Fig. 7b4 and c4). This morphology is intermediate between the modern Panthera (showing narrower nares and smaller
and more laterally-oriented orbits; Fig. 7d4) and Acinonyx (having
very large nares and orbits; Fig. 7a4). Similarly to Acinonyx, the cat
from Pantalla has smaller infraorbital foramina than Panthera.
The quite high development of the sagittal and nuchal crests can
be pointed out observing the crania from Pantalla in lateral and
dorsal view. In this feature, they resemble more Panthera than
Acinonyx (Fig. 7a1, c1 and d1). The morphology of the caudal part of
the cranium in A. pardinensis from Saint Vallier (Fig. 7a2) should be
considered with prudence, because this portion was reconstructed
(Geraads, 2008).
Taking into account all the above cranial characters, the felid
from Pantalla strongly recalls the morphology of A. pardinensis
linxiaensis from Longdan (Qiu et al., 2004, Plates 20 and 21),
although the Chinese specimens are slightly larger than the Italian
ones.
The upper dentition of the Pantalla felid is much more similar to
cheetah-like than to pantherine-like cats. The quite slender canines
and the narrow cheek teeth are two of the most important characters identifying cheetahs according to ORegan (2002). All the
characters of the P3 (the relative development of the cusps, the high
and vertical crown, the narrow occlusal outline) clearly resemble
the extant A. jubatus as well as A. pardinensis from Saint Vallier. In
particular, the P3 of the Pantalla specimens is more similar to the

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M. Cherin et al. / Quaternary Science Reviews 87 (2014) 82e97

one of the skull fragment QSV 116 (Viret, 1954, Plate 12, Fig. 3b)
than to that of the cranium QSV 112 (Viret, 1954, Plate 12, Fig. 1b),
which appears broader in occlusal view. In this, the latter resembles
the P3 of A. p. linxiaensis (Qiu et al., 2004, Plate 20, Fig. 1c).
The P4 is the most characteristic tooth, showing a marked
reduction of the protocone. Such morphology has been recognized
as typical for both extant and extinct cheetahs by many authors
(Viret, 1954; Van Valkenburgh et al., 1990; Geraads, 1997; ORegan,
2002; Argant, 2004; Spassov, 2011). From a morphometric point of
view, the reduction of the protocone corresponds to a reduction of
the ratio between the P4 maximum breadth and length, which is on
average always lower than 45% in cheetah-like cats (i.e., Acinonyx
and Miracinonyx species; Fig. 8). On the contrary, the upper
carnassial of Panthera and Puma always shows a strong and
lingually-enlarged protocone, more or less oriented in mesial direction (Fig. 8). From this perspective, the P4 from Pantalla differs
from the recently described Acinonyx kurteni from the Late Pliocene
of China (Christiansen and Mazk, 2009), which is characterized by
a Panthera-like strong protocone. As a matter of fact, the holotype

Fig. 8. Occlusal view of the left upper carnassial of selected extinct and extant Felidae,
drawn not in scale. Acinonyx pardinensis and A. jubatus are characterized by a strongly
reduced and distally-placed protocone. For each species, the ratio between the
maximum breadth and length (average 100) is indicated with bold numbers (sample
size in brackets). This ratio is <45% in Acinonyx and >45% in Panthera and Puma. The
specimens used for the making of the drawings are indicated below each species name.

and single specimen of A. kurteni seems to be a fossil forgery, since

most of the skull was articially modeled (Deng, 2011). As a
consequence, the validity of this taxon was seriously challenged
(Spassov, 2011). Christiansen and Mazk (2009) also pointed out
that one of the most peculiar features of the cheetahs upper
carnassial is the presence of a well-developed ectoparastyle, an
accessory style located mesiolabially to the parastyle. Although the
extant A. jubatus is characterized by a prominent ectoparastyle, we
believe that it cannot be considered as a diagnostic character. In fact
this is present with different extent in many other large felids such
as P. onca and P. tigris, moreover showing a remarkable individual
variability. The dimensional values of the P4 from Pantalla fall into
the range of A. pardinensis from Eurasia and North Africa. The upper
carnassial IGF 4372 from Olivola cited by Kurtn and Crusafont
Pair (1977) was not taken into consideration in our morphometric comparisons since it does not belong to A. pardinensis but to
P. gombaszoegensis. Similarly, the tooth IGF 15358 from Montopoli
labeled as Acinonyx sp. in the catalogue of the Florence paleontological museum was recently referred to Puma pardoides (Cherin
et al., 2013c).
SBAU 337627 resembles a typical cheetahs mandible in all its
morphological characters. On the contrary, pantherine cats normally have longer and more robust mandibles, with a longer diastema between the canine and the P3, and a deeper and wider
masseteric fossa. The overall morphology of the hemimandible
from Pantalla is very similar to the complete specimens of
A. pardinensis from Saint Vallier, Untermassfeld and Longdan.
However, it has to be pointed out that the corpus of the French
specimen QSV 112 (Viret, 1954, Plate 12, Fig. 2a) in occlusal view is
more curved than the one form Pantalla, which is almost
completely straight. In addition, the two mandibles from Untermassfeld IQW 1980/15503 and 15504 are characterized by a very
caudally-inclined coronoid process, which is quite vertical in SBAU
337627. Nevertheless, this character seems to be quite variable in
cheetah-like cats, given that the two complete mandibles of A. p.
linxiaensis gured by Qiu et al. (2004, Plate 20, Fig. 2 and Plate 21,
Fig. 2) are clearly different from this perspective.
The angular outline of the rostroventral margin is another
interesting mandibular character linking the Pantalla mandible to
fossil and living cheetahs. On the contrary, in pantherine cats and
pumas this margin is more receding, probably due to the greater
development of the canine root. This reects also in the symphysis
shape, which is very short in cheetahs and long in Panthera and
Puma.
The lower postcanine teeth of SBAU 337627 are again very
similar to those of cheetah-like cats. The partial overlapping of the
three teeth in occlusal view and the quite short P3 compared to the
M1 are two characters pointed out for A. pardinensis by Schaub
(1949) and Viret (1954). In labial view, the high-crowned postcanine teeth have more or less the same height, as in all cheetahs
and differently form Panthera, in which the P3 is normally much
lower than the P4 and M1. In addition, the pantherine lower premolars never show the characteristic cheetah-like eur-de-lis
morphology. The single right P4 of A. pardinensis reported at Casa
Sgherri (Italy) by Marcolini et al. (2000) cannot be referred to this
taxon because it does not show this peculiar morphology, but a
heavier and sturdier pantherine-like appearance. Unfortunately,
Marcolini et al. (2000) did not publish any measure of the tooth, so
it is not possible to offer an alternative taxonomic attribution.
In the light of the above comparisons, the felid from Pantalla is
here referred to the large-sized cheetah-like cat A. pardinensis.
Further taxonomic considerations are hard to be carried on because
the available fossil record is not enough to shine a light on this
carnivores taxonomic status. The macrospecies concept proposed
by Hemmer et al. (2008, 2011) and supported, at least in part, by

M. Cherin et al. / Quaternary Science Reviews 87 (2014) 82e97

Spassov (2011) represents a good, parsimonious model that however should be reviewed starting from the Pantalla sample. In fact,
the recognizing of some primitive, pantherine-like characters in the
Pantalla skull material (discussed from a functional and ecological
perspective in the next paragraph) makes it more similar to the
middleelate Villafranchian North African and Chinese subspecies
(respectively A. p. aicha and A. p. linxiaensis) rather than to the one
occurring in Europe and Central Asia in the same period (A. p.
pardinensis). As a consequence, the relations among the above three
forms should be reconsidered. Given the quite continuous chronologic and geographic distribution (Fig. 1), the middle to late
Villafranchian Old World cheetah-like cats may belong to a single
wide-ranging taxon. In this framework, the complete skull QSV 112
from Saint Vallier, which was frequently used in the past as a
reference specimen for describing the anatomy and inferring the
ecology of A. pardinensis (cfr. Turner and Antn, 1997), should be restudied taking into account the hard work of restoration of its
caudal portion (Geraads, 2008). Similarly, as previously suggested
by Spassov (2011), the relationships between the early Villafranchian A. p. arvernensis and its chronologic successor A. p. pardinensis need to be re-examined in the light of the recent ndings of
the former taxon in Tuozidong at about 2 Ma (Dong et al., 2013).
5. Predatory behavior and ecological role of the giant Plioe
Pleistocene cheetah
In the last decades a number of authors tried to infer information about the biology and ecology of the giant cheetah
A. pardinensis starting from its skeletal morphology (Kurtn, 1968;
Turner and Antn, 1997; Hemmer et al., 2011). Most of these studies
were focused on comparisons with the modern cheetah A. jubatus,
one of the most charismatic large carnivorans in the World, whose
predatory behavior was subject of several studies.
On the other hand, the direct similarity between the hunting
strategy in A. pardinensis and A. jubatus is questionable in the light
of at least two considerations:
(1) Although the fossil record is rather scanty, paleontological data
at our disposal do not clarify the phylogenetic relationship
between the two species. Acinonyx jubatus probably appears in
Africa in the MiddleeLate Pliocene (cfr. Werdelin and Peign,
2010), while the earliest records of cheetah-like cats in Eurasia and North Africa are not older than the PlioePleistocene

93

boundary, even if some undescribed materials of Sivapanthera sp. from the Middle Pliocene of Yushe (Qiu, 2006) could
represent the rst occurrence of this group. In addition,
phylogenetic proximity is not necessarily related to similar
predatory behaviors.
(2) Thanks to the outstanding running capabilities the extant
cheetah is considered a highly specialized predator, with a
strong hunting adaptation in open savannah environments.
Nevertheless, such considerations are inuenced by the fact
that most of the studies on the cheetahs ecology were
focused on the Serengeti area in East Africa. On the contrary,
recent researches in different areas (Broomhall et al., 2004;
Mills et al., 2004; Bissett and Bernard, 2007) demonstrate
that A. jubatus is a quite adaptable species, with a great
behavioral and ecological exibility. This is indirectly supported considering that the cheetah former range was much
wider than today, including large part of Africa and southwestern Asia, from Arabia to India (Kingdon, 1977).
Although more ecologically exible than previously thought,
A. jubatus is an atypical large-sized cat, which evolved a very
peculiar predatory behavior (see Section 1). Differently from pantherine cats, the cheetahs small and rounded skull, which is
modied for speed as well as the postcranial skeleton, appears as an
apparent anatomical deciency for hunting large prey (Eaton,
1970). In fact, pantherine cats have very strong and large canines
adapted to being wedged between the preys cervical vertebrae,
causing a rapid death. In addition, their large postcanine teeth
(especially the upper carnassial, bearing a strong protocone) can
crush parts of the preys neck and skull (Eaton, 1970). Some jaguars
from Mato Grosso (Brasil) were adapted in killing the capybara by
punching their temporal bones with the canines (Schaller and
Vasconcelos, 1978). The cheetah is not equipped with similar
tooth lethal weapons, so normally kills the prey by strangulation,
through a prolonged (even some minutes) bite in the ventral part of
the neck (Eaton, 1970).
The above observations represent the starting point of our analyses on the morphology and function of the jaw muscles in
A. pardinensis (Fig. 9), through the comparison with this muscle
system in the extant A. jubatus and P. onca (Fig. 10).
The temporal muscle of A. pardinensis resembles the one of
P. onca in morphology and dimensions, while the overall skull
shape is intermediate between the jaguar and the cheetah (Fig. 10).

Fig. 9. 3D reconstruction sequence of the masticatory muscles of Acinonyx pardinensis from Pantalla (Italy). a) Reconstruction of the skull. b) Highlighting of the insertions areas of
the temporal muscle on the cranium (iatm) and of the masseter muscle on the zygomatic process (iamm) and the masseteric fossa (mf). c) Final reconstruction with the temporal
muscle (tm), masseter muscle (mm) and orbicularis oculi muscle (oom). (Online version in color.)

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M. Cherin et al. / Quaternary Science Reviews 87 (2014) 82e97

Fig. 10. 3D reconstruction and morphological comparison of the masticatory muscles in Acinonyx jubatus MCZR 367 (a), Acinonyx pardinensis SBAU 337624 from Pantalla (b) and
Panthera onca MCZR 258 (c). (Online version in color.)

The short and domed morphology of the cheetahs cranium is

related to the small size and crowding of the teeth, as well as to the
greater arching of the zygomatic bones, which are more distant
from the tooth occlusal plane (Fig. 10a). Such morphology causes a
greater extension of the masseter muscle during the jaw opening
compared to pantherine cats. The strength of any given muscle is
proportional to its cross sectional area (Liem et al., 2001), so the
higher the muscle extension, the lower its strength. In addition, the
muscle strength is also related to the mechanics of the mandibular
lever. In felids with a short muzzle the distance between the canines and the lever fulcrum (i.e., the temporomandibular joint) is
lower. This reects in the decrease of the moment arm of resistance
and the enhancing of bite force (Kitchener et al., 2010). In the light
of this biomechanical principle Hemmer (2001) suggested that the
skull shortening in A. jubatus and A. pardinensis allows these species
to have a more powerful bite than pantherine cats. On the other
hand, in large felids with elongated crania the decrease of the bite
force due to the greater distance between the canines and the
mandibular joint is balanced by a greater development of jaw
muscles (Slater and Van Valkenburgh, 2009). This suggests that the
bite force is nearly constant in felids as the body size increases, so
the splanchnocranium shortening in the cheetah-like cats does not
entail a more powerful bite.
With the above considerations in mind, it can be understood
why the musculoskeletal skull structure in pantherine cats (in
which the canines are very strong and stout, the temporalis has an
upside down-L shape extending in the occipital region, and the
masseter insertion area on the cranium is close to the occlusal plane
due to the low-bowed zygomatic arch; Fig. 10c) is built to achieve a
very strong bite. This allows these carnivores to focus on large-sized
prey, whose catch and kill can cause high mechanical stresses to the
predators skull (Van Valkenburgh and Hertel, 1998; Radloff and du
Toit, 2004; Hayward and Kerley, 2005; Meachen-Samuels and Van
Valkenburgh, 2009).
The skull morphology of A. pardinensis suggests us that this
extinct felids masticatory system could be more similar to that of
the extant jaguar than that of the cheetah (Figs. 10 and 11). Similarly, the mean body size of A. pardinensis (about 80 kg) was much
closer to that of a large pantherine cat (about 80 kg for P. onca) than
that of A. jubatus (about 40 kg) (details in Table 4). As a consequence, in the light of the musculoskeletal skull morphology and
body size, it is likely that A. pardinensis could catch large prey
through a killing strategy more similar to pantherine cats than to
the small and slender cheetah. The interspecic competition with
P. gombaszoegensis in the areas where both species occurred (e.g.,
Dmanisi, Untermassfeld, Pirro Nord) could be reduced by temporal
(e.g., diurnal against nocturnal hunting activity) and/or spatial (e.g.,

preference for more or less thicket areas in heterogeneous ecosystems) niche displacements.
The prey of A. jubatus ranges from the hare (about 2 kg) to
adult wildebeest and zebra (up to 270 kg), although this felid
usually catches medium to small-sized ungulates like gazelle,
impala and springbok with a mass of 23e56 kg (Turner and
Antn, 1997; Hayward et al., 2006). The hunting target is inuenced by the local availability but also by the vegetation structure: cheetahs hunting in thicket areas focus on relatively larger
prey (Hayward et al., 2006). This is related to the lower presence
of kleptoparasites like lions, hyenas, jackals and vultures in such
habitats than in grasslands, where these ecological interactions
(which frequently conclude with a prey loss by the cheetah) are
rather common (Eaton, 1970; Hayward et al., 2006; Hunter et al.,
2006). A similar preference for relatively larger prey can be
observed in the Asiatic cheetah A. j. venaticus, which occupies
areas with very low probability of meat robbery (Farhadinia and
Hemami, 2010). Most of the PlioePleistocene paleoecosystems
in which A. pardinensis occurred were characterized by a very rich
and complex carnivore guild. Hemmer et al. (2011) hypothesized
that in Dmanisi the hunting activity of the giant cheetah (probably focused on 50e100 kg prey) could provide a large amount of
fresh meat available for kleptoparasites, including early Homo.
Nevertheless, this hypothesis is again based on a direct correlation between the hunting strategy of A. pardinensis and that of
A. jubatus, which normally leaves uneaten large part of the preys
carcass (Hayward et al., 2006). In the light of this presumed
ecological and behavioral correlation, A. pardinensis was considered a low-ranked species in the PlioePleistocene carnivore hierarchy compared to other taxa such as saber-toothed cats or
P. gombaszoegensis (Hemmer, 2001; Hemmer et al., 2011). Our
results suggest that the ecological role of the giant cheetah should
be dealt with caution, since this predators ecology and behavior
were probably more complex than previously thought and rather
different from those of the extant cheetah. The supposed adaptation of A. pardinensis for high speed in open environments was
already brought into consideration by Turner and Antn (1997).
The few postcranial elements at our disposal, such as those from
touaires (Schaub, 1949), Montopoli (Ficcarelli, 1984), Untermassfeld (Hemmer, 2001) and Dmanisi (Hemmer et al., 2011),
suggest that A. pardinensis had body proportions similar to the
living cheetah. On the other hand, we cannot demonstrate that
this body structure was really related to a hunting strategy based
on high speed. While the skull anatomy is more strictly related to
the feeding behavior, the postcranial one could be inuenced by
other ecological and environmental factors. This is demonstrated
by the fact that a number of living medium- to large-sized felids

M. Cherin et al. / Quaternary Science Reviews 87 (2014) 82e97

95

Table 4
Predicted body masses (kg) for Acinonyx pardinensis from various Old World sites
based on the length of the lower carnassial (mm), calculated using the prediction
equation by Van Valkenburgh (1990). The source of morphometric data is indicated
in brackets. The body mass of A. pardinensis ranges from 60 to 121 kg, with an
average value of 83 kg, which is very close to the one estimated for the giant cheetah
from Pantalla (80 kg). The prediction equation by Van Valkenburgh (1990) was also
tested by calculating the body mass of individuals of some living felid species (SPE
and HNHM collections) and then comparing these analytic results with the average
weights published by Marker and Dickman (2003; for A. jubatus) and Macdonald
et al. (2010; for the other four species). The estimated mean body masses (EBM;
sample size in brackets) are very close to the actual mean body masses (ABM).
Locality and references

Specimen

M1
length

Body
mass

Pantalla, Italy (original data)

Montopoli, Italy (original data)
Villafranca dAsti, Italy
(Kurtn and Crusafont
Pair, 1977)
Mosbach, Germany
(Hemmer et al., 2008)
La Cte dArd, France
(Kurtn and Crusafont
Pair, 1977)
Les touaires, France
(Kurtn and Crusafont
Pair, 1977)
Saint Vallier, France
(Viret, 1954; Kurtn and
Crusafont Pair, 1977;
Argant, 2004)

SBAU 337627
IGF 12477
MHNB V.I.132

21.3
19.6
20.5

80
62
71

SMF PA/F.6236

19.2

60

MHNP e

20.3

69

MHNP e

24.4

121

MHNL QSV.110

19.5

61

MHNL QSV.112
MHNL QSV.113
MHNL QSV.117
MHNB St.V.122
MHNL SV.98.624
IQW 1980/15503

20.3
21.4
21.0
19.7
20.1
23.9

69
81
76
63
67
113

IQW 1980/15504
IGME FP1-2002-1027

23.4
19.7

106
63

ICP V.133

21.9

87

INSAP AaO-1325

22.7

97

INSAP AaO-3187
HMV 1221

20.8
22.5

74
94

IVPP V.13537
HMV 1223
e

23.2
19.4
23.8

103
60
112

23.6

109

EBM
44 (5)
76 (8)
44 (16)
149 (11)
41 (3)

83
ABM
43
86
42
147
45

Untermassfeld, Germany
(Hemmer, 2001)
Fonelas P-1, Spain
(Garrido, 2008)
Villarroya, Spain
(Kurtn and Crusafont
Pair, 1977)
Ahl Al Oughlam,
Morocco (Geraads, 1997)
Longdan, China
(Qiu et al., 2004)

Yuanqu Loc. 105,

China (Zdansky, 1925)
Siwalik Hills, Pinjor
NHM 16573
Stage (original data)
Mean body mass (Pantalla excluded)
Living species
Acinonyx jubatus
Panthera onca
Panthera pardus
Panthera leo
Puma concolor

Fig. 11. Reconstruction sequence of the head shape of Acinonyx pardinensis from
Pantalla. Artwork by D.A. Iurino.

are characterized by similar proportions but very different

hunting behaviors compared to the cheetah. For instance, the
extant snow leopard resembles to some extent the cheetah in
having a slender skeleton with long limbs, but these features are
related to this cats outstanding performance in mountainous
terrain (Turner and Antn, 1997). In the next future, a systematic
study of all the reported postcranial material of A. pardinensis,
including a detailed comparative analysis of muscle insertions,
may allow us to better explore such aspects of this carnivores
function and ecology.

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M. Cherin et al. / Quaternary Science Reviews 87 (2014) 82e97

Acknowledgments
We gratefully thank all the people that helped and supported us
in analyzing museum collections: M.C. De Angelis (SBAU), E. Cioppi
(IGF), G. Csorba and M. Gasparik (HNHM), A. Lister and P. Brewer
(NHM) and R. Carlini (MCZR). M. Danti and W.S. Della Sala (M.G.
Vannini Hospital, Rome) were essential for the making of the CT
scanning. C. Eleni and C. Cocumelli (Istituto Zooprolattico Sperimentale del Lazio e della Toscana, Italy) gave us useful advices in
the rst phase of this work. Thanks are also due to L. Santini, N.
Spassov and T. Deng for their precious suggestions and help. We are
indebted to M.J. Salesa and an anonymous reviewer for the very
useful comments on the manuscript.
Part of the comparative analyses for this study were carried out
thanks to the support from the SYNTHESYS Project http://www.
synthesys.info/, which is nanced by European Community
Research Infrastructure Action under the FP7 Capacities Program
(project number: HU-TAF-2306 and GB-TAF-2953 for MC).
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